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<span>Toggle Developments leading up to the synthesis subsection</span> </button> <ul id="toc-Developments_leading_up_to_the_synthesis-sublist" class="vector-toc-list"> <li id="toc-Darwin's_evolution_by_natural_selection,_1859" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Darwin's_evolution_by_natural_selection,_1859"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Darwin's evolution by natural selection, 1859</span> </div> </a> <ul id="toc-Darwin's_evolution_by_natural_selection,_1859-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-The_eclipse_of_Darwinism,_1880s_onwards" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#The_eclipse_of_Darwinism,_1880s_onwards"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>The eclipse of Darwinism, 1880s onwards</span> </div> </a> <ul id="toc-The_eclipse_of_Darwinism,_1880s_onwards-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Weismann's_germ_plasm,_1892" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Weismann's_germ_plasm,_1892"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Weismann's germ plasm, 1892</span> </div> </a> <ul id="toc-Weismann's_germ_plasm,_1892-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Disputed_beginnings" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Disputed_beginnings"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Disputed beginnings</span> </div> </a> <button aria-controls="toc-Disputed_beginnings-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 Disputed beginnings subsection</span> </button> <ul id="toc-Disputed_beginnings-sublist" class="vector-toc-list"> <li id="toc-Genetics,_mutationism_and_biometrics,_1900–1918" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetics,_mutationism_and_biometrics,_1900–1918"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Genetics, mutationism and biometrics, 1900–1918</span> </div> </a> <ul id="toc-Genetics,_mutationism_and_biometrics,_1900–1918-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Castle's_hooded_rats,_1911" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Castle's_hooded_rats,_1911"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Castle's hooded rats, 1911</span> </div> </a> <ul id="toc-Castle's_hooded_rats,_1911-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Morgan's_fruit_flies,_1912" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Morgan's_fruit_flies,_1912"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Morgan's fruit flies, 1912</span> </div> </a> <ul id="toc-Morgan's_fruit_flies,_1912-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-An_obstruction:_Woodger's_positivism,_1929" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#An_obstruction:_Woodger's_positivism,_1929"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>An obstruction: Woodger's positivism, 1929</span> </div> </a> <ul id="toc-An_obstruction:_Woodger's_positivism,_1929-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Elements_of_the_synthesis" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Elements_of_the_synthesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Elements of the synthesis</span> </div> </a> <button aria-controls="toc-Elements_of_the_synthesis-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 Elements of the synthesis subsection</span> </button> <ul id="toc-Elements_of_the_synthesis-sublist" class="vector-toc-list"> <li id="toc-Fisher_and_Haldane's_mathematical_population_genetics,_1918–1930" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Fisher_and_Haldane's_mathematical_population_genetics,_1918–1930"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Fisher and Haldane's mathematical population genetics, 1918–1930</span> </div> </a> <ul id="toc-Fisher_and_Haldane's_mathematical_population_genetics,_1918–1930-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-De_Beer's_embryology,_1930" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#De_Beer's_embryology,_1930"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>De Beer's embryology, 1930</span> </div> </a> <ul id="toc-De_Beer's_embryology,_1930-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Wright's_adaptive_landscape,_1932" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Wright's_adaptive_landscape,_1932"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Wright's adaptive landscape, 1932</span> </div> </a> <ul id="toc-Wright's_adaptive_landscape,_1932-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dobzhansky's_evolutionary_genetics,_1937" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dobzhansky's_evolutionary_genetics,_1937"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Dobzhansky's evolutionary genetics, 1937</span> </div> </a> <ul id="toc-Dobzhansky's_evolutionary_genetics,_1937-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ford's_ecological_genetics,_1940" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ford's_ecological_genetics,_1940"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5</span> <span>Ford's ecological genetics, 1940</span> </div> </a> <ul id="toc-Ford's_ecological_genetics,_1940-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Schmalhausen's_stabilizing_selection,_1941" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Schmalhausen's_stabilizing_selection,_1941"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.6</span> <span>Schmalhausen's stabilizing selection, 1941</span> </div> </a> <ul id="toc-Schmalhausen's_stabilizing_selection,_1941-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Huxley's_popularising_synthesis,_1942" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Huxley's_popularising_synthesis,_1942"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.7</span> <span>Huxley's popularising synthesis, 1942</span> </div> </a> <ul id="toc-Huxley's_popularising_synthesis,_1942-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mayr's_allopatric_speciation,_1942" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mayr's_allopatric_speciation,_1942"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.8</span> <span>Mayr's allopatric speciation, 1942</span> </div> </a> <ul id="toc-Mayr's_allopatric_speciation,_1942-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Simpson's_palaeontology,_1944" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Simpson's_palaeontology,_1944"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.9</span> <span>Simpson's palaeontology, 1944</span> </div> </a> <ul id="toc-Simpson's_palaeontology,_1944-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Society_for_the_Study_of_Evolution,_1946" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Society_for_the_Study_of_Evolution,_1946"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.10</span> <span>Society for the Study of Evolution, 1946</span> </div> </a> <ul id="toc-Society_for_the_Study_of_Evolution,_1946-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stebbins's_botany,_1950" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Stebbins's_botany,_1950"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.11</span> <span>Stebbins's botany, 1950</span> </div> </a> <ul id="toc-Stebbins's_botany,_1950-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Definitions_by_the_founders" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Definitions_by_the_founders"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Definitions by the founders</span> </div> </a> <ul id="toc-Definitions_by_the_founders-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-After_the_synthesis" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#After_the_synthesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>After the synthesis</span> </div> </a> <button aria-controls="toc-After_the_synthesis-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 After the synthesis subsection</span> </button> <ul id="toc-After_the_synthesis-sublist" class="vector-toc-list"> <li id="toc-Hamilton's_inclusive_fitness,_1964" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Hamilton's_inclusive_fitness,_1964"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Hamilton's inclusive fitness, 1964</span> </div> </a> <ul id="toc-Hamilton's_inclusive_fitness,_1964-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Williams's_gene-centred_evolution,_1966" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Williams's_gene-centred_evolution,_1966"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Williams's gene-centred evolution, 1966</span> </div> </a> <ul id="toc-Williams's_gene-centred_evolution,_1966-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Wilson's_sociobiology,_1975" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Wilson's_sociobiology,_1975"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>Wilson's sociobiology, 1975</span> </div> </a> <ul id="toc-Wilson's_sociobiology,_1975-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Lewis's_homeotic_genes,_1978" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Lewis's_homeotic_genes,_1978"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.4</span> <span>Lewis's homeotic genes, 1978</span> </div> </a> <ul id="toc-Lewis's_homeotic_genes,_1978-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Later_syntheses" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Later_syntheses"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Later syntheses</span> </div> </a> <button aria-controls="toc-Later_syntheses-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 Later syntheses subsection</span> </button> <ul id="toc-Later_syntheses-sublist" class="vector-toc-list"> <li id="toc-Pigliucci's_extended_evolutionary_synthesis,_2007" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Pigliucci's_extended_evolutionary_synthesis,_2007"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Pigliucci's extended evolutionary synthesis, 2007</span> </div> </a> <ul id="toc-Pigliucci's_extended_evolutionary_synthesis,_2007-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Koonin's_'post-modern'_evolutionary_synthesis,_2009" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Koonin's_'post-modern'_evolutionary_synthesis,_2009"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Koonin's 'post-modern' evolutionary synthesis, 2009</span> </div> </a> <ul id="toc-Koonin's_'post-modern'_evolutionary_synthesis,_2009-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Towards_a_replacement_synthesis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Towards_a_replacement_synthesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Towards a replacement synthesis</span> </div> </a> <ul id="toc-Towards_a_replacement_synthesis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Historiography" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Historiography"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Historiography</span> </div> </a> <ul id="toc-Historiography-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Notes" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Notes"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Notes</span> </div> </a> <ul id="toc-Notes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sources" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Sources"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>Sources</span> </div> </a> <ul id="toc-Sources-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-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">Modern synthesis (20th century)</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 37 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-37" 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">37 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/%D8%A7%D8%B5%D8%B7%D9%86%D8%A7%D8%B9_%D8%AA%D8%B7%D9%88%D8%B1%D9%8A_%D8%AD%D8%AF%D9%8A%D8%AB" title="اصطناع تطوري حديث – Arabic" lang="ar" hreflang="ar" data-title="اصطناع تطوري حديث" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%86%E0%A6%A7%E0%A7%81%E0%A6%A8%E0%A6%BF%E0%A6%95_%E0%A6%AC%E0%A6%BF%E0%A6%AC%E0%A6%B0%E0%A7%8D%E0%A6%A4%E0%A6%A8%E0%A6%BF%E0%A6%95_%E0%A6%B8%E0%A6%82%E0%A6%B6%E0%A7%8D%E0%A6%B2%E0%A7%87%E0%A6%B7%E0%A6%A3" title="আধুনিক বিবর্তনিক সংশ্লেষণ – Bangla" lang="bn" hreflang="bn" data-title="আধুনিক বিবর্তনিক সংশ্লেষণ" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A1%D0%B8%D0%BD%D1%82%D0%B5%D1%82%D0%B8%D1%87%D0%BD%D0%B0_%D1%82%D0%B5%D0%BE%D1%80%D0%B8%D1%8F_%D0%B7%D0%B0_%D0%B5%D0%B2%D0%BE%D0%BB%D1%8E%D1%86%D0%B8%D1%8F%D1%82%D0%B0" title="Синтетична теория за еволюцията – Bulgarian" lang="bg" hreflang="bg" data-title="Синтетична теория за еволюцията" 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/Moderna_evolucijska_sinteza" title="Moderna evolucijska sinteza – Bosnian" lang="bs" hreflang="bs" data-title="Moderna evolucijska sinteza" 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/S%C3%ADntesi_evolutiva_moderna" title="Síntesi evolutiva moderna – Catalan" lang="ca" hreflang="ca" data-title="Síntesi evolutiva moderna" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Modern%C3%AD_evolu%C4%8Dn%C3%AD_synt%C3%A9za" title="Moderní evoluční syntéza – Czech" lang="cs" hreflang="cs" data-title="Moderní evoluční syntéza" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Synthetische_Evolutionstheorie" title="Synthetische Evolutionstheorie – German" lang="de" hreflang="de" data-title="Synthetische Evolutionstheorie" 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/S%C3%BCnteetiline_evolutsiooniteooria" title="Sünteetiline evolutsiooniteooria – Estonian" lang="et" hreflang="et" data-title="Sünteetiline evolutsiooniteooria" 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/S%C3%ADntesis_evolutiva_moderna" title="Síntesis evolutiva moderna – Spanish" lang="es" hreflang="es" data-title="Síntesis evolutiva moderna" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Eboluzioaren_teoria_sintetikoa" title="Eboluzioaren teoria sintetikoa – Basque" lang="eu" hreflang="eu" data-title="Eboluzioaren teoria sintetikoa" 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/%D8%B3%D9%86%D8%AA%D8%B2_%D9%86%D9%88%DB%8C%D9%86" title="سنتز نوین – Persian" lang="fa" hreflang="fa" data-title="سنتز نوین" 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/Th%C3%A9orie_synth%C3%A9tique_de_l%27%C3%A9volution" title="Théorie synthétique de l'évolution – French" lang="fr" hreflang="fr" data-title="Théorie synthétique de l'évolution" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/S%C3%ADntese_moderna_(s%C3%A9culo_XX)" title="Síntese moderna (século XX) – Galician" lang="gl" hreflang="gl" data-title="Síntese moderna (século XX)" 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%98%84%EB%8C%80_%EC%A7%84%ED%99%94_%EC%9D%B4%EB%A1%A0" 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title="This is a good article. Click here for more information."><img alt="This is a good article. Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Fusion of natural selection with Mendelian inheritance</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 other uses, see <a href="/wiki/Modern_synthesis_(disambiguation)" class="mw-disambig" title="Modern synthesis (disambiguation)">Modern synthesis (disambiguation)</a>.</div> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Modern_Synthesis.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/97/Modern_Synthesis.svg/390px-Modern_Synthesis.svg.png" decoding="async" width="390" height="293" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/97/Modern_Synthesis.svg/585px-Modern_Synthesis.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/97/Modern_Synthesis.svg/780px-Modern_Synthesis.svg.png 2x" data-file-width="900" data-file-height="675" /></a><figcaption>Several major ideas about <a href="/wiki/Evolution" title="Evolution">evolution</a> came together in the <a href="/wiki/Population_genetics" title="Population genetics">population genetics</a> of the early 20th century to form the modern synthesis, including <a href="/wiki/Genetic_variation" title="Genetic variation">genetic variation</a>, <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>, and particulate (<a href="/wiki/Mendelian_genetics" class="mw-redirect" title="Mendelian genetics">Mendelian</a>) inheritance.<sup id="cite_ref-Gould216_1-0" class="reference"><a href="#cite_note-Gould216-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This ended the <a href="/wiki/Eclipse_of_Darwinism" class="mw-redirect" title="Eclipse of Darwinism">eclipse of Darwinism</a> and supplanted a variety of <a href="/wiki/Alternatives_to_Darwinism" class="mw-redirect" title="Alternatives to Darwinism">non-Darwinian theories of evolution</a>.</figcaption></figure> <p>The <b>modern synthesis</b><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup> was the early 20th-century synthesis of <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a>'s theory of <a href="/wiki/Evolution" title="Evolution">evolution</a> and <a href="/wiki/Gregor_Mendel" title="Gregor Mendel">Gregor Mendel</a>'s ideas on heredity into a joint mathematical framework. <a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a> coined the term in his 1942 book, <i><a href="/wiki/Evolution:_The_Modern_Synthesis" title="Evolution: The Modern Synthesis">Evolution: The Modern Synthesis</a></i>. The synthesis combined the ideas of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>, <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian genetics</a>, and <a href="/wiki/Population_genetics" title="Population genetics">population genetics</a>. It also related the broad-scale <a href="/wiki/Macroevolution" title="Macroevolution">macroevolution</a> seen by <a href="/wiki/Paleontology" title="Paleontology">palaeontologists</a> to the small-scale <a href="/wiki/Microevolution" title="Microevolution">microevolution</a> of local <a href="/wiki/Population" title="Population">populations.</a> </p><p>The synthesis was defined differently by its founders, with <a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Ernst Mayr</a> in 1959, <a href="/wiki/G._Ledyard_Stebbins" title="G. Ledyard Stebbins">G. Ledyard Stebbins</a> in 1966, and <a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a> in 1974 offering differing basic postulates, though they all include natural selection, working on heritable variation supplied by mutation. Other major figures in the synthesis included <a href="/wiki/E._B._Ford" title="E. B. Ford">E. B. Ford</a>, <a href="/wiki/Bernhard_Rensch" title="Bernhard Rensch">Bernhard Rensch</a>, <a href="/wiki/Ivan_Schmalhausen" title="Ivan Schmalhausen">Ivan Schmalhausen</a>, and <a href="/wiki/George_Gaylord_Simpson" title="George Gaylord Simpson">George Gaylord Simpson</a>. An early event in the modern synthesis was <a href="/wiki/R._A._Fisher" class="mw-redirect" title="R. A. Fisher">R. A. Fisher</a>'s 1918 paper on mathematical population genetics, though <a href="/wiki/William_Bateson" title="William Bateson">William Bateson</a>, and separately <a href="/wiki/Udny_Yule" title="Udny Yule">Udny Yule</a>, had already started to show how Mendelian genetics could work in evolution in 1902. </p><p>Different syntheses followed, including with <a href="/wiki/Social_behaviour" class="mw-redirect" title="Social behaviour">social behaviour</a> in <a href="/wiki/E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a>'s <a href="/wiki/Sociobiology" title="Sociobiology">sociobiology</a> in 1975, <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a>'s integration of <a href="/wiki/Embryology" title="Embryology">embryology</a> with genetics and evolution, starting in 1977, and <a href="/wiki/Massimo_Pigliucci" title="Massimo Pigliucci">Massimo Pigliucci</a>'s and <a href="/wiki/Gerd_B._M%C3%BCller" title="Gerd B. Müller">Gerd B. Müller</a>'s proposed <a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">extended evolutionary synthesis</a> of 2007. In the view of evolutionary biologist <a href="/wiki/Eugene_Koonin" title="Eugene Koonin">Eugene Koonin</a> in 2009, the modern synthesis will be replaced by a 'post-modern' synthesis that will include revolutionary changes in <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a>, the study of <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a> and the resulting <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">tree of life</a>, and <a href="/wiki/Genomics" title="Genomics">genomics</a>.