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Evolution of sexual reproduction - Wikipedia
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class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Advantages of sex and sexual reproduction</span> </div> </a> <button aria-controls="toc-Advantages_of_sex_and_sexual_reproduction-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 Advantages of sex and sexual reproduction subsection</span> </button> <ul id="toc-Advantages_of_sex_and_sexual_reproduction-sublist" class="vector-toc-list"> <li id="toc-Advantages_due_to_genetic_variation,_DNA_repair_and_genetic_complementation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Advantages_due_to_genetic_variation,_DNA_repair_and_genetic_complementation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Advantages due to genetic variation, DNA repair and genetic complementation</span> </div> </a> <ul id="toc-Advantages_due_to_genetic_variation,_DNA_repair_and_genetic_complementation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Protection_from_major_genetic_mutation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Protection_from_major_genetic_mutation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Protection from major genetic mutation</span> </div> </a> <ul id="toc-Protection_from_major_genetic_mutation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Novel_genotypes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Novel_genotypes"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Novel genotypes</span> </div> </a> <ul id="toc-Novel_genotypes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Increased_resistance_to_parasites" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Increased_resistance_to_parasites"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Increased resistance to parasites</span> </div> </a> <ul id="toc-Increased_resistance_to_parasites-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Disadvantages_of_sex_and_sexual_reproduction" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Disadvantages_of_sex_and_sexual_reproduction"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Disadvantages of sex and sexual reproduction</span> </div> </a> <button aria-controls="toc-Disadvantages_of_sex_and_sexual_reproduction-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 Disadvantages of sex and sexual reproduction subsection</span> </button> <ul id="toc-Disadvantages_of_sex_and_sexual_reproduction-sublist" class="vector-toc-list"> <li id="toc-Population_expansion_cost_of_sex" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Population_expansion_cost_of_sex"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Population expansion cost of sex</span> </div> </a> <ul id="toc-Population_expansion_cost_of_sex-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Selfish_cytoplasmic_genes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Selfish_cytoplasmic_genes"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Selfish cytoplasmic genes</span> </div> </a> <ul id="toc-Selfish_cytoplasmic_genes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetic_heritability_cost_of_sex" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_heritability_cost_of_sex"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Genetic heritability cost of sex</span> </div> </a> <ul id="toc-Genetic_heritability_cost_of_sex-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-DNA_repair_and_complementation" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#DNA_repair_and_complementation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>DNA repair and complementation</span> </div> </a> <ul id="toc-DNA_repair_and_complementation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Deleterious_mutation_clearance" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Deleterious_mutation_clearance"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Deleterious mutation clearance</span> </div> </a> <button aria-controls="toc-Deleterious_mutation_clearance-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 Deleterious mutation clearance subsection</span> </button> <ul id="toc-Deleterious_mutation_clearance-sublist" class="vector-toc-list"> <li id="toc-Evading_harmful_mutation_build-up" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evading_harmful_mutation_build-up"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Evading harmful mutation build-up</span> </div> </a> <ul id="toc-Evading_harmful_mutation_build-up-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Removal_of_deleterious_genes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Removal_of_deleterious_genes"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Removal of deleterious genes</span> </div> </a> <ul id="toc-Removal_of_deleterious_genes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Other_explanations" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Other_explanations"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Other explanations</span> </div> </a> <button aria-controls="toc-Other_explanations-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 Other explanations subsection</span> </button> <ul id="toc-Other_explanations-sublist" class="vector-toc-list"> <li id="toc-Geodakyan's_evolutionary_theory_of_sex" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Geodakyan's_evolutionary_theory_of_sex"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Geodakyan's evolutionary theory of sex</span> </div> </a> <ul id="toc-Geodakyan's_evolutionary_theory_of_sex-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Speed_of_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Speed_of_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Speed