<sup id="cite_ref-Koonin2009_4-0" class="reference"><a href="#cite_note-Koonin2009-4"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Developments_leading_up_to_the_synthesis">Developments leading up to the synthesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=1" title="Edit section: Developments leading up to the synthesis"><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:Darwin%27s_Pangenesis.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/54/Darwin%27s_Pangenesis.svg/300px-Darwin%27s_Pangenesis.svg.png" decoding="async" width="300" height="103" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/54/Darwin%27s_Pangenesis.svg/450px-Darwin%27s_Pangenesis.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/54/Darwin%27s_Pangenesis.svg/600px-Darwin%27s_Pangenesis.svg.png 2x" data-file-width="925" data-file-height="317" /></a><figcaption><a href="/wiki/Charles_Darwin" title="Charles Darwin">Darwin</a>'s <a href="/wiki/Pangenesis" title="Pangenesis">pangenesis</a> theory. Every part of the body emits tiny gemmules which migrate to the <a href="/wiki/Gonad" title="Gonad">gonads</a> and contribute to the next generation via the fertilised egg. Changes to the body during an organism's life would be inherited, as in <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a>.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History of evolutionary thought</a></div> <div class="mw-heading mw-heading3"><h3 id="Darwin's_evolution_by_natural_selection,_1859"><span id="Darwin.27s_evolution_by_natural_selection.2C_1859"></span>Darwin's evolution by natural selection, 1859</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=2" title="Edit section: Darwin's evolution by natural selection, 1859"><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 articles: <a href="/wiki/Evolution" title="Evolution">Evolution</a> and <a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></div> <p><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a>'s 1859 book, <i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a>,</i> convinced most biologists that <a href="/wiki/Evolution" title="Evolution">evolution</a> had occurred, but not that <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> was its primary mechanism. In the 19th and early 20th centuries, variations of <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a> (inheritance of acquired characteristics), <a href="/wiki/Orthogenesis" title="Orthogenesis">orthogenesis</a> (progressive evolution), <a href="/wiki/Saltationism" class="mw-redirect" title="Saltationism">saltationism</a> (evolution by jumps) and <a href="/wiki/Mutationism" title="Mutationism">mutationism</a> (evolution driven by mutations) were discussed as alternatives.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Darwin himself had sympathy for Lamarckism, but <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> advocated natural selection and totally rejected Lamarckism.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In 1880, Samuel Butler labelled Wallace's view <a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">neo-Darwinism</a>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Blending_Inheritance.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Blending_Inheritance.svg/240px-Blending_Inheritance.svg.png" decoding="async" width="240" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Blending_Inheritance.svg/360px-Blending_Inheritance.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3b/Blending_Inheritance.svg/480px-Blending_Inheritance.svg.png 2x" data-file-width="900" data-file-height="675" /></a><figcaption><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a>, implied by pangenesis, causes the <a href="/wiki/Mean" title="Mean">averaging out</a> of every characteristic, which as the engineer <a href="/wiki/Fleeming_Jenkin" title="Fleeming Jenkin">Fleeming Jenkin</a> pointed out, would make evolution by <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> impossible.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="The_eclipse_of_Darwinism,_1880s_onwards"><span id="The_eclipse_of_Darwinism.2C_1880s_onwards"></span>The eclipse of Darwinism, 1880s onwards</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=3" title="Edit section: The eclipse of Darwinism, 1880s onwards"><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_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></div> <p>From the 1880s onwards, biologists grew skeptical of Darwinian evolution. This <a href="/wiki/Eclipse_of_Darwinism" class="mw-redirect" title="Eclipse of Darwinism">eclipse of Darwinism</a> (in <a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a>'s words) grew out of the weaknesses in Darwin's account, with respect to his view of inheritance. Darwin believed in <a href="/wiki/Blending_inheritance" title="Blending inheritance">blending inheritance</a>, which implied that any new variation, even if beneficial, would be weakened by 50% at each generation, as the engineer <a href="/wiki/Fleeming_Jenkin" title="Fleeming Jenkin">Fleeming Jenkin</a> noted in 1868.<sup id="cite_ref-Bowler196-250_9-0" class="reference"><a href="#cite_note-Bowler196-250-9"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Larson105-129_10-0" class="reference"><a href="#cite_note-Larson105-129-10"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> This in turn meant that small variations would not survive long enough to be selected. Blending would therefore directly oppose natural selection. In addition, Darwin and others considered Lamarckian inheritance of acquired characteristics entirely possible, and Darwin's 1868 theory of <a href="/wiki/Pangenesis" title="Pangenesis">pangenesis</a>, with contributions to the next generation (gemmules) flowing from all parts of the body, actually implied Lamarckism as well as blending.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Darwin68_12-0" class="reference"><a href="#cite_note-Darwin68-12"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Weismann%27s_Germ_Plasm.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Weismann%27s_Germ_Plasm.svg/300px-Weismann%27s_Germ_Plasm.svg.png" decoding="async" width="300" height="155" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/Weismann%27s_Germ_Plasm.svg/450px-Weismann%27s_Germ_Plasm.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/32/Weismann%27s_Germ_Plasm.svg/600px-Weismann%27s_Germ_Plasm.svg.png 2x" data-file-width="1240" data-file-height="642" /></a><figcaption><a href="/wiki/August_Weismann" title="August Weismann">August Weismann</a>'s <a href="/wiki/Germ_plasm" title="Germ plasm">germ plasm</a> theory. The hereditary material, the germplasm, is confined to the <a href="/wiki/Gonad" title="Gonad">gonads</a> and the <a href="/wiki/Gamete" title="Gamete">gametes</a>. Somatic cells (of the body) <a href="/wiki/Embryology" title="Embryology">develop</a> afresh in each generation from the germplasm.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Weismann's_germ_plasm,_1892"><span id="Weismann.27s_germ_plasm.2C_1892"></span>Weismann's germ plasm, 1892</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=4" title="Edit section: Weismann's germ plasm, 1892"><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/Germ_plasm" title="Germ plasm">Germ plasm</a></div> <p><a href="/wiki/August_Weismann" title="August Weismann">August Weismann</a>'s idea, set out in his 1892 book <i>Das Keimplasma: eine Theorie der Vererbung</i> ("The Germ Plasm: a Theory of Inheritance"),<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> was that the hereditary material, which he called the <a href="/wiki/Germ_plasm" title="Germ plasm">germ plasm</a>, and the rest of the body (the <a href="/wiki/Somatic_(biology)" title="Somatic (biology)">soma</a>) had a one-way relationship: the germ-plasm formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a population subject to natural selection. If correct, this made Darwin's pangenesis wrong, and Lamarckian inheritance impossible. His experiment on mice, cutting off their tails and showing that their offspring had normal tails, demonstrated that inheritance was 'hard'.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>b<span class="cite-bracket">]</span></a></sup> He argued strongly and dogmatically<sup id="cite_ref-FOOTNOTEBowler1989248_17-0" class="reference"><a href="#cite_note-FOOTNOTEBowler1989248-17"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> for Darwinism and against Lamarckism, polarising opinions among other scientists. This increased anti-Darwinian feeling, contributing to its eclipse.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEBowler1989247–253,_257_19-0" class="reference"><a href="#cite_note-FOOTNOTEBowler1989247–253,_257-19"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Disputed_beginnings">Disputed beginnings</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=5" title="Edit section: Disputed beginnings"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Genetics,_mutationism_and_biometrics,_1900–1918"><span id="Genetics.2C_mutationism_and_biometrics.2C_1900.E2.80.931918"></span>Genetics, mutationism and biometrics, 1900–1918</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=6" title="Edit section: Genetics, mutationism and biometrics, 1900–1918"><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 articles: <a href="/wiki/Mutationism" title="Mutationism">Mutationism</a> and <a href="/wiki/Biostatistics" title="Biostatistics">Biostatistics</a></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:William_Bateson.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/William_Bateson.jpg/170px-William_Bateson.jpg" decoding="async" width="170" height="257" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/8/8b/William_Bateson.jpg 1.5x" data-file-width="226" data-file-height="342" /></a><figcaption><a href="/wiki/William_Bateson" title="William Bateson">William Bateson</a> championed <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelism</a>.</figcaption></figure> <p>While carrying out breeding experiments to clarify the mechanism of inheritance in 1900, <a href="/wiki/Hugo_de_Vries" title="Hugo de Vries">Hugo de Vries</a> and <a href="/wiki/Carl_Correns" title="Carl Correns">Carl Correns</a> independently rediscovered <a href="/wiki/Gregor_Mendel" title="Gregor Mendel">Gregor Mendel</a>'s work. News of this reached <a href="/wiki/William_Bateson" title="William Bateson">William Bateson</a> in <a href="/wiki/England" title="England">England</a>, who reported on the paper during a presentation to the <a href="/wiki/Royal_Horticultural_Society" title="Royal Horticultural Society">Royal Horticultural Society</a> in May 1900.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a>, the contributions of each parent retain their integrity, rather than blending with the contribution of the other parent. In the case of a cross between two true-breeding varieties such as Mendel's round and wrinkled peas, the first-generation offspring are all alike, in this case, all round. Allowing these to cross, the original characteristics reappear (segregation): about 3/4 of their offspring are round, 1/4 wrinkled. There is a discontinuity between the appearance of the offspring; de Vries coined the term <a href="/wiki/Allele" title="Allele">allele</a> for a variant form of an <a href="/wiki/Gene" title="Gene">inherited characteristic</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> This reinforced a major division of thought, already present in the 1890s, between gradualists who followed Darwin, and saltationists such as Bateson.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>The two schools were the Mendelians, such as Bateson and de Vries, who favoured mutationism, evolution driven by mutation, based on genes whose alleles <a href="/wiki/Particulate_inheritance" title="Particulate inheritance">segregated discretely</a> like Mendel's peas;<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> and the <a href="/wiki/Biostatistics" title="Biostatistics">biometric school</a>, led by <a href="/wiki/Karl_Pearson" title="Karl Pearson">Karl Pearson</a> and <a href="/wiki/Walter_Frank_Raphael_Weldon" class="mw-redirect" title="Walter Frank Raphael Weldon">Walter Weldon</a>. The biometricians argued vigorously against mutationism, saying that empirical evidence indicated that variation was continuous in most organisms, not discrete as Mendelism seemed to predict; they wrongly believed that Mendelism inevitably implied evolution in discontinuous jumps.<sup id="cite_ref-grafen_69_25-0" class="reference"><a href="#cite_note-grafen_69-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Karl_Pearson,_1912.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Karl_Pearson%2C_1912.jpg/170px-Karl_Pearson%2C_1912.jpg" decoding="async" width="170" height="222" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Karl_Pearson%2C_1912.jpg/255px-Karl_Pearson%2C_1912.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/18/Karl_Pearson%2C_1912.jpg/340px-Karl_Pearson%2C_1912.jpg 2x" data-file-width="713" data-file-height="933" /></a><figcaption><a href="/wiki/Karl_Pearson" title="Karl Pearson">Karl Pearson</a> led the <a href="/wiki/Biostatistics" title="Biostatistics">biometric school</a>.</figcaption></figure> <p>A traditional view is that the biometricians and the Mendelians rejected natural selection and argued for their separate theories for 20 years, the debate only resolved by the development of population genetics.<sup id="cite_ref-grafen_69_25-1" class="reference"><a href="#cite_note-grafen_69-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> A more recent view is that Bateson, de Vries, <a href="/wiki/Thomas_Hunt_Morgan" title="Thomas Hunt Morgan">Thomas Hunt Morgan</a> and <a href="/wiki/Reginald_Punnett" title="Reginald Punnett">Reginald Punnett</a> had by 1918 formed a synthesis of Mendelism and mutationism. The understanding achieved by these geneticists spanned the action of natural selection on alleles (alternative forms of a gene), the <a href="/wiki/Hardy%E2%80%93Weinberg_equilibrium" class="mw-redirect" title="Hardy–Weinberg equilibrium">Hardy–Weinberg equilibrium</a>, the evolution of continuously varying traits (like height), and the probability that a new mutation will become fixed. In this view, the early geneticists accepted natural selection but rejected Darwin's non-Mendelian ideas about variation and heredity, and the synthesis began soon after 1900.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Stoltzfus_29-0" class="reference"><a href="#cite_note-Stoltzfus-29"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The traditional claim that Mendelians rejected the idea of continuous variation is false; as early as 1902, Bateson and Saunders wrote that "If there were even so few as, say, four or five pairs of possible allelomorphs, the various homo- and heterozygous combinations might, on seriation, give so near an approach to a continuous curve, that the purity of the elements would be unsuspected".<sup id="cite_ref-BatesonSaunders1902_30-0" class="reference"><a href="#cite_note-BatesonSaunders1902-30"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Also in 1902, the statistician <a href="/wiki/Udny_Yule" title="Udny Yule">Udny Yule</a> showed mathematically that given multiple factors, Mendel's theory enabled continuous variation. Yule criticised Bateson's approach as confrontational,<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> but failed to prevent the Mendelians and the biometricians from falling out.<sup id="cite_ref-FOOTNOTEProvine200181–82_32-0" class="reference"><a href="#cite_note-FOOTNOTEProvine200181–82-32"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Castle's_hooded_rats,_1911"><span id="Castle.27s_hooded_rats.2C_1911"></span>Castle's hooded rats, 1911</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=7" title="Edit section: Castle's hooded rats, 1911"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Starting in 1906, <a href="/wiki/William_Ernest_Castle" class="mw-redirect" title="William Ernest Castle">William Castle</a> carried out a long study of the effect of selection on coat colour in <a href="/wiki/Rat" title="Rat">rats</a>. The <a href="/wiki/Piebald" title="Piebald">piebald</a> or hooded pattern was <a href="/wiki/Recessive" class="mw-redirect" title="Recessive">recessive</a> to the grey wild type. He crossed hooded rats with both wild and "Irish" types, and then back-crossed the offspring with pure hooded rats. The dark stripe on the back was bigger. He then tried selecting different groups for bigger or smaller stripes for 5 generations and found that it was possible to change the characteristics considerably beyond the initial range of variation. This effectively refuted de Vries's claim that continuous variation was caused by the environment and could not be inherited. By 1911, Castle noted that the results could be explained by Darwinian selection on a heritable variation of a sufficient number of Mendelian genes.<sup id="cite_ref-FOOTNOTEProvine2001109–114_33-0" class="reference"><a href="#cite_note-FOOTNOTEProvine2001109–114-33"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Morgan's_fruit_flies,_1912"><span id="Morgan.27s_fruit_flies.2C_1912"></span>Morgan's fruit flies, 1912</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=8" title="Edit section: Morgan's fruit flies, 1912"><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/Thomas_Hunt_Morgan" title="Thomas Hunt Morgan">Thomas Hunt Morgan</a></div> <p>Thomas Hunt Morgan began his career in genetics as a <a href="/wiki/Saltation_(biology)" title="Saltation (biology)">saltationist</a> and started out trying to demonstrate that mutations could produce new species in fruit flies. However, the experimental work at his lab with the fruit fly, <i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>c<span class="cite-bracket">]</span></a></sup> showed that rather than creating new species in a single step, mutations increased the supply of genetic variation in the population.<sup id="cite_ref-auto_36-1" class="reference"><a href="#cite_note-auto-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> By 1912, after years of work on the genetics of fruit flies, Morgan showed that these insects had many small Mendelian factors (discovered as mutant flies) on which Darwinian evolution could work as if the variation was fully continuous. The way was open for geneticists to conclude that Mendelism supported Darwinism.<sup id="cite_ref-FOOTNOTEProvine2001120–121_38-0" class="reference"><a href="#cite_note-FOOTNOTEProvine2001120–121-38"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p><span class="anchor" id="Woodger"></span> </p> <div class="mw-heading mw-heading2"><h2 id="An_obstruction:_Woodger's_positivism,_1929"><span id="An_obstruction:_Woodger.27s_positivism.2C_1929"></span>An obstruction: Woodger's positivism, 1929</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=9" title="Edit section: An obstruction: Woodger's positivism, 1929"><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">Further information: <a href="/wiki/Joseph_Henry_Woodger" title="Joseph Henry Woodger">Joseph Henry Woodger</a></div> <p>The <a href="/wiki/Theoretical_biology" class="mw-redirect" title="Theoretical biology">theoretical biologist</a> and <a href="/wiki/Philosophy_of_biology" title="Philosophy of biology">philosopher of biology</a> <a href="/wiki/Joseph_Henry_Woodger" title="Joseph Henry Woodger">Joseph Henry Woodger</a> led the introduction of <a href="/wiki/Logical_positivism" title="Logical positivism">positivism</a> into biology with his 1929 book <i>Biological Principles</i>. He saw a mature <a href="/wiki/Science" title="Science">science</a> as being characterised by a framework of <a href="/wiki/Hypothesis" title="Hypothesis">hypotheses</a> that could be verified by facts established by <a href="/wiki/Experiment" title="Experiment">experiments</a>. He criticised the traditional <a href="/wiki/Natural_history" title="Natural history">natural history</a> style of <a href="/wiki/Biology" title="Biology">biology</a>, including the study of <a href="/wiki/Evolution" title="Evolution">evolution</a>, as immature science, since it relied on <a href="/wiki/Narrative" title="Narrative">narrative</a>.<sup id="cite_ref-Smocovitis100_39-0" class="reference"><a href="#cite_note-Smocovitis100-39"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Woodger set out to play the role of <a href="/wiki/Robert_Boyle" title="Robert Boyle">Robert Boyle</a>'s 1661 <i><a href="/wiki/Sceptical_Chymist" class="mw-redirect" title="Sceptical Chymist">Sceptical Chymist</a></i>, intending to convert the subject of biology into a formal, unified science, and ultimately, following the <a href="/wiki/Vienna_Circle" title="Vienna Circle">Vienna Circle</a> of logical positivists like <a href="/wiki/Otto_Neurath" title="Otto Neurath">Otto Neurath</a> and <a href="/wiki/Rudolf_Carnap" title="Rudolf Carnap">Rudolf Carnap</a>, <a href="/wiki/Reductionism" title="Reductionism">to reduce</a> biology to physics and chemistry. His efforts stimulated the biologist <a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a> to push for the axiomatisation of biology, and by influencing thinkers such as Huxley, helped to bring about the modern synthesis.<sup id="cite_ref-Smocovitis100_39-1" class="reference"><a href="#cite_note-Smocovitis100-39"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> The positivist climate made natural history unfashionable, and in America, research and university-level teaching on evolution declined almost to nothing by the late 1930s. The Harvard physiologist <a href="/wiki/William_John_Crozier" title="William John Crozier">William John Crozier</a> told his students that evolution was not even a science: "You can't experiment with two million years!"<sup id="cite_ref-Smocovitis114_40-0" class="reference"><a href="#cite_note-Smocovitis114-40"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p><p>The tide of opinion turned with the adoption of <a href="/wiki/Mathematical_model" title="Mathematical model">mathematical modelling</a> and <a href="/wiki/Controlled_experiment" class="mw-redirect" title="Controlled experiment">controlled experimentation</a> in population genetics, combining genetics, ecology and evolution in a framework acceptable to positivism.<sup id="cite_ref-Smocovitis119_41-0" class="reference"><a href="#cite_note-Smocovitis119-41"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Elements_of_the_synthesis">Elements of the synthesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=10" title="Edit section: Elements of the synthesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Fisher_and_Haldane's_mathematical_population_genetics,_1918–1930"><span id="Fisher_and_Haldane.27s_mathematical_population_genetics.2C_1918.E2.80.931930"></span>Fisher and Haldane's mathematical population genetics, 1918–1930</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=11" title="Edit section: Fisher and Haldane's mathematical population genetics, 1918–1930"><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/A_Mathematical_Theory_of_Natural_and_Artificial_Selection" title="A Mathematical Theory of Natural and Artificial Selection">A Mathematical Theory of Natural and Artificial Selection</a></div> <p>In 1918, R. A. Fisher wrote "<a href="/wiki/The_Correlation_between_Relatives_on_the_Supposition_of_Mendelian_Inheritance" title="The Correlation between Relatives on the Supposition of Mendelian Inheritance">The Correlation between Relatives on the Supposition of Mendelian Inheritance</a>,"<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> which showed how continuous variation could come from a number of discrete <a href="/wiki/Locus_(genetics)" title="Locus (genetics)">genetic loci</a>. In this and other papers, culminating in his 1930 book <i><a href="/wiki/The_Genetical_Theory_of_Natural_Selection" title="The Genetical Theory of Natural Selection">The Genetical Theory of Natural Selection</a></i>,<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> Fisher showed how Mendelian genetics was consistent with the idea of evolution by natural selection.<sup id="cite_ref-Larson221-243_44-0" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>d<span class="cite-bracket">]</span></a></sup> </p><p>In the 1920s, <a href="/wiki/A_Mathematical_Theory_of_Natural_and_Artificial_Selection" title="A Mathematical Theory of Natural and Artificial Selection">a series of papers</a> by <a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a> analyzed real-world examples of natural selection, such as the <a href="/wiki/Peppered_moth_evolution" title="Peppered moth evolution">evolution of industrial melanism in peppered moths</a>.<sup id="cite_ref-Larson221-243_44-1" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> and showed that natural selection could work even faster than Fisher had assumed.<sup id="cite_ref-Bowler325-339_47-0" class="reference"><a href="#cite_note-Bowler325-339-47"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Both of these scholars, and others, such as Dobzhansky and Wright, wanted to raise biology to the standards of the physical sciences by basing it on mathematical modeling and empirical testing. Natural selection, once considered unverifiable, was becoming predictable, measurable, and testable.<sup id="cite_ref-Smocovitis122_48-0" class="reference"><a href="#cite_note-Smocovitis122-48"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="De_Beer's_embryology,_1930"><span id="De_Beer.27s_embryology.2C_1930"></span>De Beer's embryology, 1930</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=12" title="Edit section: De Beer's embryology, 1930"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The traditional view is that <a href="/wiki/Developmental_biology" title="Developmental biology">developmental biology</a> played little part in the modern synthesis,<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> but in his 1930 book <i><a href="/wiki/Embryos_and_Ancestors" class="mw-redirect" title="Embryos and Ancestors">Embryos and Ancestors</a></i>, the evolutionary embryologist <a href="/wiki/Gavin_de_Beer" title="Gavin de Beer">Gavin de Beer</a> anticipated <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a><sup id="cite_ref-Held_50-0" class="reference"><a href="#cite_note-Held-50"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> by showing that evolution could occur by <a href="/wiki/Heterochrony" title="Heterochrony">heterochrony</a>,<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> such as in <a href="/wiki/Paedomorphosis" class="mw-redirect" title="Paedomorphosis">the retention of juvenile features in the adult</a>.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> This, de Beer argued, could cause apparently sudden changes in the <a href="/wiki/Fossil_record" class="mw-redirect" title="Fossil record">fossil record</a>, since embryos fossilise poorly. As the gaps in the fossil record had been used as an argument against Darwin's gradualist evolution, de Beer's explanation supported the Darwinian position.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> However, despite de Beer, the modern synthesis largely ignored embryonic development when explaining the form of organisms, since population genetics appeared to be an adequate explanation of how such forms evolved.<sup id="cite_ref-Gilbert1991_54-0" class="reference"><a href="#cite_note-Gilbert1991-54"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>e<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Wright's_adaptive_landscape,_1932"><span id="Wright.27s_adaptive_landscape.2C_1932"></span>Wright's adaptive landscape, 1932</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=13" title="Edit section: Wright's adaptive landscape, 1932"><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:Fitness-landscape-cartoon.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Fitness-landscape-cartoon.png/220px-Fitness-landscape-cartoon.png" decoding="async" width="220" height="108" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Fitness-landscape-cartoon.png/330px-Fitness-landscape-cartoon.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/67/Fitness-landscape-cartoon.png/440px-Fitness-landscape-cartoon.png 2x" data-file-width="484" data-file-height="238" /></a><figcaption><a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a> introduced the idea of a <a href="/wiki/Fitness_landscape" title="Fitness landscape">fitness landscape</a> with local optima.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Population_genetics#History" title="Population genetics">Population genetics § History</a></div> <p>The population geneticist <a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a> focused on combinations of genes that interacted as complexes, and the effects of <a href="/wiki/Inbreeding" title="Inbreeding">inbreeding</a> on small relatively isolated populations, which could be subject to <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a>. In a 1932 paper, he introduced the concept of an <a href="/wiki/Fitness_landscape" title="Fitness landscape">adaptive landscape</a> in which phenomena such as cross breeding and genetic drift in small populations could push them away from adaptive peaks, which would in turn allow natural selection to push them towards new adaptive peaks.<sup id="cite_ref-Larson221-243_44-2" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> Wright's model would appeal to field naturalists such as Theodosius Dobzhansky and Ernst Mayr who were becoming aware of the importance of geographical isolation in real world populations.<sup id="cite_ref-Bowler325-339_47-1" class="reference"><a href="#cite_note-Bowler325-339-47"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> The work of Fisher, Haldane and Wright helped to found the discipline of theoretical population genetics.<sup id="cite_ref-auto1_59-0" class="reference"><a href="#cite_note-auto1-59"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dobzhansky's_evolutionary_genetics,_1937"><span id="Dobzhansky.27s_evolutionary_genetics.2C_1937"></span>Dobzhansky's evolutionary genetics, 1937</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=14" title="Edit section: Dobzhansky's evolutionary genetics, 1937"><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">Further information: <a href="/wiki/Genetics_and_the_Origin_of_Species" title="Genetics and the Origin of Species">Genetics and the Origin of Species</a></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Drosophila_pseudoobscura-Male.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Drosophila_pseudoobscura-Male.png/220px-Drosophila_pseudoobscura-Male.png" decoding="async" width="220" height="98" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/c/cb/Drosophila_pseudoobscura-Male.png 1.5x" data-file-width="256" data-file-height="114" /></a><figcaption><i><a href="/wiki/Drosophila_pseudoobscura" title="Drosophila pseudoobscura">Drosophila pseudoobscura</a></i>, the fruit fly which served as <a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a>'s <a href="/wiki/Model_organism" title="Model organism">model organism</a></figcaption></figure> <p><a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Theodosius Dobzhansky</a>, an immigrant from the <a href="/wiki/Soviet_Union" title="Soviet Union">Soviet Union</a> to the <a href="/wiki/United_States" title="United States">United States</a>, who had been a postdoctoral worker in Morgan's fruit fly lab, was one of the first to apply genetics to natural populations. He worked mostly with <i><a href="/wiki/Drosophila_pseudoobscura" title="Drosophila pseudoobscura">Drosophila pseudoobscura</a></i>. He says pointedly: "Russia has a variety of climates from the Arctic to sub-tropical... Exclusively laboratory workers who neither possess nor wish to have any knowledge of living beings in nature were and are in a minority."<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Not surprisingly, there were other <a href="/wiki/Russia" title="Russia">Russian</a> geneticists with similar ideas, though for some time their work was known to only a few in the <a href="/wiki/Western_world" title="Western world">West</a>. His 1937 work <i><a href="/wiki/Genetics_and_the_Origin_of_Species" title="Genetics and the Origin of Species">Genetics and the Origin of Species</a></i><sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> was a key step in bridging the gap between population geneticists and field naturalists. It presented the conclusions reached by Fisher, Haldane, and especially Wright in their highly mathematical papers in a form that was easily accessible to others.<sup id="cite_ref-Larson221-243_44-3" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bowler325-339_47-2" class="reference"><a href="#cite_note-Bowler325-339-47"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Further, Dobzhansky asserted the physicality, and hence the biological reality, of the mechanisms of inheritance: that evolution was based on material genes, arranged in a string on physical hereditary structures, the <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a>, and <a href="/wiki/Genetic_linkage" title="Genetic linkage">linked</a> more or less strongly to each other according to their actual physical distances on the chromosomes. As with Haldane and Fisher, Dobzhansky's "evolutionary genetics"<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> was a genuine science, now unifying cell biology, genetics, and both micro and macroevolution.<sup id="cite_ref-Smocovitis122_48-1" class="reference"><a href="#cite_note-Smocovitis122-48"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> His work emphasized that real-world populations had far more genetic variability than the early population geneticists had assumed in their models and that genetically distinct sub-populations were important. Dobzhansky argued that natural selection worked to maintain genetic diversity as well as by driving change. He was influenced by his exposure in the 1920s to the work of <a href="/wiki/Sergei_Chetverikov" title="Sergei Chetverikov">Sergei Chetverikov</a>, who had looked at the role of recessive genes in maintaining a reservoir of genetic variability in a population, before his work was shut down by the rise of <a href="/wiki/Lysenkoism" title="Lysenkoism">Lysenkoism</a> in the <a href="/wiki/Soviet_Union" title="Soviet Union">Soviet Union</a>.<sup id="cite_ref-Larson221-243_44-4" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bowler325-339_47-3" class="reference"><a href="#cite_note-Bowler325-339-47"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> By 1937, Dobzhansky was able to argue that mutations were the main source of evolutionary changes and variability, along with chromosome rearrangements, effects of genes on their neighbours during development, and polyploidy. Next, genetic drift (he used the term in 1941), selection, migration, and geographical isolation could change gene frequencies. Thirdly, mechanisms like ecological or sexual isolation and hybrid sterility could fix the results of the earlier processes.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ford's_ecological_genetics,_1940"><span id="Ford.27s_ecological_genetics.2C_1940"></span>Ford's ecological genetics, 1940</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=15" title="Edit section: Ford's ecological genetics, 1940"><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:Callimorpha.dominula.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Callimorpha.dominula.jpg/220px-Callimorpha.dominula.jpg" decoding="async" width="220" height="123" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Callimorpha.dominula.jpg/330px-Callimorpha.dominula.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/28/Callimorpha.dominula.jpg/440px-Callimorpha.dominula.jpg 2x" data-file-width="1095" data-file-height="612" /></a><figcaption><a href="/wiki/E._B._Ford" title="E. B. Ford">E. B. Ford</a> studied <a href="/wiki/Polymorphism_(biology)#Scarlet_tiger_moth" title="Polymorphism (biology)">polymorphism</a> in the <a href="/wiki/Scarlet_tiger_moth" title="Scarlet tiger moth">scarlet tiger moth</a> for many years.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></div> <p><a href="/wiki/E._B._Ford" title="E. B. Ford">E. B. Ford</a> was an experimental naturalist who wanted to test natural selection in nature, virtually inventing the field of <a href="/wiki/Ecological_genetics" title="Ecological genetics">ecological genetics</a>.<sup id="cite_ref-Ford1964_66-0" class="reference"><a href="#cite_note-Ford1964-66"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> His work on natural selection in wild populations of butterflies and moths was the first to show that predictions made by R. A. Fisher were correct. In 1940, he was the first to describe and define <a href="/wiki/Polymorphism_(biology)#Genetic_polymorphism" title="Polymorphism (biology)">genetic polymorphism</a>, and to predict that <a href="/wiki/Human_blood_group_systems" title="Human blood group systems">human blood group polymorphisms</a> might be maintained in the population by providing some protection against disease.<sup id="cite_ref-Ford1964_66-1" class="reference"><a href="#cite_note-Ford1964-66"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> His 1949 book <i>Mendelism and Evolution</i><sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> helped to persuade Dobzhansky to change the emphasis in the third edition of his famous textbook <i>Genetics and the Origin of Species</i> from drift to selection.<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Schmalhausen's_stabilizing_selection,_1941"><span id="Schmalhausen.27s_stabilizing_selection.2C_1941"></span>Schmalhausen's stabilizing selection, 1941</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=16" title="Edit section: Schmalhausen's stabilizing selection, 1941"><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">Further information: <a href="/wiki/Stabilizing_selection" title="Stabilizing selection">Stabilizing selection</a></div> <p><a href="/wiki/Ivan_Schmalhausen" title="Ivan Schmalhausen">Ivan Schmalhausen</a> developed the theory of <a href="/wiki/Stabilizing_selection" title="Stabilizing selection">stabilizing selection</a>, the idea that selection can preserve a trait at some value, publishing a paper in Russian titled "Stabilizing selection and its place among factors of evolution" in 1941 and a monograph <i>Factors of Evolution: The Theory of Stabilizing Selection</i><sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> in 1945. He developed it from J. M. Baldwin's 1902 concept that changes induced by the environment will ultimately be replaced by hereditary changes (including the <a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a> on behaviour), following that theory's implications to their Darwinian conclusion, and bringing him into conflict with Lysenkoism. Schmalhausen observed that stabilizing selection would remove most variations from the norm, most mutations being harmful.<sup id="cite_ref-Levit2006_71-0" class="reference"><a href="#cite_note-Levit2006-71"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Adams1988_72-0" class="reference"><a href="#cite_note-Adams1988-72"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> Dobzhansky called the work "an important missing link in the modern view of evolution".<sup id="cite_ref-FOOTNOTEMayrProvine1998ix_74-0" class="reference"><a href="#cite_note-FOOTNOTEMayrProvine1998ix-74"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Huxley's_popularising_synthesis,_1942"><span id="Huxley.27s_popularising_synthesis.2C_1942"></span>Huxley's popularising synthesis, 1942</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=17" title="Edit section: Huxley's popularising synthesis, 1942"><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/Evolution:_The_Modern_Synthesis" title="Evolution: The Modern Synthesis">Evolution: The Modern Synthesis</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Julian_Huxley_1964.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Julian_Huxley_1964.jpg/170px-Julian_Huxley_1964.jpg" decoding="async" width="170" height="219" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Julian_Huxley_1964.jpg/255px-Julian_Huxley_1964.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Julian_Huxley_1964.jpg/340px-Julian_Huxley_1964.jpg 2x" data-file-width="1760" data-file-height="2272" /></a><figcaption><a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a> presented a serious but popularising version of the theory in his 1942 book <i><a href="/wiki/Evolution:_The_Modern_Synthesis" title="Evolution: The Modern Synthesis">Evolution: The Modern Synthesis</a></i>.