of evolution</span> </div> </a> <ul id="toc-Speed_of_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Libertine_bubble_theory" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Libertine_bubble_theory"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>Libertine bubble theory</span> </div> </a> <ul id="toc-Libertine_bubble_theory-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Origin_of_sexual_reproduction" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Origin_of_sexual_reproduction"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Origin of sexual reproduction</span> </div> </a> <button aria-controls="toc-Origin_of_sexual_reproduction-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 Origin of sexual reproduction subsection</span> </button> <ul id="toc-Origin_of_sexual_reproduction-sublist" class="vector-toc-list"> <li id="toc-Diploidy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Diploidy"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Diploidy</span> </div> </a> <ul id="toc-Diploidy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Meiosis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Meiosis"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Meiosis</span> </div> </a> <ul id="toc-Meiosis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Virus-like_RNA-based_origin" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Virus-like_RNA-based_origin"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Virus-like RNA-based origin</span> </div> </a> <ul id="toc-Virus-like_RNA-based_origin-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Parasitic_DNA_elements" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Parasitic_DNA_elements"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.4</span> <span>Parasitic DNA elements</span> </div> </a> <ul id="toc-Parasitic_DNA_elements-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Partial_predation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Partial_predation"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.5</span> <span>Partial predation</span> </div> </a> <ul id="toc-Partial_predation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Vaccination-like_process" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Vaccination-like_process"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.6</span> <span>Vaccination-like process</span> </div> </a> <ul id="toc-Vaccination-like_process-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mechanistic_origin_of_sexual_reproduction" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Mechanistic_origin_of_sexual_reproduction"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Mechanistic origin of sexual reproduction</span> </div> </a> <button aria-controls="toc-Mechanistic_origin_of_sexual_reproduction-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 Mechanistic origin of sexual reproduction subsection</span> </button> <ul id="toc-Mechanistic_origin_of_sexual_reproduction-sublist" class="vector-toc-list"> <li id="toc-Viral_eukaryogenesis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Viral_eukaryogenesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Viral eukaryogenesis</span> </div> </a> <ul id="toc-Viral_eukaryogenesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Neomuran_revolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Neomuran_revolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.2</span> <span>Neomuran revolution</span> </div> </a> <ul id="toc-Neomuran_revolution-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Questions" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Questions"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Questions</span> </div> </a> <ul id="toc-Questions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>References</span> </div> </a> <ul id="toc-References-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">11</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> 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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">Evolution of sexual reproduction</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 15 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-15" 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">15 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%AA%D8%B7%D9%88%D8%B1_%D8%A7%D9%84%D8%AA%D9%83%D8%A7%D8%AB%D8%B1_%D8%A7%D9%84%D8%AC%D9%86%D8%B3%D9%8A" title="تطور التكاثر الجنسي – Arabic" lang="ar" hreflang="ar" data-title="تطور التكاثر الجنسي" 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%AF%E0%A7%8C%E0%A6%A8_%E0%A6%AA%E0%A7%8D%E0%A6%B0%E0%A6%9C%E0%A6%A8%E0%A6%A8%E0%A7%87%E0%A6%B0_%E0%A6%AC%E0%A6%BF%E0%A6%AC%E0%A6%B0%E0%A7%8D%E0%A6%A4%E0%A6%A8" title="যৌন প্রজননের বিবর্তন – Bangla" lang="bn" hreflang="bn" data-title="যৌন প্রজননের বিবর্তন" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://es.wikipedia.org/wiki/Evoluci%C3%B3n_de_la_reproducci%C3%B3n_sexual" title="Evolución de la reproducción sexual – Spanish" lang="es" hreflang="es" data-title="Evolución de la reproducción sexual" 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/Sexu-ugalketaren_eboluzio" title="Sexu-ugalketaren eboluzio – Basque" lang="eu" hreflang="eu" data-title="Sexu-ugalketaren eboluzio" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%81%D8%B1%DA%AF%D8%B4%D8%AA_%D8%AA%D9%88%D9%84%DB%8C%D8%AF%D9%85%D8%AB%D9%84_%D8%AC%D9%86%D8%B3%DB%8C" title="فرگشت تولیدمثل جنسی – Persian" lang="fa" hreflang="fa" data-title="فرگشت تولیدمثل جنسی" 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/%C3%89volution_de_la_reproduction_sexu%C3%A9e" title="Évolution de la reproduction sexuée – French" lang="fr" hreflang="fr" data-title="Évolution