</figcaption></figure> <p>In 1942, <a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a>'s serious but popularising<sup id="cite_ref-Ruse_75-0" class="reference"><a href="#cite_note-Ruse-75"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lamm_76-0" class="reference"><a href="#cite_note-Lamm-76"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> <i><a href="/wiki/Evolution:_The_Modern_Synthesis" title="Evolution: The Modern Synthesis">Evolution: The Modern Synthesis</a></i><sup id="cite_ref-FOOTNOTEHuxley2010_2-1" class="reference"><a href="#cite_note-FOOTNOTEHuxley2010-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> introduced a name for the synthesis and intentionally set out to promote a "synthetic point of view" on the evolutionary process. He imagined a wide synthesis of many sciences: genetics, developmental physiology, ecology, systematics, palaeontology, cytology, and mathematical analysis of biology, and assumed that evolution would proceed differently in different groups of organisms according to how their genetic material was organised and their strategies for reproduction, leading to progressive but varying evolutionary trends.<sup id="cite_ref-Lamm_76-1" class="reference"><a href="#cite_note-Lamm-76"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> His vision was of an "evolutionary humanism",<sup id="cite_ref-Smocovitis138_77-0" class="reference"><a href="#cite_note-Smocovitis138-77"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> with a system of ethics and a meaningful place for "Man" in the world grounded in a unified theory of evolution which would demonstrate progress leading to humanity at its summit. Natural selection was in his view a "fact of nature capable of verification by observation and experiment", while the "period of synthesis" of the 1920s and 1930s had formed a "more unified science",<sup id="cite_ref-Smocovitis138_77-1" class="reference"><a href="#cite_note-Smocovitis138-77"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> rivalling physics and enabling the "rebirth of Darwinism".<sup id="cite_ref-Smocovitis138_77-2" class="reference"><a href="#cite_note-Smocovitis138-77"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> </p><p>However, the book was not the research text that it appeared to be. In the view of the philosopher of science <a href="/wiki/Michael_Ruse" title="Michael Ruse">Michael Ruse</a>, and in Huxley's own opinion, Huxley was "a generalist, a synthesizer of ideas, rather than a specialist".<sup id="cite_ref-Ruse_75-1" class="reference"><a href="#cite_note-Ruse-75"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> Ruse observes that Huxley wrote as if he were adding empirical evidence to the mathematical framework established by Fisher and the population geneticists, but that this was not so. Huxley avoided mathematics, for instance not even mentioning <a href="/wiki/Fisher%27s_fundamental_theorem_of_natural_selection" title="Fisher's fundamental theorem of natural selection">Fisher's fundamental theorem of natural selection</a>. Instead, Huxley used a mass of examples to demonstrate that natural selection is powerful and that it works on Mendelian genes. The book was successful in its goal of persuading readers of the reality of evolution, effectively illustrating topics such as <a href="/wiki/Island_biogeography" class="mw-redirect" title="Island biogeography">island biogeography</a>, <a href="/wiki/Speciation" title="Speciation">speciation</a>, and competition. Huxley further showed that the appearance of long-term <a href="/wiki/Orthogenesis" title="Orthogenesis">orthogenetic trends</a> – predictable directions for evolution – in the fossil record were readily explained as <a href="/wiki/Allometry" title="Allometry">allometric growth</a> (since parts are interconnected). All the same, Huxley did not reject orthogenesis out of hand, but maintained a belief in progress all his life, with <i><a href="/wiki/Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a></i> as the endpoint, and he had since 1912 been influenced by the <a href="/wiki/Vitalism" title="Vitalism">vitalist</a> philosopher <a href="/wiki/Henri_Bergson" title="Henri Bergson">Henri Bergson</a>, though in public he maintained an atheistic position on evolution.<sup id="cite_ref-Ruse_75-2" class="reference"><a href="#cite_note-Ruse-75"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> Huxley's belief in progress within evolution and evolutionary humanism was shared in various forms by Dobzhansky, Mayr, Simpson and Stebbins, all of them writing about "the future of Mankind". Both Huxley and Dobzhansky admired the palaeontologist priest <a href="/wiki/Pierre_Teilhard_de_Chardin" title="Pierre Teilhard de Chardin">Pierre Teilhard de Chardin</a>, Huxley writing the introduction to Teilhard's 1955 book on orthogenesis, <i><a href="/wiki/The_Phenomenon_of_Man" title="The Phenomenon of Man">The Phenomenon of Man</a></i>. This vision required evolution to be seen as the central and guiding principle of biology.<sup id="cite_ref-Smocovitis138_77-3" class="reference"><a href="#cite_note-Smocovitis138-77"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Mayr's_allopatric_speciation,_1942"><span id="Mayr.27s_allopatric_speciation.2C_1942"></span>Mayr's allopatric speciation, 1942</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=18" title="Edit section: Mayr's allopatric speciation, 1942"><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 articles: <a href="/wiki/Systematics_and_the_Origin_of_Species" title="Systematics and the Origin of Species">Systematics and the Origin of Species</a> and <a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric speciation</a></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Ernst_Mayr_PLoS.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/af/Ernst_Mayr_PLoS.jpg/220px-Ernst_Mayr_PLoS.jpg" decoding="async" width="220" height="154" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/af/Ernst_Mayr_PLoS.jpg/330px-Ernst_Mayr_PLoS.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/af/Ernst_Mayr_PLoS.jpg/440px-Ernst_Mayr_PLoS.jpg 2x" data-file-width="1964" data-file-height="1378" /></a><figcaption><a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Ernst Mayr</a> argued that <a href="/wiki/Allopatric_speciation" title="Allopatric speciation">geographic isolation</a> was needed to provide sufficient <a href="/wiki/Reproductive_isolation" title="Reproductive isolation">reproductive isolation</a> for <a href="/wiki/Speciation" title="Speciation">new species to form</a>.</figcaption></figure> <p><a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Ernst Mayr</a>'s key contribution to the synthesis was <i><a href="/wiki/Systematics_and_the_Origin_of_Species" title="Systematics and the Origin of Species">Systematics and the Origin of Species</a></i>, published in 1942.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> It asserted the importance of and set out to explain population variation in evolutionary processes including speciation. He analysed in particular the effects of <a href="/wiki/Taxon" title="Taxon">polytypic</a> species, geographic variation, and isolation by geographic and other means.<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> Mayr emphasized the importance of <a href="/wiki/Allopatric_speciation" title="Allopatric speciation">allopatric speciation</a>, where geographically isolated sub-populations diverge so far that <a href="/wiki/Reproductive_isolation" title="Reproductive isolation">reproductive isolation</a> occurs. He was skeptical of the reality of <a href="/wiki/Sympatric_speciation" title="Sympatric speciation">sympatric speciation</a> believing that geographical isolation was a prerequisite for building up intrinsic (reproductive) isolating mechanisms. Mayr also introduced the <a href="/wiki/Species_problem#Biological_species_concept" class="mw-redirect" title="Species problem">biological species concept</a> that defined a species as a group of interbreeding or potentially interbreeding populations that were reproductively isolated from all other populations.<sup id="cite_ref-Larson221-243_44-5" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bowler325-339_47-4" class="reference"><a href="#cite_note-Bowler325-339-47"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-80" class="reference"><a href="#cite_note-80"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEMayr1982_81-0" class="reference"><a href="#cite_note-FOOTNOTEMayr1982-81"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> Before he left <a href="/wiki/Germany" title="Germany">Germany</a> for the United States in 1930, Mayr had been influenced by the work of the German biologist <a href="/wiki/Bernhard_Rensch" title="Bernhard Rensch">Bernhard Rensch</a>, who in the 1920s had analyzed the geographic distribution of polytypic species, paying particular attention to how variations between populations correlated with factors such as differences in climate.<sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Equine_evolution.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Equine_evolution.jpg/260px-Equine_evolution.jpg" decoding="async" width="260" height="222" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Equine_evolution.jpg/390px-Equine_evolution.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Equine_evolution.jpg/520px-Equine_evolution.jpg 2x" data-file-width="9623" data-file-height="8223" /></a><figcaption><a href="/wiki/George_Gaylord_Simpson" title="George Gaylord Simpson">George Gaylord Simpson</a> argued against the naive view that <a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">evolution such as of the horse</a> took place in a "straight-line". He noted that any chosen line is one path in a complex branching tree, <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> having no <a href="/wiki/Orthogenesis" title="Orthogenesis">imposed direction</a>.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Simpson's_palaeontology,_1944"><span id="Simpson.27s_palaeontology.2C_1944"></span>Simpson's palaeontology, 1944</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=19" title="Edit section: Simpson's palaeontology, 1944"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/George_Gaylord_Simpson" title="George Gaylord Simpson">George Gaylord Simpson</a> was responsible for showing that the modern synthesis was compatible with palaeontology in his 1944 book <i><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and Mode in Evolution</a></i>. Simpson's work was crucial because so many palaeontologists had disagreed, in some cases vigorously, with the idea that natural selection was the main mechanism of evolution. It showed that the trends of linear progression (in for example the <a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">evolution of the horse</a>) that earlier palaeontologists had used as support for <a href="/wiki/Lamarckism#Neo-Lamarckism" title="Lamarckism">neo-Lamarckism</a> and orthogenesis did not hold up under careful examination. Instead, the <a href="/wiki/Fossil#Dating" title="Fossil">fossil record</a> was consistent with the irregular, branching, and non-directional pattern predicted by the modern synthesis.<sup id="cite_ref-Larson221-243_44-6" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bowler325-339_47-5" class="reference"><a href="#cite_note-Bowler325-339-47"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Society_for_the_Study_of_Evolution,_1946"><span id="Society_for_the_Study_of_Evolution.2C_1946"></span>Society for the Study of Evolution, 1946</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=20" title="Edit section: Society for the Study of Evolution, 1946"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>During <a href="/wiki/World_War_II" title="World War II">World War II</a>, Mayr edited a series of bulletins of the Committee on Common Problems of Genetics, Paleontology, and Systematics, formed in 1943, reporting on discussions of a "synthetic attack" on the interdisciplinary problems of evolution. In 1946, the committee became the Society for the Study of Evolution, with Mayr, Dobzhansky and Sewall Wright the first of the signatories. Mayr became the editor of its journal, <i><a href="/wiki/Evolution_(journal)" title="Evolution (journal)">Evolution</a></i>. From Mayr and Dobzhansky's point of view, suggests the historian of science Betty Smocovitis, Darwinism was reborn, evolutionary biology was legitimised, and genetics and evolution were synthesised into a newly unified science. Everything fitted into the new framework, except "heretics" like <a href="/wiki/Richard_Goldschmidt" title="Richard Goldschmidt">Richard Goldschmidt</a> who annoyed Mayr and Dobzhansky by insisting on the possibility of <a href="/wiki/Mutationism" title="Mutationism">speciation by macromutation</a>, creating "hopeful monsters". The result was "bitter controversy".<sup id="cite_ref-Smocovitis153_56-1" class="reference"><a href="#cite_note-Smocovitis153-56"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Polyploidization.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/04/Polyploidization.svg/220px-Polyploidization.svg.png" decoding="async" width="220" height="132" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/04/Polyploidization.svg/330px-Polyploidization.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/04/Polyploidization.svg/440px-Polyploidization.svg.png 2x" data-file-width="200" data-file-height="120" /></a><figcaption><a href="/wiki/Speciation" title="Speciation">Speciation</a> via <a href="/wiki/Polyploidy" title="Polyploidy">polyploidy</a>: a <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a> cell may fail to separate during <a href="/wiki/Meiosis" title="Meiosis">meiosis</a>, producing diploid <a href="/wiki/Gamete" title="Gamete">gametes</a>, which self-fertilize to produce a fertile tetraploid <a href="/wiki/Zygote" title="Zygote">zygote</a> that cannot interbreed with its parent species.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Stebbins's_botany,_1950"><span id="Stebbins.27s_botany.2C_1950"></span>Stebbins's botany, 1950</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=21" title="Edit section: Stebbins's botany, 1950"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The botanist <a href="/wiki/G._Ledyard_Stebbins" title="G. Ledyard Stebbins">G. Ledyard Stebbins</a> extended the synthesis to encompass <a href="/wiki/Botany" title="Botany">botany</a>. He described the important effects on <a href="/wiki/Speciation" title="Speciation">speciation</a> of <a href="/wiki/Hybridisation_(biology)" class="mw-redirect" title="Hybridisation (biology)">hybridization</a> and <a href="/wiki/Polyploidy" title="Polyploidy">polyploidy</a> in plants in his 1950 book <i><a href="/wiki/Variation_and_Evolution_in_Plants" title="Variation and Evolution in Plants">Variation and Evolution in Plants</a></i>. These permitted evolution to proceed rapidly at times, polyploidy in particular evidently being able to create new species effectively instantaneously.<sup id="cite_ref-Larson221-243_44-7" class="reference"><a href="#cite_note-Larson221-243-44"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smocovitis2001_85-0" class="reference"><a href="#cite_note-Smocovitis2001-85"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Definitions_by_the_founders">Definitions by the founders</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=22" title="Edit section: Definitions by the founders"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The modern synthesis was defined differently by its various founders, with differing numbers of basic postulates, as shown in the table. </p> <table class="wikitable"> <caption>Definitions of the modern synthesis by its founders, as they numbered them </caption> <tbody><tr> <th>Component</th> <th><a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Mayr</a> 1959</th> <th><a href="/wiki/G._Ledyard_Stebbins" title="G. Ledyard Stebbins">Stebbins</a>, 1966</th> <th><a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Dobzhansky</a>, 1974 </th></tr> <tr> <td><b><a href="/wiki/Mutation" title="Mutation">Mutation</a></b></td> <td>(1) <a href="/wiki/Random" class="mw-redirect" title="Random">Randomness</a> in all events that produce new genotypes, e.g. mutation <sup id="cite_ref-MayrEssence1959_86-0" class="reference"><a href="#cite_note-MayrEssence1959-86"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup></td> <td>(1) a source of variability, but not of <a href="/wiki/Orthogenesis" title="Orthogenesis">direction</a><sup id="cite_ref-Stebbins1966_87-0" class="reference"><a href="#cite_note-Stebbins1966-87"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup></td> <td>(1) yields genetic raw materials<sup id="cite_ref-Dobzhansky1974_88-0" class="reference"><a href="#cite_note-Dobzhansky1974-88"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><b><a href="/wiki/Genetic_recombination" title="Genetic recombination">Recombination</a></b></td> <td>(1) Randomness in recombination, <a href="/wiki/Fertilisation" title="Fertilisation">fertilisation</a><sup id="cite_ref-MayrEssence1959_86-1" class="reference"><a href="#cite_note-MayrEssence1959-86"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup></td> <td>(2) a source of variability, but not of <a href="/wiki/Orthogenesis" title="Orthogenesis">direction</a><sup id="cite_ref-Stebbins1966_87-1" class="reference"><a href="#cite_note-Stebbins1966-87"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup></td> <td> </td></tr> <tr> <td><b><a href="/wiki/Chromosome" title="Chromosome">Chromosomal organisation</a></b></td> <td></td> <td>(3) affects genetic linkage, arranges variation in <a href="/wiki/Gene_pool" title="Gene pool">gene pool</a><sup id="cite_ref-Stebbins1966_87-2" class="reference"><a href="#cite_note-Stebbins1966-87"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup></td> <td> </td></tr> <tr> <td><b><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></b></td> <td>(2) is only direction-giving factor,<sup id="cite_ref-MayrEssence1959_86-2" class="reference"><a href="#cite_note-MayrEssence1959-86"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-MayrSynth1980_89-0" class="reference"><a href="#cite_note-MayrSynth1980-89"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> as seen in <a href="/wiki/Adaptation" title="Adaptation">adaptations</a> to physical and biotic environment<sup id="cite_ref-MayrEssence1959_86-3" class="reference"><a href="#cite_note-MayrEssence1959-86"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup></td> <td>(4) guides changes to <a href="/wiki/Gene_pool" title="Gene pool">gene pool</a><sup id="cite_ref-Stebbins1966_87-3" class="reference"><a href="#cite_note-Stebbins1966-87"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup></td> <td>(2) constructs evolutionary changes from genetic raw materials<sup id="cite_ref-Dobzhansky1974_88-1" class="reference"><a href="#cite_note-Dobzhansky1974-88"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><b><a href="/wiki/Reproductive_isolation" title="Reproductive isolation">Reproductive isolation</a></b></td> <td></td> <td>(5) limits direction in which selection can guide the population<sup id="cite_ref-Stebbins1966_87-4" class="reference"><a href="#cite_note-Stebbins1966-87"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup></td> <td>(3) makes divergence irreversible in <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual organisms</a><sup id="cite_ref-Dobzhansky1974_88-2" class="reference"><a href="#cite_note-Dobzhansky1974-88"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="After_the_synthesis">After the synthesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=23" title="Edit section: After the synthesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>After the synthesis, evolutionary biology continued to develop with major contributions from workers including W. D. Hamilton,<sup id="cite_ref-Hamilton_90-0" class="reference"><a href="#cite_note-Hamilton-90"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> George C. Williams,<sup id="cite_ref-Williams_91-0" class="reference"><a href="#cite_note-Williams-91"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> E. O. Wilson,<sup id="cite_ref-YudellDesalle2000_92-0" class="reference"><a href="#cite_note-YudellDesalle2000-92"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> Edward B. Lewis<sup id="cite_ref-Gilbert2000_93-0" class="reference"><a href="#cite_note-Gilbert2000-93"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> and others. </p> <div class="mw-heading mw-heading3"><h3 id="Hamilton's_inclusive_fitness,_1964"><span id="Hamilton.27s_inclusive_fitness.2C_1964"></span>Hamilton's inclusive fitness, 1964</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=24" title="Edit section: Hamilton's inclusive fitness, 1964"><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">Further information: <a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive fitness</a> and <a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></div> <p>In 1964, <a href="/wiki/W._D._Hamilton" title="W. D. Hamilton">W. D. Hamilton</a> published two papers on "The Genetical Evolution of Social Behaviour". These defined <a href="/wiki/Inclusive_fitness" title="Inclusive fitness">inclusive fitness</a> as the number of offspring equivalents an individual rears, rescues or otherwise supports through its behaviour. This was contrasted with personal reproductive fitness, the number of offspring that the individual directly begets. Hamilton, and others such as <a href="/wiki/John_Maynard_Smith" title="John Maynard Smith">John Maynard Smith</a>, argued that a gene's success consisted in maximising the number of copies of itself, either by begetting them or by indirectly encouraging begetting by related individuals who shared the gene, the theory of <a href="/wiki/Kin_selection" title="Kin selection">kin selection</a>.<sup id="cite_ref-Hamilton_90-1" class="reference"><a href="#cite_note-Hamilton-90"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-94" class="reference"><a href="#cite_note-94"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Williams's_gene-centred_evolution,_1966"><span id="Williams.27s_gene-centred_evolution.2C_1966"></span>Williams's gene-centred evolution, 1966</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=25" title="Edit section: Williams's gene-centred evolution, 1966"><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">Further information: <a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a> and <a href="/wiki/Adaptation_and_Natural_Selection" title="Adaptation and Natural Selection">Adaptation and Natural Selection</a></div> <p>In 1966, <a href="/wiki/George_C._Williams_(biologist)" class="mw-redirect" title="George C. Williams (biologist)">George C. Williams</a> published <i><a href="/wiki/Adaptation_and_Natural_Selection" title="Adaptation and Natural Selection">Adaptation and Natural Selection</a></i>, outlined a <a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">gene-centred view of evolution</a> following Hamilton's concepts, disputing the idea of <a href="/wiki/Evolutionary_progress" class="mw-redirect" title="Evolutionary progress">evolutionary progress</a>, and attacking the then widespread theory of <a href="/wiki/Group_selection" title="Group selection">group selection</a>. Williams argued that natural selection worked by changing the frequency of alleles, and could not work at the level of groups.<sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Williams_91-1" class="reference"><a href="#cite_note-Williams-91"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> Gene-centred evolution was popularised by <a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a> in his 1976 book <i><a href="/wiki/The_Selfish_Gene" title="The Selfish Gene">The Selfish Gene</a></i> and developed in his more technical writings.<sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-extendedphenotype_97-0" class="reference"><a href="#cite_note-extendedphenotype-97"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Wilson's_sociobiology,_1975"><span id="Wilson.27s_sociobiology.2C_1975"></span>Wilson's sociobiology, 1975</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=26" title="Edit section: Wilson's sociobiology, 1975"><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:Atta.cephalotes.gamut.selection.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Atta.cephalotes.gamut.selection.jpg/220px-Atta.cephalotes.gamut.selection.jpg" decoding="async" width="220" height="134" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Atta.cephalotes.gamut.selection.jpg/330px-Atta.cephalotes.gamut.selection.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Atta.cephalotes.gamut.selection.jpg/440px-Atta.cephalotes.gamut.selection.jpg 2x" data-file-width="1941" data-file-height="1179" /></a><figcaption><a href="/wiki/Ant" title="Ant">Ant</a> <a href="/wiki/Sociobiology" title="Sociobiology">societies have evolved</a> elaborate <a href="/wiki/Caste_(biology)" class="mw-redirect" title="Caste (biology)">caste</a> structures, widely different in size and function.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></div> <p>In 1975, <a href="/wiki/E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a> published his controversial<sup id="cite_ref-Fisher1994_98-0" class="reference"><a href="#cite_note-Fisher1994-98"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> book <i><a href="/wiki/Sociobiology:_The_New_Synthesis" title="Sociobiology: The New Synthesis">Sociobiology: The New Synthesis</a></i>, the subtitle alluding to the modern synthesis<sup id="cite_ref-YudellDesalle2000_92-1" class="reference"><a href="#cite_note-YudellDesalle2000-92"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> as he attempted to bring the study of animal society into the evolutionary fold. This appeared radically new, although Wilson was following Darwin, Fisher, Dawkins and others.<sup id="cite_ref-YudellDesalle2000_92-2" class="reference"><a href="#cite_note-YudellDesalle2000-92"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> Critics such as <a href="/wiki/Gerhard_Lenski" title="Gerhard Lenski">Gerhard Lenski</a> noted that he was following Huxley, Simpson and Dobzhansky's approach, which Lenski considered needlessly reductive as far as human society was concerned.<sup id="cite_ref-Lenski1976_99-0" class="reference"><a href="#cite_note-Lenski1976-99"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> By 2000, the proposed discipline of <a href="/wiki/Sociobiology" title="Sociobiology">sociobiology</a> had morphed into the relatively well-accepted discipline of <a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">evolutionary psychology</a>.<sup id="cite_ref-YudellDesalle2000_92-3" class="reference"><a href="#cite_note-YudellDesalle2000-92"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Lewis's_homeotic_genes,_1978"><span id="Lewis.27s_homeotic_genes.2C_1978"></span>Lewis's homeotic genes, 1978</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=27" title="Edit section: Lewis's homeotic genes, 1978"><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:Genes_hox.jpeg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0c/Genes_hox.jpeg/220px-Genes_hox.jpeg" decoding="async" width="220" height="137" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0c/Genes_hox.jpeg/330px-Genes_hox.jpeg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0c/Genes_hox.jpeg/440px-Genes_hox.jpeg 2x" data-file-width="4131" data-file-height="2571" /></a><figcaption><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a> has formed a synthesis of evolutionary and <a href="/wiki/Developmental_biology" title="Developmental biology">developmental biology</a>, discovering <a href="/wiki/Deep_homology" title="Deep homology">deep homology</a> between the <a href="/wiki/Embryogenesis" class="mw-redirect" title="Embryogenesis">embryogenesis</a> of such different animals as <a href="/wiki/Insect" title="Insect">insects</a> and <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a>.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></div> <p>In 1977, <a href="/wiki/Recombinant_DNA" title="Recombinant DNA">recombinant DNA</a> technology enabled biologists to start to explore the genetic control of development. The growth of <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a> from 1978, when <a href="/wiki/Edward_B._Lewis" title="Edward B. Lewis">Edward B. Lewis</a> discovered <a href="/wiki/Homeotic" class="mw-redirect" title="Homeotic">homeotic</a> genes, showed that many so-called <a href="/wiki/Toolkit_gene" class="mw-redirect" title="Toolkit gene">toolkit genes</a> act to regulate development, influencing the expression of other genes. It also revealed that some of the regulatory genes are extremely ancient, so that animals as different as insects and mammals share control mechanisms; for example, the <i><a href="/wiki/Pax6" class="mw-redirect" title="Pax6">Pax6</a></i> gene is involved in forming the <a href="/wiki/Eye" title="Eye">eyes</a> of mice and of fruit flies. Such <a href="/wiki/Deep_homology" title="Deep homology">deep homology</a> provided strong evidence for evolution and indicated the paths that evolution had taken.<sup id="cite_ref-Gilbert2000_93-1" class="reference"><a href="#cite_note-Gilbert2000-93"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Later_syntheses">Later syntheses</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=28" title="Edit section: Later syntheses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1982, a historical note on a series of evolutionary biology books<sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">[</span>f<span class="cite-bracket">]</span></a></sup> could state without qualification that evolution is the central organizing principle of biology. Smocovitis commented on this that "What the architects of the synthesis had worked to construct had by 1982 become a matter of fact", adding in a footnote that "the centrality of evolution had thus been rendered <a href="/wiki/Tacit_knowledge" title="Tacit knowledge">tacit knowledge</a>, part of the <a href="/wiki/Received_wisdom" class="mw-redirect" title="Received wisdom">received wisdom</a> of the profession".<sup id="cite_ref-FOOTNOTESmocovitis1996186–187_101-0" class="reference"><a href="#cite_note-FOOTNOTESmocovitis1996186–187-101"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> </p><p>By the late 20th century, however, the modern synthesis was showing its age, and fresh syntheses to remedy its defects and fill in its gaps were proposed from different directions. These have included such diverse fields as the <a href="/wiki/Sociology" title="Sociology">study of society</a>,<sup id="cite_ref-YudellDesalle2000_92-4" class="reference"><a href="#cite_note-YudellDesalle2000-92"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> developmental biology,<sup id="cite_ref-Gilbert1991_54-1" class="reference"><a href="#cite_note-Gilbert1991-54"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> epigenetics,<sup id="cite_ref-pigliucci_102-0" class="reference"><a href="#cite_note-pigliucci-102"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a>, <a href="/wiki/Microbiology" title="Microbiology">microbiology</a>, <a href="/wiki/Genomics" title="Genomics">genomics</a>,<sup id="cite_ref-Koonin2009_4-1" class="reference"><a href="#cite_note-Koonin2009-4"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a>, and <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>.<sup id="cite_ref-Noble2011_103-0" class="reference"><a href="#cite_note-Noble2011-103"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> The physiologist <a href="/wiki/Denis_Noble" title="Denis Noble">Denis Noble</a> argues that these additions render neo-Darwinism in the sense of the early 20th century's modern synthesis "at the least, incomplete as a theory of evolution",<sup id="cite_ref-Noble2011_103-1" class="reference"><a href="#cite_note-Noble2011-103"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> and one that has been falsified by later biological research.<sup id="cite_ref-Noble2011_103-2" class="reference"><a href="#cite_note-Noble2011-103"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> </p><p>Michael Rose and Todd Oakley note that evolutionary biology, formerly divided and "<a href="/wiki/Balkanization" title="Balkanization">Balkanized</a>", has been brought together by genomics. It has in their view discarded at least five common assumptions from the modern synthesis, namely that the genome is always a well-organised set of genes; that each gene has a single function; that species are well adapted biochemically to their ecological niches; that species are the durable units of evolution, and all levels from organism to organ, cell and molecule within the species are characteristic of it; and that the design of every organism and cell is efficient. They argue that the "new biology" integrates genomics, <a href="/wiki/Bioinformatics" title="Bioinformatics">bioinformatics</a>, and evolutionary genetics into a general-purpose toolkit for a "Postmodern Synthesis".<sup id="cite_ref-auto1_59-1" class="reference"><a href="#cite_note-auto1-59"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Pigliucci's_extended_evolutionary_synthesis,_2007"><span id="Pigliucci.27s_extended_evolutionary_synthesis.2C_2007"></span>Pigliucci's extended evolutionary synthesis, 2007</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=29" title="Edit section: Pigliucci's extended evolutionary synthesis, 2007"><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/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></div> <p>In 2007, more than half a century after the modern synthesis, <a href="/wiki/Massimo_Pigliucci" title="Massimo Pigliucci">Massimo Pigliucci</a> called for an <a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">extended evolutionary synthesis</a> to incorporate aspects of biology that had not been included or had not existed in the mid-20th century.<sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smee_2015_105-0" class="reference"><a href="#cite_note-Smee_2015-105"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup> It revisits the relative importance of different factors, challenges assumptions made in the modern synthesis, and adds new factors<sup id="cite_ref-Smee_2015_105-1" class="reference"><a href="#cite_note-Smee_2015-105"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-106" class="reference"><a href="#cite_note-106"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup> such as <a href="/wiki/Group_selection#Multilevel_selection_theory" title="Group selection">multilevel selection</a>, <a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">transgenerational epigenetic inheritance</a>, <a href="/wiki/Niche_construction" title="Niche construction">niche construction</a>, and <a href="/wiki/Evolvability" title="Evolvability">evolvability</a>.<sup id="cite_ref-beyonddna_107-0" class="reference"><a href="#cite_note-beyonddna-107"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pigliucci_102-1" class="reference"><a href="#cite_note-pigliucci-102"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Laubichler_2015_108-0" class="reference"><a href="#cite_note-Laubichler_2015-108"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Koonin's_'post-modern'_evolutionary_synthesis,_2009"><span id="Koonin.27s_.27post-modern.27_evolutionary_synthesis.2C_2009"></span>Koonin's 'post-modern' evolutionary synthesis, 2009</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=30" title="Edit section: Koonin's 'post-modern' evolutionary synthesis, 2009"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Tree_Of_Life_(with_horizontal_gene_transfer).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/220px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png" decoding="async" width="220" height="293" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/330px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg/440px-Tree_Of_Life_%28with_horizontal_gene_transfer%29.svg.png 2x" data-file-width="1782" data-file-height="2370" /></a><figcaption>A 21st century <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">tree of life</a> showing <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfers</a> among <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a> and the saltational <a href="/wiki/Endosymbiosis" class="mw-redirect" title="Endosymbiosis">endosymbiosis</a> events that created the <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>, neither fitting into the 20th century's modern synthesis</figcaption></figure> <p>In 2009, Darwin's 200th anniversary, the <i>Origin of Species</i><span class="nowrap" style="padding-left:0.1em;">'</span> 150th, and the 200th of Lamarck's "early evolutionary synthesis",<sup id="cite_ref-Koonin2009_4-2" class="reference"><a href="#cite_note-Koonin2009-4"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <i><a href="/wiki/Philosophie_Zoologique" class="mw-redirect" title="Philosophie Zoologique">Philosophie Zoologique</a></i>, the evolutionary biologist <a href="/wiki/Eugene_Koonin" title="Eugene Koonin">Eugene Koonin</a> stated that while "the edifice of the [early 20th century] Modern Synthesis has crumbled, apparently, beyond repair",<sup id="cite_ref-Koonin2009_4-3" class="reference"><a href="#cite_note-Koonin2009-4"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> a new 21st-century synthesis could be glimpsed. Three interlocking revolutions had, he argued, taken place in evolutionary biology: molecular, microbiological, and genomic. The <a href="/wiki/Molecular_biology" title="Molecular biology">molecular revolution</a> included the <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral theory</a>, that most mutations are neutral and that <a href="/wiki/Negative_selection_(natural_selection)" title="Negative selection (natural selection)">negative selection</a> happens more often than the <a href="/wiki/Directional_selection" title="Directional selection">positive</a> form, and that all current life evolved from <a href="/wiki/Last_Universal_Common_Ancestor" class="mw-redirect" title="Last Universal Common Ancestor">a single common ancestor</a>. In microbiology, the synthesis has expanded to cover the <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>, using <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNA</a> to form a <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">tree of life</a>. Finally, <a href="/wiki/Genomics" title="Genomics">genomics</a> brought together the molecular and microbiological syntheses - in particular, <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a> between <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> shows that prokaryotes can freely share genes. Many of these points had already been made by other researchers such as Ulrich Kutschera and <a href="/wiki/Karl_J._Niklas" title="Karl J. Niklas">Karl J. Niklas</a>.<sup id="cite_ref-Kutschera_Niklas_2004_109-0" class="reference"><a href="#cite_note-Kutschera_Niklas_2004-109"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Towards_a_replacement_synthesis">Towards a replacement synthesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=31" title="Edit section: Towards a replacement synthesis"><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:Modern_Synthesis_Limits.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Modern_Synthesis_Limits.svg/480px-Modern_Synthesis_Limits.svg.png" decoding="async" width="480" height="296" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Modern_Synthesis_Limits.svg/720px-Modern_Synthesis_Limits.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Modern_Synthesis_Limits.svg/960px-Modern_Synthesis_Limits.svg.png 2x" data-file-width="1027" data-file-height="633" /></a><figcaption>Inputs to the modern synthesis, with other topics (inverted colours) such as developmental biology that were not joined with evolutionary biology until the turn of the 21st century<sup id="cite_ref-Kutschera_Niklas_2004_109-1" class="reference"><a href="#cite_note-Kutschera_Niklas_2004-109"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Biologists, alongside scholars of the history and philosophy of biology, have continued to debate the need for, and possible nature of, a replacement synthesis. For example, in 2017 Philippe Huneman and Denis M. Walsh stated in their book <i>Challenging the Modern Synthesis</i> that numerous theorists had pointed out that the disciplines of embryological developmental theory, morphology, and ecology had been omitted. They noted that all such arguments amounted to a continuing desire to replace the modern synthesis with one that united "all biological fields of research related to evolution, adaptation, and diversity in a single theoretical framework."<sup id="cite_ref-110" class="reference"><a href="#cite_note-110"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> They observed further that there are two groups of challenges to the way the modern synthesis viewed inheritance. The first is that other modes such as <a href="/wiki/Epigenetic_inheritance" class="mw-redirect" title="Epigenetic inheritance">epigenetic inheritance</a>, <a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">phenotypic plasticity</a>, the <a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a>, and the <a href="/wiki/Maternal_effect" title="Maternal effect">maternal effect</a> allow new characteristics to arise and be passed on and for the genes to catch up with the new adaptations later. The second is that all such mechanisms are part, not of an inheritance system, but a <a href="/wiki/Developmental_systems_theory" title="Developmental systems theory">developmental system</a>: the fundamental unit is not a discrete selfishly competing gene, but a collaborating system that works at all levels from genes and cells to organisms and cultures to guide evolution.