de la reproduction sexuée" 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/Evoluci%C3%B3n_da_reproduci%C3%B3n_sexual" title="Evolución da reprodución sexual – Galician" lang="gl" hreflang="gl" data-title="Evolución da reprodución sexual" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Evolusi_reproduksi_seksual" title="Evolusi reproduksi seksual – Indonesian" lang="id" hreflang="id" data-title="Evolusi reproduksi seksual" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%90%D7%91%D7%95%D7%9C%D7%95%D7%A6%D7%99%D7%94_%D7%A9%D7%9C_%D7%94%D7%A8%D7%91%D7%99%D7%99%D7%94_%D7%94%D7%9E%D7%99%D7%A0%D7%99%D7%AA" title="האבולוציה של הרבייה המינית – Hebrew" lang="he" hreflang="he" data-title="האבולוציה של הרבייה המינית" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Evolusi_pembiakan_seks" title="Evolusi pembiakan seks – Malay" lang="ms" hreflang="ms" data-title="Evolusi pembiakan seks" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%AF_%D8%B2%D9%88%D8%AC%D9%8A_%D8%AA%DA%A9%D8%AB%D8%B1_%D8%AA%DA%A9%D8%A7%D9%85%D9%84" title="د زوجي تکثر تکامل – Pashto" lang="ps" hreflang="ps" data-title="د زوجي تکثر تکامل" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Origem_e_evolu%C3%A7%C3%A3o_do_sexo_eucari%C3%B3tico" title="Origem e evolução do sexo eucariótico – Portuguese" lang="pt" hreflang="pt" data-title="Origem e evolução do sexo eucariótico" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/E%C5%9Feyli_%C3%BCremenin_evrimi" title="Eşeyli üremenin evrimi – Turkish" lang="tr" hreflang="tr" data-title="Eşeyli üremenin evrimi" 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class="sidebar sidebar-collapse nomobile nowraplinks" style="border:2px solid #90ee90"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#90ee90;padding:0.2em;font-size:175%;font-weight:bold"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></th></tr><tr><td class="sidebar-image"><span class="notpageimage" typeof="mw:File"><a href="/wiki/File:Darwin%27s_finches_by_Gould.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/180px-Darwin%27s_finches_by_Gould.jpg" decoding="async" width="180" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/270px-Darwin%27s_finches_by_Gould.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/360px-Darwin%27s_finches_by_Gould.jpg 2x" 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href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Natural history</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/Human_evolution" title="Human evolution">Human evolution</a> <ul><li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li></ul></li> <li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogeny</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Classification</a></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Evolutionary taxonomy</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Extinction_event" title="Extinction event">Extinction event</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">History of evolutionary theory</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Overview</a></li> <li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Before Darwin</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Darwin</a></li> <li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">Origin of Species</a></i></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">Before synthesis</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evo-devo</a></li> <li><a href="/wiki/Current_research_in_evolutionary_biology" class="mw-redirect" title="Current research in evolutionary biology">Current research</a></li> <li><a href="/wiki/History_of_speciation" title="History of speciation">History of speciation</a></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a> (<a href="/wiki/Timeline_of_paleontology" title="Timeline of paleontology">timeline</a>)</li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Fields and applications</div><div class="sidebar-list-content mw-collapsible-content plainlist"> <ul><li><a href="/wiki/Applications_of_evolution" title="Applications of evolution">Applications of evolution</a></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Biosocial criminology</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_aesthetics" title="Evolutionary aesthetics">Evolutionary aesthetics</a></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Evolutionary anthropology</a></li> <li><a href="/wiki/Evolutionary_computation" title="Evolutionary computation">Evolutionary computation</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Evolutionary_economics" title="Evolutionary economics">Evolutionary economics</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Evolutionary ethics</a></li> <li><a href="/wiki/Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Evolutionary linguistics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/Evolutionary_physiology" title="Evolutionary physiology">Evolutionary physiology</a></li> <li><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></li> <li><a href="/wiki/Experimental_evolution" title="Experimental evolution">Experimental evolution</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a></li> <li><a href="/wiki/Paleontology" title="Paleontology">Paleontology</a></li> <li><a href="/wiki/Selective_breeding" title="Selective breeding">Selective breeding</a></li> <li><a href="/wiki/Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Speciation experiments</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li> <li><a href="/wiki/Island_biogeography" class="mw-redirect" title="Island biogeography">Island biogeography</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Universal_Darwinism" title="Universal