<sup id="cite_ref-111" class="reference"><a href="#cite_note-111"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> The molecular biologist <a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Sean B. Carroll</a> has commented that had Huxley had access to <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a>, "embryology would have been a cornerstone of his Modern Synthesis, and so evo-devo is today a key element of a more complete, expanded evolutionary synthesis."<sup id="cite_ref-Carroll_2008_112-0" class="reference"><a href="#cite_note-Carroll_2008-112"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Historiography">Historiography</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=32" title="Edit section: Historiography"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div><p> Looking back at the conflicting accounts of the modern synthesis, the historian Betty Smocovitis notes in her 1996 book <i>Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology</i> that both historians and philosophers of biology have attempted to grasp its scientific meaning, but have found it "a moving target";<sup id="cite_ref-Smocovitis187_113-0" class="reference"><a href="#cite_note-Smocovitis187-113"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> the only thing they agreed on was that it was a historical event.<sup id="cite_ref-Smocovitis187_113-1" class="reference"><a href="#cite_note-Smocovitis187-113"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> In her words</p><blockquote><p>"by the late 1980s the notoriety of the evolutionary synthesis was recognized ... So notorious did 'the synthesis' become, that few serious historically minded analysts would touch the subject, let alone know where to begin to sort through the interpretive mess left behind by the numerous critics and commentators".<sup id="cite_ref-114" class="reference"><a href="#cite_note-114"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup></p></blockquote> <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=Modern_synthesis_(20th_century)&action=edit&section=33" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Objections_to_evolution" title="Objections to evolution">Objections to evolution</a></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=Modern_synthesis_(20th_century)&action=edit&section=34" 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-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">Also known variously as the <b>New Synthesis</b>, the <b>Modern Evolutionary Synthesis</b>, the <b>Evolutionary Synthesis</b>, and the <b>neo-Darwinian Synthesis</b>. These alternative terms are ambiguous as they could possibly include later syntheses, so this article uses <a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a>'s 1942 "modern synthesis"<sup id="cite_ref-FOOTNOTEHuxley2010_2-0" class="reference"><a href="#cite_note-FOOTNOTEHuxley2010-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> throughout.</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">Peter Gauthier has however argued that <a href="/wiki/Weismann%27s_experiment" class="mw-redirect" title="Weismann's experiment">Weismann's experiment</a> showed only that injury did not affect the germplasm. It did not test the effect of Lamarckian use and disuse.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-37"><span class="mw-cite-backlink"><b><a href="#cite_ref-37">^</a></b></span> <span class="reference-text">Morgan's work with fruit flies helped establish the link between Mendelian genetics and the <a href="/wiki/Chromosome" title="Chromosome">chromosomal</a> theory of inheritance, that the hereditary material was embodied in these bodies within the cell nucleus.<sup id="cite_ref-auto_36-0" class="reference"><a href="#cite_note-auto-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-46"><span class="mw-cite-backlink"><b><a href="#cite_ref-46">^</a></b></span> <span class="reference-text">Fisher also analysed <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a> in his book, but his work was largely ignored, and Darwin's case for such selection misunderstood, so it formed no substantial part of the modern synthesis.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-57"><span class="mw-cite-backlink"><b><a href="#cite_ref-57">^</a></b></span> <span class="reference-text">Though <a href="/wiki/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a> had called for <a href="/wiki/Embryology" title="Embryology">embryology</a> to be added to the synthesis in his 1953 paper "Epigenetics and Evolution".<sup id="cite_ref-Smocovitis153_56-0" class="reference"><a href="#cite_note-Smocovitis153-56"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-100"><span class="mw-cite-backlink"><b><a href="#cite_ref-100">^</a></b></span> <span class="reference-text">In a reissue of Dobzhansky's <i>Genetics and the Origin of Species</i>.</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=Modern_synthesis_(20th_century)&action=edit&section=35" 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"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Gould216-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Gould216_1-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFGould2002">Gould 2002</a>, p. 216</span> </li> <li id="cite_note-FOOTNOTEHuxley2010-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-FOOTNOTEHuxley2010_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-FOOTNOTEHuxley2010_2-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFHuxley2010">Huxley 2010</a>.</span> </li> <li id="cite_note-Koonin2009-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Koonin2009_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Koonin2009_4-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Koonin2009_4-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Koonin2009_4-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFKoonin2009" class="citation journal cs1"><a href="/wiki/Eugene_Koonin" title="Eugene Koonin">Koonin, Eugene V.</a> (November 2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2784144">"The Origin at 150: is a new evolutionary synthesis in sight?"</a>. <i><a href="/wiki/Trends_in_Genetics" class="mw-redirect" title="Trends in Genetics">Trends in Genetics</a></i>. <b>25</b> (11): 473–475. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.tig.2009.09.007">10.1016/j.tig.2009.09.007</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2784144">2784144</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/19836100">19836100</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+Genetics&rft.atitle=The+Origin+at+150%3A+is+a+new+evolutionary+synthesis+in+sight%3F&rft.volume=25&rft.issue=11&rft.pages=473-475&rft.date=2009-11&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2784144%23id-name%3DPMC&rft_id=info%3Apmid%2F19836100&rft_id=info%3Adoi%2F10.1016%2Fj.tig.2009.09.007&rft.aulast=Koonin&rft.aufirst=Eugene+V.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2784144&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><a href="#CITEREFBowler2003">Bowler 2003</a>, pp. 236–256</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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKutschera2003" class="citation journal cs1"><a href="/wiki/Ulrich_Kutschera" title="Ulrich Kutschera">Kutschera, Ulrich</a> (December 2003). "A comparative analysis of the Darwin–Wallace papers and the development of the concept of natural selection". <i>Theory in Biosciences</i>. <b>122</b> (4): 343–359. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs12064-003-0063-6">10.1007/s12064-003-0063-6</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:24297627">24297627</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Theory+in+Biosciences&rft.atitle=A+comparative+analysis+of+the+Darwin%E2%80%93Wallace+papers+and+the+development+of+the+concept+of+natural+selection&rft.volume=122&rft.issue=4&rft.pages=343-359&rft.date=2003-12&rft_id=info%3Adoi%2F10.1007%2Fs12064-003-0063-6&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A24297627%23id-name%3DS2CID&rft.aulast=Kutschera&rft.aufirst=Ulrich&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFButler1880" class="citation book cs1"><a href="/wiki/Samuel_Butler_(novelist)" title="Samuel Butler (novelist)">Butler, Samuel</a> (1880). <a rel="nofollow" class="external text" href="https://archive.org/details/unconsciousmemo00hartgoog"><i>Unconscious Memory</i></a>. David Bogue. p. 280. <q>I may predict with some certainty that before long we shall find the original Darwinism of Dr. <a href="/wiki/Erasmus_Darwin" title="Erasmus Darwin">Erasmus Darwin</a> … generally accepted instead of the neo-Darwinism of to-day, and that the variations whose accumulation results in species will be recognised as due to the wants and endeavours of the living forms in which they appear, instead of being ascribed to chance, or, in other words, to unknown causes, as by Mr. Charles Darwin's system</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Unconscious+Memory&rft.pages=280&rft.pub=David+Bogue&rft.date=1880&rft.aulast=Butler&rft.aufirst=Samuel&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Funconsciousmemo00hartgoog&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></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 id="CITEREFBeccaloni2013" class="citation web cs1">Beccaloni, George (2013). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20190411231439/http://wallacefund.info/terms-darwinism-and-neo-darwinism">"On the Terms "Darwinism" and "Neo-Darwinism"<span class="cs1-kern-right"></span>"</a>. <i>A. R. Wallace Website</i>. Archived from <a rel="nofollow" class="external text" href="http://wallacefund.info/terms-darwinism-and-neo-darwinism">the original</a> on 2019-04-11<span class="reference-accessdate">. Retrieved <span class="nowrap">2017-09-14</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=A.+R.+Wallace+Website&rft.atitle=On+the+Terms+%22Darwinism%22+and+%22Neo-Darwinism%22&rft.date=2013&rft.aulast=Beccaloni&rft.aufirst=George&rft_id=http%3A%2F%2Fwallacefund.info%2Fterms-darwinism-and-neo-darwinism&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></span> </li> <li id="cite_note-Bowler196-250-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-Bowler196-250_9-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFBowler2003">Bowler 2003</a>, pp. 196–253</span> </li> <li id="cite_note-Larson105-129-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-Larson105-129_10-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFLarson2004">Larson 2004</a>, pp. 105–129</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"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGayon,_Jean1998" class="citation book cs1">Gayon, Jean (1998). <i>Darwinism's Struggle for Survival: Heredity and the Hypothesis of Natural Selection</i>. 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John Murray. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4191-8660-8" title="Special:BookSources/978-1-4191-8660-8"><bdi>978-1-4191-8660-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+variation+of+animals+and+plants+under+domestication&rft.pub=John+Murray&rft.date=1868&rft.isbn=978-1-4191-8660-8&rft.aulast=Darwin&rft.aufirst=Charles&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DF880.1%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" 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 id="CITEREFHolterhoff,_Kate2014" class="citation journal cs1">Holterhoff, Kate (2014). 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Retrieved <span class="nowrap">14 December</span> 2017</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Mendel%27s+Peas&rft.place=Norwich%2C+UK&rft.pub=Germplasm+Resources+Unit%2C+John+Innes+Centre&rft.aulast=Ambrose&rft.aufirst=Mike&rft_id=https%3A%2F%2Fwww.jic.ac.uk%2Fgermplas%2FPISUM%2FZGS4F.HTM&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></span> </li> <li id="cite_note-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-21">^</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://www.phschool.com/science/biology_place/biocoach/inheritance/laws.html">"Reviewing Mendel's Laws"</a>. Pearson<span class="reference-accessdate">. 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"The modern theory of biological evolution: an expanded synthesis". <i><a href="/wiki/Naturwissenschaften" class="mw-redirect" title="Naturwissenschaften">Naturwissenschaften</a></i>. <b>91</b> (6): 255–276. <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/2004NW.....91..255K">2004NW.....91..255K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00114-004-0515-y">10.1007/s00114-004-0515-y</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15241603">15241603</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:10731711">10731711</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Naturwissenschaften&rft.atitle=The+modern+theory+of+biological+evolution%3A+an+expanded+synthesis&rft.volume=91&rft.issue=6&rft.pages=255-276&rft.date=2004-06&rft_id=info%3Adoi%2F10.1007%2Fs00114-004-0515-y&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A10731711%23id-name%3DS2CID&rft_id=info%3Apmid%2F15241603&rft_id=info%3Abibcode%2F2004NW.....91..255K&rft.aulast=Kutschera&rft.aufirst=Ulrich&rft.au=Niklas%2C+Karl+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></span> </li> <li id="cite_note-110"><span class="mw-cite-backlink"><b><a href="#cite_ref-110">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHunemanWalsh2017" class="citation book cs1">Huneman, Philippe; Walsh, Denis M. (2017). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=72EwDwAAQBAJ&pg=PT76"><i>Challenging the Modern Synthesis: Adaptation, Development, and Inheritance</i></a>. <a href="/wiki/Oxford_University_Press" title="Oxford University Press">Oxford University Press</a>. pp. 76 (Chapter 2). <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-068145-6" title="Special:BookSources/978-0-19-068145-6"><bdi>978-0-19-068145-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Challenging+the+Modern+Synthesis%3A+Adaptation%2C+Development%2C+and+Inheritance&rft.pages=76+%28Chapter+2%29&rft.pub=Oxford+University+Press&rft.date=2017&rft.isbn=978-0-19-068145-6&rft.aulast=Huneman&rft.aufirst=Philippe&rft.au=Walsh%2C+Denis+M.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D72EwDwAAQBAJ%26pg%3DPT76&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></span> </li> <li id="cite_note-111"><span class="mw-cite-backlink"><b><a href="#cite_ref-111">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHunemanWalsh2017" class="citation book cs1">Huneman, Philippe; Walsh, Denis M. (2017). <i>Challenging the Modern Synthesis: Adaptation, Development, and Inheritance</i>. <a href="/wiki/Oxford_University_Press" title="Oxford University Press">Oxford University Press</a>. pp. Introduction. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-068145-6" title="Special:BookSources/978-0-19-068145-6"><bdi>978-0-19-068145-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Challenging+the+Modern+Synthesis%3A+Adaptation%2C+Development%2C+and+Inheritance&rft.pages=Introduction&rft.pub=Oxford+University+Press&rft.date=2017&rft.isbn=978-0-19-068145-6&rft.aulast=Huneman&rft.aufirst=Philippe&rft.au=Walsh%2C+Denis+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></span> </li> <li id="cite_note-Carroll_2008-112"><span class="mw-cite-backlink"><b><a href="#cite_ref-Carroll_2008_112-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarroll2008" class="citation journal cs1"><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Carroll, Sean B.</a> (2008). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cell.2008.06.030">"Evo-Devo and an Expanding Evolutionary Synthesis: A Genetic Theory of Morphological Evolution"</a>. <i>Cell</i>. <b>134</b> (1): 25–36. <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.cell.2008.06.030">10.1016/j.cell.2008.06.030</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/18614008">18614008</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:2513041">2513041</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell&rft.atitle=Evo-Devo+and+an+Expanding+Evolutionary+Synthesis%3A+A+Genetic+Theory+of+Morphological+Evolution&rft.volume=134&rft.issue=1&rft.pages=25-36&rft.date=2008&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A2513041%23id-name%3DS2CID&rft_id=info%3Apmid%2F18614008&rft_id=info%3Adoi%2F10.1016%2Fj.cell.2008.06.030&rft.aulast=Carroll&rft.aufirst=Sean+B.&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.cell.2008.06.030&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></span> </li> <li id="cite_note-Smocovitis187-113"><span class="mw-cite-backlink">^ <a href="#cite_ref-Smocovitis187_113-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Smocovitis187_113-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFSmocovitis1996">Smocovitis 1996</a>, p. 187</span> </li> <li id="cite_note-114"><span class="mw-cite-backlink"><b><a href="#cite_ref-114">^</a></b></span> <span class="reference-text"><a href="#CITEREFSmocovitis1996">Smocovitis 1996</a>, p. 43</span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Sources">Sources</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=36" title="Edit section: Sources"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBateson1894" class="citation book cs1"><a href="/wiki/William_Bateson" title="William Bateson">Bateson, William</a> (1894). <a rel="nofollow" class="external text" href="https://archive.org/details/materialsforstud00bate"><i>Materials for the Study of Variation Treated with Especial Regard to Discontinuity in the Origin of Species</i></a>. London; New York: <a href="/wiki/Macmillan_Publishers" title="Macmillan Publishers">Macmillan</a>. <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/07001821">07001821</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/1944285">1944285</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Materials+for+the+Study+of+Variation+Treated+with+Especial+Regard+to+Discontinuity+in+the+Origin+of+Species&rft.place=London%3B+New+York&rft.pub=Macmillan&rft.date=1894&rft_id=info%3Aoclcnum%2F1944285&rft_id=info%3Alccn%2F07001821&rft.aulast=Bateson&rft.aufirst=William&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmaterialsforstud00bate&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBowler2003" class="citation book cs1"><a href="/wiki/Peter_J._Bowler" title="Peter J. Bowler">Bowler, Peter J.</a> (2003). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/evolutionhistory0000bowl_n7y8"><i>Evolution: The History of an Idea</i></a></span> (3rd completely rev. and expanded ed.). Berkeley, CA: <a href="/wiki/University_of_California_Press" title="University of California Press">University of California Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-520-23693-6" title="Special:BookSources/978-0-520-23693-6"><bdi>978-0-520-23693-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution%3A+The+History+of+an+Idea&rft.place=Berkeley%2C+CA&rft.edition=3rd+completely+rev.+and+expanded&rft.pub=University+of+California+Press&rft.date=2003&rft.isbn=978-0-520-23693-6&rft.aulast=Bowler&rft.aufirst=Peter+J.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fevolutionhistory0000bowl_n7y8&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBowler1989" class="citation book cs1"><a href="/wiki/Peter_J._Bowler" title="Peter J. Bowler">Bowler, Peter J.</a> (1989). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/evolutionhistory0000bowl"><i>Evolution: The History of an Idea</i></a></span> (2nd revised ed.). Berkeley, CA: <a href="/wiki/University_of_California_Press" title="University of California Press">University of California Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-520-06386-0" title="Special:BookSources/978-0-520-06386-0"><bdi>978-0-520-06386-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution%3A+The+History+of+an+Idea&rft.place=Berkeley%2C+CA&rft.edition=2nd+revised&rft.pub=University+of+California+Press&rft.date=1989&rft.isbn=978-0-520-06386-0&rft.aulast=Bowler&rft.aufirst=Peter+J.