Darwinism">Universal Darwinism</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Social implications</div><div class="sidebar-list-content mw-collapsible-content plainlist"> <ul><li><a href="/wiki/Eugenics" title="Eugenics">Eugenics</a></li> <li><a href="/wiki/Evolution_as_fact_and_theory" title="Evolution as fact and theory">Evolution as fact and theory</a></li> <li><a href="/wiki/Dysgenics" title="Dysgenics">Dysgenics</a></li> <li><a href="/wiki/Social_effects_of_evolutionary_theory" title="Social effects of evolutionary theory">Social effects</a></li> <li><a href="/wiki/Rejection_of_evolution_by_religious_groups" title="Rejection of evolution by religious groups">Creation–evolution controversy</a></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic evolution</a></li> <li><a href="/wiki/Objections_to_evolution" title="Objections to evolution">Objections to evolution</a></li> <li><a href="/wiki/Level_of_support_for_evolution" title="Level of support for evolution">Level of support</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature-nurture controversy</a></li></ul></div></div></td> </tr><tr><td class="sidebar-below hlist"> <ul><li><span class="nowrap"><span class="nowrap"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Tree_of_life.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/16px-Tree_of_life.svg.png" decoding="async" width="16" height="13" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/24px-Tree_of_life.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/32px-Tree_of_life.svg.png 2x" data-file-width="915" data-file-height="760" /></a></span> </span><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></span></li> <li><span class="nowrap"><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></span></li></ul></td></tr><tr><td class="sidebar-navbar"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolutionary_biology" title="Template:Evolutionary biology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_biology" title="Template talk:Evolutionary biology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_biology" title="Special:EditPage/Template:Evolutionary biology"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Ladybird-Coccinellidae-mating.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/dc/Ladybird-Coccinellidae-mating.jpg/220px-Ladybird-Coccinellidae-mating.jpg" decoding="async" width="220" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/dc/Ladybird-Coccinellidae-mating.jpg/330px-Ladybird-Coccinellidae-mating.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/dc/Ladybird-Coccinellidae-mating.jpg/440px-Ladybird-Coccinellidae-mating.jpg 2x" data-file-width="604" data-file-height="434" /></a><figcaption><a href="/wiki/Ladybird" class="mw-redirect" title="Ladybird">Ladybirds</a> mating</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:2012-04-03_16-43-53-amaryllis-sp.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/2012-04-03_16-43-53-amaryllis-sp.jpg/220px-2012-04-03_16-43-53-amaryllis-sp.jpg" decoding="async" width="220" height="146" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/2012-04-03_16-43-53-amaryllis-sp.jpg/330px-2012-04-03_16-43-53-amaryllis-sp.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/76/2012-04-03_16-43-53-amaryllis-sp.jpg/440px-2012-04-03_16-43-53-amaryllis-sp.jpg 2x" data-file-width="4288" data-file-height="2848" /></a><figcaption><a href="/wiki/Pollen" title="Pollen">Pollen</a> production is an essential step in sexual reproduction of <a href="/wiki/Seed_plant" title="Seed plant">seed plants</a>.</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1233989161">.mw-parser-output .unsolved{margin:0.5em 0 1em 1em;border:#ccc solid;padding:0.35em 0.35em 0.35em 2.2em;background-color:var(--background-color-interactive-subtle);background-image:url("https://upload.wikimedia.org/wikipedia/commons/2/26/Question%2C_Web_Fundamentals.svg");background-position:top 50%left 0.35em;background-size:1.5em;background-repeat:no-repeat}@media(min-width:720px){.mw-parser-output .unsolved{clear:right;float:right;max-width:25%}}.mw-parser-output .unsolved-label{font-weight:bold}.mw-parser-output .unsolved-body{margin:0.35em;font-style:italic}.mw-parser-output .unsolved-more{font-size:smaller}</style> <div role="note" aria-labelledby="unsolved-label-biology" class="unsolved"> <div><span class="unsolved-label" id="unsolved-label-biology">Unsolved problem in biology</span>:</div> <div class="unsolved-body">What <a href="/wiki/Selection_pressure" class="mw-redirect" title="Selection pressure">selection pressures</a> led to the evolution and maintenance of sexual reproduction?</div> <div class="unsolved-more"><a href="/wiki/List_of_unsolved_problems_in_biology" title="List of unsolved problems in biology">(more unsolved problems in biology)</a></div> </div> <p><b>Evolution of sexual reproduction</b> describes how <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexually reproducing</a> <a href="/wiki/Animal" title="Animal">animals</a>, <a href="/wiki/Plant" title="Plant">plants</a>, <a href="/wiki/Fungus" title="Fungus">fungi</a> and <a href="/wiki/Protist" title="Protist">protists</a> could have <a href="/wiki/Evolution" title="Evolution">evolved</a> from a <a href="/wiki/Common_descent" title="Common descent">common ancestor</a> that was a single-celled <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> <a href="/wiki/Species" title="Species">species</a>.