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fevolutionhistory0000bowl&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li></ul></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDobzhansky1937" class="citation book cs1"><a href="/wiki/Theodosius_Dobzhansky" title="Theodosius Dobzhansky">Dobzhansky, Theodosius</a> (1937). <a href="/wiki/Genetics_and_the_Origin_of_Species" title="Genetics and the Origin of Species"><i>Genetics and the Origin of Species</i></a>. <a href="/wiki/Columbia_University_Biological_Series" title="Columbia University Biological Series">Columbia University Biological Series</a>. New York: <a href="/wiki/Columbia_University_Press" title="Columbia University Press">Columbia University Press</a>. <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/37033383">37033383</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/766405">766405</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Genetics+and+the+Origin+of+Species&rft.place=New+York&rft.series=Columbia+University+Biological+Series&rft.pub=Columbia+University+Press&rft.date=1937&rft_id=info%3Aoclcnum%2F766405&rft_id=info%3Alccn%2F37033383&rft.aulast=Dobzhansky&rft.aufirst=Theodosius&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDobzhansky1951" class="citation book cs1">—— (1951). <a rel="nofollow" class="external text" href="https://archive.org/details/in.ernet.dli.2015.547790"><i>Genetics and the Origin of Species</i></a>. Columbia University Biological Series (3rd revised ed.). New York: Columbia University Press. <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/51014816">51014816</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/295774">295774</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Genetics+and+the+Origin+of+Species&rft.place=New+York&rft.series=Columbia+University+Biological+Series&rft.edition=3rd+revised&rft.pub=Columbia+University+Press&rft.date=1951&rft_id=info%3Aoclcnum%2F295774&rft_id=info%3Alccn%2F51014816&rft.aulast=Dobzhansky&rft.aufirst=Theodosius&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fin.ernet.dli.2015.547790&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li></ul></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFisher1999" class="citation book cs1"><a href="/wiki/Ronald_Fisher" title="Ronald Fisher">Fisher, R. A.</a> (1999) [1930]. <a href="/wiki/The_Genetical_Theory_of_Natural_Selection" title="The Genetical Theory of Natural Selection"><i>The Genetical Theory of Natural Selection</i></a>. Edited with a foreword and notes by J. H. Bennett (A complete variorum ed.). Oxford, UK: Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-850440-5" title="Special:BookSources/978-0-19-850440-5"><bdi>978-0-19-850440-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Genetical+Theory+of+Natural+Selection&rft.place=Oxford%2C+UK&rft.edition=A+complete+variorum&rft.pub=Oxford+University+Press&rft.date=1999&rft.isbn=978-0-19-850440-5&rft.aulast=Fisher&rft.aufirst=R.+A.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFord1964" class="citation journal cs1"><a href="/wiki/E._B._Ford" title="E. B. Ford">Ford, E. B.</a> (1964). "Ecological Genetics". <i>Advancement of Science</i>. <b>25</b> (124). London; New York: <a href="/wiki/Methuen_Publishing" title="Methuen Publishing">Methuen Publishing</a>; <a href="/wiki/John_Wiley_%26_Sons" class="mw-redirect" title="John Wiley & Sons">John Wiley & Sons</a>: 227–35. <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/64009038">64009038</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/875507052">875507052</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/5701915">5701915</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Advancement+of+Science&rft.atitle=Ecological+Genetics&rft.volume=25&rft.issue=124&rft.pages=227-35&rft.date=1964&rft_id=info%3Aoclcnum%2F875507052&rft_id=info%3Apmid%2F5701915&rft_id=info%3Alccn%2F64009038&rft.aulast=Ford&rft.aufirst=E.+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFord1975" class="citation journal cs1">—— (1975). "Ecological Genetics". <i>Advancement of Science</i>. <b>25</b> (124) (4th ed.). London; New York: <a href="/wiki/Chapman_%26_Hall" title="Chapman & Hall">Chapman and Hall</a>; John Wiley & Sons: 227–35. <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/75002165">75002165</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/1890603">1890603</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/5701915">5701915</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Advancement+of+Science&rft.atitle=Ecological+Genetics&rft.volume=25&rft.issue=124&rft.pages=227-35&rft.date=1975&rft_id=info%3Aoclcnum%2F1890603&rft_id=info%3Apmid%2F5701915&rft_id=info%3Alccn%2F75002165&rft.aulast=Ford&rft.aufirst=E.+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li></ul></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGould1977" class="citation book cs1"><a href="/wiki/Stephen_Jay_Gould" title="Stephen Jay Gould">Gould, Stephen Jay</a> (1977). <a href="/wiki/Ontogeny_and_Phylogeny_(book)" class="mw-redirect" title="Ontogeny and Phylogeny (book)"><i>Ontogeny and Phylogeny</i></a>. <a href="/wiki/Harvard_University_Press" title="Harvard University Press">Belknap Press of Harvard University Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-674-63940-9" title="Special:BookSources/978-0-674-63940-9"><bdi>978-0-674-63940-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Ontogeny+and+Phylogeny&rft.pub=Belknap+Press+of+Harvard+University+Press&rft.date=1977&rft.isbn=978-0-674-63940-9&rft.aulast=Gould&rft.aufirst=Stephen+Jay&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGould2002" class="citation book cs1">Gould, Stephen Jay (2002). <a href="/wiki/The_Structure_of_Evolutionary_Theory" title="The Structure of Evolutionary Theory"><i>The Structure of Evolutionary Theory</i></a>. Belknap Press of Harvard University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-674-00613-3" title="Special:BookSources/978-0-674-00613-3"><bdi>978-0-674-00613-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Structure+of+Evolutionary+Theory&rft.pub=Belknap+Press+of+Harvard+University+Press&rft.date=2002&rft.isbn=978-0-674-00613-3&rft.aulast=Gould&rft.aufirst=Stephen+Jay&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGrafenRidley2006" class="citation book cs1"><a href="/wiki/Alan_Grafen" title="Alan Grafen">Grafen, Alan</a>; <a href="/wiki/Mark_Ridley_(zoologist)" title="Mark Ridley (zoologist)">Ridley, Mark</a>, eds. (2006). <a href="/wiki/Richard_Dawkins:_How_a_Scientist_Changed_the_Way_We_Think" title="Richard Dawkins: How a Scientist Changed the Way We Think"><i>Richard Dawkins: How a Scientist Changed the Way We Think</i></a>. Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-929116-8" title="Special:BookSources/978-0-19-929116-8"><bdi>978-0-19-929116-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Richard+Dawkins%3A+How+a+Scientist+Changed+the+Way+We+Think&rft.pub=Oxford+University+Press&rft.date=2006&rft.isbn=978-0-19-929116-8&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuxley2010" class="citation book cs1"><a href="/wiki/Julian_Huxley" title="Julian Huxley">Huxley, Julian</a> (2010) [1942]. <a href="/wiki/Evolution:_The_Modern_Synthesis" title="Evolution: The Modern Synthesis"><i>Evolution: The Modern Synthesis</i></a>. With a new foreword by <a href="/wiki/Massimo_Pigliucci" title="Massimo Pigliucci">Massimo Pigliucci</a> and <a href="/wiki/Gerd_M%C3%BCller_(theoretical_biologist)" class="mw-redirect" title="Gerd Müller (theoretical biologist)">Gerd B. Müller</a> (Definitive ed.). Cambridge, MA: <a href="/wiki/MIT_Press" title="MIT Press">MIT Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-262-51366-1" title="Special:BookSources/978-0-262-51366-1"><bdi>978-0-262-51366-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution%3A+The+Modern+Synthesis&rft.place=Cambridge%2C+MA&rft.edition=Definitive&rft.pub=MIT+Press&rft.date=2010&rft.isbn=978-0-262-51366-1&rft.aulast=Huxley&rft.aufirst=Julian&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLarson2004" class="citation book cs1"><a href="/wiki/Edward_J._Larson" title="Edward J. Larson">Larson, Edward J.</a> (2004). <a rel="nofollow" class="external text" href="https://archive.org/details/evolutionremarka00lars"><i>Evolution: The Remarkable History of a Scientific Theory</i></a>. Modern Library Chronicles. Vol. 17. New York: <a href="/wiki/Modern_Library" title="Modern Library">Modern Library</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-679-64288-6" title="Special:BookSources/978-0-679-64288-6"><bdi>978-0-679-64288-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution%3A+The+Remarkable+History+of+a+Scientific+Theory&rft.place=New+York&rft.series=Modern+Library+Chronicles&rft.pub=Modern+Library&rft.date=2004&rft.isbn=978-0-679-64288-6&rft.aulast=Larson&rft.aufirst=Edward+J.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fevolutionremarka00lars&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMayr1982" class="citation book cs1"><a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Mayr, Ernst</a> (1982). <a rel="nofollow" class="external text" href="https://archive.org/details/growthofbiologic00mayr"><i>The Growth of Biological Thought: Diversity, Evolution, and Inheritance</i></a>. Cambridge, MA: Belknap Press of Harvard University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-674-36445-5" title="Special:BookSources/978-0-674-36445-5"><bdi>978-0-674-36445-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Growth+of+Biological+Thought%3A+Diversity%2C+Evolution%2C+and+Inheritance&rft.place=Cambridge%2C+MA&rft.pub=Belknap+Press+of+Harvard+University+Press&rft.date=1982&rft.isbn=978-0-674-36445-5&rft.aulast=Mayr&rft.aufirst=Ernst&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fgrowthofbiologic00mayr&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMayr1999" class="citation book cs1">Mayr, Ernst (1999) [1942]. <a href="/wiki/Systematics_and_the_Origin_of_Species" title="Systematics and the Origin of Species"><i>Systematics and the Origin of Species from the Viewpoint of a Zoologist</i></a> (1st Harvard University Press pbk. ed.). <a href="/wiki/Harvard_University_Press" title="Harvard University Press">Harvard University Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-674-86250-0" title="Special:BookSources/978-0-674-86250-0"><bdi>978-0-674-86250-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Systematics+and+the+Origin+of+Species+from+the+Viewpoint+of+a+Zoologist&rft.edition=1st+Harvard+University+Press+pbk.&rft.pub=Harvard+University+Press&rft.date=1999&rft.isbn=978-0-674-86250-0&rft.aulast=Mayr&rft.aufirst=Ernst&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMayrProvine1998" class="citation book cs1">Mayr, Ernst; <a href="/wiki/Will_Provine" class="mw-redirect" title="Will Provine">Provine, William B.</a>, eds. (1998) [1980]. <i>The Evolutionary Synthesis: Perspectives on the Unification of Biology</i>. With a new preface by Ernst Mayr (1st paperback ed.). Cambridge, MA: Harvard University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-674-27226-2" title="Special:BookSources/978-0-674-27226-2"><bdi>978-0-674-27226-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Evolutionary+Synthesis%3A+Perspectives+on+the+Unification+of+Biology&rft.place=Cambridge%2C+MA&rft.edition=1st+paperback&rft.pub=Harvard+University+Press&rft.date=1998&rft.isbn=978-0-674-27226-2&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFProvine2001" class="citation book cs1">Provine, W. B. (2001). <i>The Origins of Theoretical Population Genetics, with a new afterword</i>. University of Chicago Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-68464-2" title="Special:BookSources/978-0-226-68464-2"><bdi>978-0-226-68464-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Origins+of+Theoretical+Population+Genetics%2C+with+a+new+afterword&rft.pub=University+of+Chicago+Press&rft.date=2001&rft.isbn=978-0-226-68464-2&rft.aulast=Provine&rft.aufirst=W.+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRensch1947" class="citation book cs1 cs1-prop-foreign-lang-source"><a href="/wiki/Bernhard_Rensch" title="Bernhard Rensch">Rensch, Bernhard</a> (1947). <i>Neuere Probleme der Abstammungslehre. Die transspezifische Evolution</i> [<i>Newer Problems of Evolutionary Theory: The trans-specific Evolution</i>] (in German). Stuttgart: Ferdinand Enke Verlag. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/2271422">2271422</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Neuere+Probleme+der+Abstammungslehre.+Die+transspezifische+Evolution&rft.place=Stuttgart&rft.pub=Ferdinand+Enke+Verlag&rft.date=1947&rft_id=info%3Aoclcnum%2F2271422&rft.aulast=Rensch&rft.aufirst=Bernhard&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRensch1959" class="citation journal cs1">—— (1959). "Evolution Above the Species Level". <i>American Journal of Physical Anthropology</i> (English translation of 2nd edition of <i>Neuere Probleme der Abstammungslehre</i> (1954)). Columbia Biological Series. <b>19</b> (3): 408–410. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fajpa.1330200329">10.1002/ajpa.1330200329</a>. <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/60002460">60002460</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/3677530">3677530</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Journal+of+Physical+Anthropology&rft.atitle=Evolution+Above+the+Species+Level&rft.volume=19&rft.issue=3&rft.pages=408-410&rft.date=1959&rft_id=info%3Aoclcnum%2F3677530&rft_id=info%3Alccn%2F60002460&rft_id=info%3Adoi%2F10.1002%2Fajpa.1330200329&rft.aulast=Rensch&rft.aufirst=Bernhard&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li></ul></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRuse1996" class="citation book cs1"><a href="/wiki/Michael_Ruse" title="Michael Ruse">Ruse, Michael</a> (1996). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/monadtomanconcep0000ruse"><i>Monad to Man: the Concept of Progress in Evolutionary Biology</i></a></span>. Harvard University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-674-03248-4" title="Special:BookSources/978-0-674-03248-4"><bdi>978-0-674-03248-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Monad+to+Man%3A+the+Concept+of+Progress+in+Evolutionary+Biology&rft.pub=Harvard+University+Press&rft.date=1996&rft.isbn=978-0-674-03248-4&rft.aulast=Ruse&rft.aufirst=Michael&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmonadtomanconcep0000ruse&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSmocovitis1996" class="citation journal cs1">Smocovitis, Vassiliki Betty (1996). "Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology". <i>Journal of the History of Biology</i>. <b>25</b> (1). Princeton, NJ: Princeton University Press: 1–65. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fbf01947504">10.1007/bf01947504</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-691-03343-3" title="Special:BookSources/978-0-691-03343-3"><bdi>978-0-691-03343-3</bdi></a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11623198">11623198</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:189833728">189833728</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+History+of+Biology&rft.atitle=Unifying+Biology%3A+The+Evolutionary+Synthesis+and+Evolutionary+Biology&rft.volume=25&rft.issue=1&rft.pages=1-65&rft.date=1996&rft_id=info%3Adoi%2F10.1007%2Fbf01947504&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A189833728%23id-name%3DS2CID&rft_id=info%3Apmid%2F11623198&rft.isbn=978-0-691-03343-3&rft.aulast=Smocovitis&rft.aufirst=Vassiliki+Betty&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWright1932" class="citation book cs1"><a href="/wiki/Sewall_Wright" title="Sewall Wright">Wright, Sewall</a> (1932). "The Roles of Mutation, Inbreeding, Crossbreeding and Selection in Evolution". In <a href="/wiki/Donald_F._Jones" title="Donald F. Jones">Jones, Donald F.</a> (ed.). <i>Proceedings of the Sixth International Congress of Genetics</i>. Vol. 1. Ithaca, NY: <a href="/wiki/Genetics_Society_of_America" title="Genetics Society of America">Genetics Society of America</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/439596433">439596433</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=The+Roles+of+Mutation%2C+Inbreeding%2C+Crossbreeding+and+Selection+in+Evolution&rft.btitle=Proceedings+of+the+Sixth+International+Congress+of+Genetics&rft.place=Ithaca%2C+NY&rft.pub=Genetics+Society+of+America&rft.date=1932&rft_id=info%3Aoclcnum%2F439596433&rft.aulast=Wright&rft.aufirst=Sewall&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Modern_synthesis_(20th_century)&action=edit&section=37" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1"><a href="/wiki/Garland_E._Allen" title="Garland E. Allen">Allen, Garland E.</a> (1978). <i>Thomas Hunt Morgan: The Man and His Science</i>. Princeton, NJ: <a href="/wiki/Princeton_University_Press" title="Princeton University Press">Princeton University Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-691-08200-4" title="Special:BookSources/978-0-691-08200-4"><bdi>978-0-691-08200-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Thomas+Hunt+Morgan%3A+The+Man+and+His+Science&rft.place=Princeton%2C+NJ&rft.pub=Princeton+University+Press&rft.date=1978&rft.isbn=978-0-691-08200-4&rft.aulast=Allen&rft.aufirst=Garland+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1"><a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Dawkins, Richard</a> (1990). <a href="/wiki/The_Blind_Watchmaker" title="The Blind Watchmaker"><i>The Blind Watchmaker</i></a>. Penguin Science. London: <a href="/wiki/Penguin_Books" title="Penguin Books">Penguin Books</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-14-014481-9" title="Special:BookSources/978-0-14-014481-9"><bdi>978-0-14-014481-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Blind+Watchmaker&rft.place=London&rft.series=Penguin+Science&rft.pub=Penguin+Books&rft.date=1990&rft.isbn=978-0-14-014481-9&rft.aulast=Dawkins&rft.aufirst=Richard&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1"><a href="/wiki/Douglas_J._Futuyma" title="Douglas J. Futuyma">Futuyma, Douglas J.</a> (1998). <i>Evolutionary Biology</i> (3rd ed.). Sunderland, MA: <a href="/wiki/Sinauer_Associates" title="Sinauer Associates">Sinauer Associates</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87893-189-7" title="Special:BookSources/978-0-87893-189-7"><bdi>978-0-87893-189-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolutionary+Biology&rft.place=Sunderland%2C+MA&rft.edition=3rd&rft.pub=Sinauer+Associates&rft.date=1998&rft.isbn=978-0-87893-189-7&rft.aulast=Futuyma&rft.aufirst=Douglas+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1"><a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">Haldane, J. B. S.</a> (1932). <a href="/wiki/The_Causes_of_Evolution" title="The Causes of Evolution"><i>The Causes of Evolution</i></a>. London; New York: <a href="/wiki/Longman" title="Longman">Longmans, Green & Co.</a> <a href="/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a> <a rel="nofollow" class="external text" href="https://lccn.loc.gov/32033284">32033284</a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/5006266">5006266</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Causes+of+Evolution&rft.place=London%3B+New+York&rft.pub=Longmans%2C+Green+%26+Co.&rft.date=1932&rft_id=info%3Aoclcnum%2F5006266&rft_id=info%3Alccn%2F32033284&rft.aulast=Haldane&rft.aufirst=J.+B.+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span> "This book is based on a series of lectures delivered in January 1931 at the Prifysgol Cymru, Aberystwyth, and entitled 'A re-examination of Darwinism'."</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1"><a href="/wiki/Julian_S._Huxley" class="mw-redirect" title="Julian S. Huxley">Huxley, Julian</a>, ed. (1976) [Originally published 1940; Oxford, UK: <a href="/wiki/Oxford_University_Press#Clarendon_Press" title="Oxford University Press">The Clarendon Press</a>]. <i>The New Systematics</i> (Reprint ed.). St. Clair Shores, MI: Scholarly Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-403-01786-7" title="Special:BookSources/978-0-403-01786-7"><bdi>978-0-403-01786-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+New+Systematics&rft.place=St.+Clair+Shores%2C+MI&rft.edition=Reprint&rft.pub=Scholarly+Press&rft.date=1976&rft.isbn=978-0-403-01786-7&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Levinson, Gene (2020). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20220521082753/https://rethinkingevolution.com/"><i>Rethinking evolution: the revolution that's hiding in plain sight</i></a>. World Scientific. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9781786347268" title="Special:BookSources/9781786347268"><bdi>9781786347268</bdi></a>. Archived from <a rel="nofollow" class="external text" href="https://rethinkingevolution.com/">the original</a> on 2022-05-21<span class="reference-accessdate">. Retrieved <span class="nowrap">2020-01-27</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Rethinking+evolution%3A+the+revolution+that%27s+hiding+in+plain+sight&rft.pub=World+Scientific&rft.date=2020&rft.isbn=9781786347268&rft.aulast=Levinson&rft.aufirst=Gene&rft_id=https%3A%2F%2Frethinkingevolution.com%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation journal cs1"><a href="/wiki/Sewall_Wright" title="Sewall Wright">Wright, Sewall</a> (March 1, 1931). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1201091">"Evolution in Mendelian Populations"</a>. <i><a href="/wiki/Genetics_(journal)" title="Genetics (journal)">Genetics</a></i>. <b>16</b> (2): 97–159. <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%2Fgenetics%2F16.2.97">10.1093/genetics/16.2.97</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1201091">1201091</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/17246615">17246615</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Genetics&rft.atitle=Evolution+in+Mendelian+Populations&rft.volume=16&rft.issue=2&rft.pages=97-159&rft.date=1931-03-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1201091%23id-name%3DPMC&rft_id=info%3Apmid%2F17246615&rft_id=info%3Adoi%2F10.1093%2Fgenetics%2F16.2.97&rft.aulast=Wright&rft.aufirst=Sewall&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1201091&rfr_id=info%3Asid%2Fen.wikipedia.org%3AModern+synthesis+%2820th+century%29" class="Z3988"></span></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 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href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a class="mw-selflink selflink">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_psychology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolutionary_psychology" title="Template:Evolutionary psychology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_psychology" title="Template talk:Evolutionary psychology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_psychology" title="Special:EditPage/Template:Evolutionary psychology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_psychology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/History_of_evolutionary_psychology" title="History of evolutionary psychology">History</a> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li></ul></li> <li><a href="/wiki/Theoretical_foundations_of_evolutionary_psychology" title="Theoretical foundations of evolutionary psychology">Theoretical foundations</a> <ul><li><a href="/wiki/Adaptationism" title="Adaptationism">Adaptationism</a></li> <li><a href="/wiki/Cognitive_revolution" title="Cognitive revolution">Cognitive revolution</a></li> <li><a href="/wiki/Cognitivism_(psychology)" title="Cognitivism (psychology)">Cognitivism</a></li> <li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene selection theory</a></li> <li><a class="mw-selflink selflink">Modern synthesis</a></li></ul></li> <li><a href="/wiki/Criticism_of_evolutionary_psychology" title="Criticism of evolutionary psychology">Criticism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;"><a href="/wiki/Human_evolution" title="Human evolution">Evolutionary<br />processes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Psychological_adaptation" title="Psychological adaptation">Adaptations</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Hamiltonian_spite" title="Hamiltonian spite">Hamiltonian spite</a></li> <li><a href="/wiki/Reciprocal_altruism_in_humans" title="Reciprocal altruism in humans">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">By-products</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness_in_humans" title="Inclusive fitness in humans">Inclusive</a></li></ul></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a> <ul><li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a> <ul><li><a href="/wiki/Costly_signaling_theory_in_evolutionary_psychology" title="Costly signaling theory in evolutionary psychology">Costly signaling</a></li> <li><a href="/wiki/Male_intrasexual_competition" title="Male intrasexual competition">Male</a>/<a href="/wiki/Female_intrasexual_competition" title="Female intrasexual competition">female intrasexual competition</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Areas</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution_of_cognition" title="Evolution of cognition">Cognition</a> /<br /><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Affect_(psychology)" title="Affect (psychology)">Affect</a> <ul><li><a href="/wiki/Affect_display" title="Affect display">Display</a></li> <li><a href="/wiki/Display_rules" title="Display rules">Display rules</a></li> <li><a href="/wiki/Facial_expression" title="Facial expression">Facial expression</a></li></ul></li> <li><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li> <li><a href="/wiki/Cognitive_module" title="Cognitive module">Cognitive module</a>/<a href="/wiki/Modularity_of_mind" title="Modularity of mind">modularity of mind</a> <ul><li><a href="/wiki/Automatic_and_controlled_processes" title="Automatic and controlled processes">Automatic and controlled processes</a></li> <li><a href="/wiki/Computational_theory_of_mind" title="Computational theory of mind">Computational theory of mind</a></li> <li><a href="/wiki/Domain-general_learning" title="Domain-general learning">Domain generality</a></li> <li><a href="/wiki/Domain_specificity" title="Domain specificity">Domain specificity</a></li> <li><a href="/wiki/Dual_process_theory" title="Dual process theory">Dual process theory</a></li></ul></li> <li><a href="/wiki/Cognitive_tradeoff_hypothesis" title="Cognitive tradeoff hypothesis">Cognitive tradeoff hypothesis</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">Evolution of the brain</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></li> <li><a href="/wiki/Fight-or-flight_response" title="Fight-or-flight response">Fight-or-flight response</a> <ul><li><a href="/wiki/Arachnophobia" title="Arachnophobia">Arachnophobia</a></li> <li><a href="/wiki/Fear_of_falling" title="Fear of falling">Basophobia</a></li> <li><a href="/wiki/Ophidiophobia" title="Ophidiophobia">Ophidiophobia</a></li></ul></li> <li><a href="/wiki/Folk_biology" title="Folk biology">Folk biology</a>/<a href="/wiki/Folk_taxonomy" title="Folk taxonomy">taxonomy</a></li> <li><a href="/wiki/Folk_psychology" title="Folk psychology">Folk psychology</a>/<a href="/wiki/Theory_of_mind" title="Theory of mind">theory of mind</a></li> <li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a> <ul><li><a href="/wiki/Flynn_effect" title="Flynn effect">Flynn effect</a></li> <li><a href="/wiki/Wason_selection_task" title="Wason selection task">Wason selection task</a></li></ul></li> <li><a href="/wiki/Motor_control" title="Motor control">Motor control</a>/<a href="/wiki/Motor_skill" title="Motor skill">skill</a></li> <li><a href="/wiki/Human_multitasking" title="Human multitasking">Multitasking</a></li> <li><a href="/wiki/Neuroscience_of_sleep" title="Neuroscience of sleep">Sleep</a></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Color vision</a></li> <li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">Eye</a></li> <li><a href="/wiki/Na%C3%AFve_physics" title="Naïve physics">Naïve physics</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_psychology_and_culture" title="Evolutionary psychology and culture">Culture</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_aesthetics" title="Evolutionary aesthetics">Aesthetics</a> <ul><li><a href="/wiki/Darwinian_literary_studies" title="Darwinian literary studies">Literary criticism</a></li> <li><a href="/wiki/Evolutionary_musicology" title="Evolutionary musicology">Musicology</a></li></ul></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Anthropology</a> <ul><li><a href="/wiki/Biological_anthropology" title="Biological anthropology">Biological</a></li></ul></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Crime</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Language</a> <ul><li><a href="/wiki/Origin_of_language" title="Origin of language">Origin</a></li> <li><a href="/wiki/Evolutionary_psychology_of_language" title="Evolutionary psychology of language">Psychology</a></li> <li><a href="/wiki/Origin_of_speech" title="Origin of speech">Speech</a></li></ul></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a> <ul><li><a href="/wiki/Moral_foundations_theory" title="Moral foundations theory">Moral foundations</a></li></ul></li> <li><a href="/wiki/Evolutionary_psychology_of_religion" title="Evolutionary psychology of religion">Religion</a> <ul><li><a href="/wiki/Evolutionary_origin_of_religions" class="mw-redirect" title="Evolutionary origin of religions">Origin</a></li></ul></li> <li><a href="/wiki/Cultural_universal" title="Cultural universal">Universals</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_developmental_psychology" title="Evolutionary developmental psychology">Development</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Attachment_theory" title="Attachment theory">Attachment</a></li> <li><a href="/wiki/Human_bonding" title="Human bonding">Bonding</a></li> <li><a href="/wiki/Affectional_bond" title="Affectional bond">Affectional</a>/<a href="/wiki/Maternal_bond" title="Maternal bond">maternal</a>/<a href="/wiki/Paternal_bond" title="Paternal bond">paternal bond</a></li> <li><a href="/wiki/Maternal_deprivation" title="Maternal deprivation">Caregiver deprivation</a></li> <li><a href="/wiki/Attachment_in_children" title="Attachment in children">Childhood attachment</a></li> <li><a href="/wiki/Cinderella_effect" title="Cinderella effect">Cinderella effect</a></li> <li><a href="/wiki/Cognitive_development" title="Cognitive development">Cognitive development</a></li> <li><a href="/wiki/Evolutionary_educational_psychology" title="Evolutionary educational psychology">Education</a></li> <li><a href="/wiki/Language_acquisition" title="Language acquisition">Language acquisition</a></li> <li><a href="/wiki/Personality_development" title="Personality development">Personality development</a></li> <li><a href="/wiki/Socialization" title="Socialization">Socialization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ergonomics" title="Ergonomics">Human factors</a> /<br /><a href="/wiki/Evolutionary_psychiatry" title="Evolutionary psychiatry">Mental health</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cognitive_ergonomics" title="Cognitive ergonomics">Cognitive ergonomics</a> <ul><li><a href="/wiki/Computer-mediated_communication" title="Computer-mediated communication">Computer-mediated communication</a></li> <li><a href="/wiki/Engineering_psychology" title="Engineering psychology">Engineering psychology</a></li> <li><a href="/wiki/Human%E2%80%93computer_interaction" title="Human–computer interaction">Human–computer interaction</a></li> <li><a href="/wiki/Media_naturalness_theory" title="Media naturalness theory">Media naturalness theory</a></li> <li><a href="/wiki/Neuroergonomics" title="Neuroergonomics">Neuroergonomics</a></li></ul></li> <li><a href="/wiki/Evolutionary_approaches_to_depression" title="Evolutionary approaches to depression">Depression</a></li> <li><a href="/wiki/Digital_media_use_and_mental_health" title="Digital media use and mental health">Digital media use and mental health</a></li> <li><a href="/wiki/Accident-proneness" title="Accident-proneness">Hypophobia</a></li> <li><a href="/wiki/Imprinted_brain_hypothesis" title="Imprinted brain hypothesis">Imprinted brain hypothesis</a></li> <li><a href="/wiki/Mind-blindness" title="Mind-blindness">Mind-blindness</a></li> <li><a href="/wiki/Psychological_effects_of_Internet_use" title="Psychological effects of Internet use">Psychological effects of Internet use</a></li> <li><a href="/wiki/Rank_theory_of_depression" title="Rank theory of depression">Rank theory of depression</a></li> <li><a href="/wiki/Evolution_of_schizophrenia" title="Evolution of schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Screen_time" title="Screen time">Screen time</a></li> <li><a href="/wiki/Smartphones_and_pedestrian_safety" title="Smartphones and pedestrian safety">Smartphones and pedestrian safety</a></li> <li><a href="/wiki/Social_aspects_of_television" title="Social aspects of television">Social aspects of television</a></li> <li><a href="/wiki/Societal_impacts_of_cars" title="Societal impacts of cars">Societal impacts of cars</a> <ul><li><a href="/wiki/Distracted_driving" title="Distracted driving">Distracted driving</a></li> <li><a href="/wiki/Lead%E2%80%93crime_hypothesis" title="Lead–crime hypothesis">Lead–crime hypothesis</a></li> <li><a href="/wiki/Mobile_phones_and_driving_safety" title="Mobile phones and driving safety">Mobile phones and driving safety</a></li> <li><a href="/wiki/Texting_while_driving" title="Texting while driving">Texting while driving</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sexology" title="Sexology">Sex</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Human_sexual_activity" title="Human sexual activity">Activity</a></li> <li><a href="/wiki/Attachment_in_adults" title="Attachment in adults">Adult attachment</a></li> <li><a href="/wiki/Age_disparity_in_sexual_relationships" title="Age disparity in sexual relationships">Age disparity</a></li> <li><a href="/wiki/Sexual_arousal" title="Sexual arousal">Arousal</a></li> <li><a href="/wiki/Concealed_ovulation" title="Concealed ovulation">Concealed ovulation</a></li> <li><a href="/wiki/Coolidge_effect" title="Coolidge effect">Coolidge effect</a></li> <li><a href="/wiki/Sexual_desire" title="Sexual desire">Desire</a></li> <li><a href="/wiki/Sexual_fantasy" title="Sexual fantasy">Fantasy</a></li> <li><a href="/wiki/Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">Hormonal motivation</a></li> <li><a href="/wiki/Sexual_jealousy" title="Sexual jealousy">Jealousy</a></li> <li><a href="/wiki/Mate_guarding_in_humans" title="Mate guarding in humans">Mate guarding</a></li> <li><a href="/wiki/Mating_preferences" title="Mating preferences">Mating preferences</a></li> <li><a href="/wiki/Human_mating_strategies" title="Human mating strategies">Mating strategies</a></li> <li><a href="/wiki/Biology_and_sexual_orientation" title="Biology and sexual orientation">Orientation</a></li> <li><a href="/wiki/Ovulatory_shift_hypothesis" title="Ovulatory shift hypothesis">Ovulatory shift hypothesis</a></li> <li><a href="/wiki/Pair_bond" title="Pair bond">Pair bond</a></li> <li><a href="/wiki/Physical_attractiveness" title="Physical attractiveness">Physical</a>/<a href="/wiki/Sexual_attraction" title="Sexual attraction">Sexual attraction</a></li> <li><a href="/wiki/Human_sexuality" title="Human sexuality">Sexuality</a>/<a href="/wiki/Human_male_sexuality" title="Human male sexuality">male</a>/<a href="/wiki/Human_female_sexuality" title="Human female sexuality">female</a></li> <li><a href="/wiki/Sexy_son_hypothesis" title="Sexy son hypothesis">Sexy son hypothesis</a></li> <li><a href="/wiki/Westermarck_effect" title="Westermarck effect">Westermarck effect</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sex_differences_in_psychology" title="Sex differences in psychology">Sex differences</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Aggression" title="Aggression">Aggression</a></li> <li><a href="/wiki/Sex_and_gender_differences_in_autism" title="Sex and gender differences in autism">Autism</a></li> <li><a href="/wiki/Sex_differences_in_cognition" title="Sex differences in cognition">Cognition</a></li> <li><a href="/wiki/Sex_differences_in_crime" title="Sex differences in crime">Crime</a></li> <li><a href="/wiki/Sexual_division_of_labour" title="Sexual division of labour">Division of labour</a></li> <li><a href="/wiki/Sex_differences_in_emotional_intelligence" title="Sex differences in emotional intelligence">Emotional intelligence</a></li> <li><a href="/wiki/Empathising%E2%80%93systemising_theory" title="Empathising–systemising theory">Empathising–systemising theory</a></li> <li><a href="/wiki/Gender_role" title="Gender role">Gender role</a></li> <li><a href="/wiki/Sex_differences_in_intelligence" title="Sex differences in intelligence">Intelligence</a></li> <li><a href="/wiki/Sex_differences_in_memory" title="Sex differences in memory">Memory</a></li> <li><a href="/wiki/Mental_disorders_and_gender" title="Mental disorders and gender">Mental health</a></li> <li><a href="/wiki/Sex_differences_in_narcissism" title="Sex differences in narcissism">Narcissism</a></li> <li><a href="/wiki/Neuroscience_of_sex_differences" title="Neuroscience of sex differences">Neuroscience</a></li> <li><a href="/wiki/Sex_differences_in_schizophrenia" title="Sex differences in schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Substance_abuse" title="Substance abuse">Substance abuse</a></li> <li><a href="/wiki/Gender_differences_in_suicide" title="Gender differences in suicide">Suicide</a></li> <li><a href="/wiki/Variability_hypothesis" title="Variability hypothesis">Variability hypothesis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Related subjects</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Academic disciplines</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Behavioral_economics" title="Behavioral economics">Behavioral</a>/<a href="/wiki/Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a></li> <li><a href="/wiki/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a>/<a href="/wiki/Behavioural_genetics" title="Behavioural genetics">genetics</a></li> <li><a href="/wiki/Affective_neuroscience" title="Affective neuroscience">Affective</a>/<a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">behavioral</a>/<a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">cognitive</a>/<a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">evolutionary neuroscience</a></li> <li><a href="/wiki/Biocultural_anthropology" title="Biocultural anthropology">Biocultural anthropology</a></li> <li><a href="/wiki/Biological_psychiatry" title="Biological psychiatry">Biological psychiatry</a></li> <li><a href="/wiki/Cognitive_psychology" title="Cognitive psychology">Cognitive psychology</a></li> <li><a href="/wiki/Cognitive_science" title="Cognitive science">Cognitive science</a></li> <li><a href="/wiki/Cross-cultural_psychology" title="Cross-cultural psychology">Cross-cultural psychology</a></li> <li><a href="/wiki/Ethology" title="Ethology">Ethology</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Functional_psychology" title="Functional psychology">Functional psychology</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a href="/wiki/Philosophy_of_mind" title="Philosophy of mind">Philosophy of mind</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Research topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Great_ape_language" title="Great ape language">Great ape language</a></li> <li><a href="/wiki/Human%E2%80%93animal_communication" title="Human–animal communication">Human–animal communication</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Primate_cognition" title="Primate cognition">Primate cognition</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li> <li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theoretical positions</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/Cultural_selection_theory" title="Cultural selection theory">Cultural selection theory</a></li> <li><a href="/wiki/Determinism" title="Determinism">Determinism</a>/<a href="/wiki/Indeterminism" title="Indeterminism">indeterminism</a> <ul><li><a href="/wiki/Biological_determinism" title="Biological determinism">Biological determinism</a></li> <li><a href="/wiki/Connectionism" title="Connectionism">Connectionism</a></li> <li><a href="/wiki/Cultural_determinism" title="Cultural determinism">Cultural determinism</a></li> <li><a href="/wiki/Environmental_determinism" title="Environmental determinism">Environmental determinism</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Psychological_nativism" title="Psychological nativism">Psychological nativism</a></li> <li><a href="/wiki/Social_constructionism" title="Social constructionism">Social constructionism</a></li> <li><a href="/wiki/Social_determinism" title="Social determinism">Social determinism</a></li> <li><a href="/wiki/Standard_social_science_model" title="Standard social science model">Standard social science 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