<sup id="cite_ref-Letunic_I_and_Bork_P_1-0" class="reference"><a href="#cite_note-Letunic_I_and_Bork_P-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Sexual reproduction is widespread in eukaryotes, though a few eukaryotic species have secondarily lost the ability to reproduce sexually, such as <a href="/wiki/Bdelloidea" title="Bdelloidea">Bdelloidea</a>, and some plants and animals routinely reproduce <a href="/wiki/Asexual_reproduction" title="Asexual reproduction">asexually</a> (by <a href="/wiki/Apomixis" title="Apomixis">apomixis</a> and <a href="/wiki/Parthenogenesis" title="Parthenogenesis">parthenogenesis</a>) without entirely having lost <a href="/wiki/Sex" title="Sex">sex</a>. The evolution of sexual reproduction contains two related yet distinct themes: its <i>origin</i> and its <i>maintenance.</i> Bacteria and Archaea (<a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>) have processes that can transfer DNA from one cell to another (<a href="/wiki/Sexual_conjugation" class="mw-redirect" title="Sexual conjugation">conjugation</a>, <a href="/wiki/Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">transformation</a>, and <a href="/wiki/Transduction_(genetics)" title="Transduction (genetics)">transduction</a><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup>), but it is unclear if these processes are evolutionarily related to sexual reproduction in Eukaryotes.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In eukaryotes, true sexual reproduction by meiosis and cell fusion is thought to have arisen in the <a href="/wiki/Last_eukaryotic_common_ancestor" class="mw-redirect" title="Last eukaryotic common ancestor">last eukaryotic common ancestor</a>, possibly via several processes of varying success, and then to have persisted.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Since hypotheses for the origin of sex are difficult to verify experimentally (outside of <a href="/wiki/Evolutionary_computation" title="Evolutionary computation">evolutionary computation</a>), most current work has focused on the persistence of sexual reproduction over evolutionary time. The maintenance of sexual reproduction (specifically, of its <a href="/wiki/Dioecious" class="mw-redirect" title="Dioecious">dioecious</a> form) by natural selection in a highly competitive world has long been one of the major mysteries of biology, since both other known mechanisms of reproduction – <a href="/wiki/Asexual_reproduction" title="Asexual reproduction">asexual reproduction</a> and <a href="/wiki/Hermaphroditism" class="mw-redirect" title="Hermaphroditism">hermaphroditism</a> – possess apparent advantages over it. Asexual reproduction can proceed by budding, fission, or spore formation and does not involve the union of gametes, which accordingly results in <a href="#xspeed">a much faster rate</a> of reproduction compared to sexual reproduction, where 50% of offspring are males and unable to produce offspring themselves. In hermaphroditic reproduction, each of the two parent organisms required for the formation of a <a href="/wiki/Zygote" title="Zygote">zygote</a> can provide either the male or the female gamete, which leads to advantages in both size and genetic variance of a population. </p><p>Sexual reproduction therefore must offer significant <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> advantages because, despite the two-fold cost of sex (see below), it dominates among multicellular forms of life, implying that the fitness of offspring produced by sexual processes outweighs the costs. Sexual reproduction derives from <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a>, where parent genotypes are reorganised and shared with the offspring. This stands in contrast to single-parent asexual <a href="/wiki/Self-replication" title="Self-replication">replication</a>, where the offspring is always identical to the parents (barring mutation). Recombination supplies two <a href="/wiki/Fault-tolerance" class="mw-redirect" title="Fault-tolerance">fault-tolerance</a> mechanisms at the molecular level: <i>recombinational <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a></i> (promoted during <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> because <a href="/wiki/Homologous_chromosome" title="Homologous chromosome">homologous chromosomes</a> pair at that time) and <i><a href="/wiki/Complementation_(genetics)" title="Complementation (genetics)">complementation</a></i> (also known as <a href="/wiki/Heterosis" title="Heterosis">heterosis</a>, hybrid vigour or masking of mutations). </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Historical_perspective">Historical perspective</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=1" title="Edit section: Historical perspective"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Reproduction, including modes of sexual reproduction, features in the writings of <a href="/wiki/Aristotle" title="Aristotle">Aristotle</a>; modern philosophical-scientific thinking on the problem dates from at least <a href="/wiki/Erasmus_Darwin" title="Erasmus Darwin">Erasmus Darwin</a> (1731–1802) in the 18th century.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> <a href="/wiki/August_Weismann" title="August Weismann">August Weismann</a> picked up the thread in 1885, arguing that sex serves to generate <a href="/wiki/Genetic_variation" title="Genetic variation">genetic variation</a>, as detailed in the majority of the explanations below.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> On the other hand, <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> (1809–1882) concluded that the effect of <a href="/wiki/Heterosis" title="Heterosis">hybrid vigor</a> (complementation) "is amply sufficient to account for the … genesis of the two sexes".<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> This is consistent with the repair and complementation hypothesis, described below. Since the emergence of the <a href="/wiki/Modern_evolutionary_synthesis" class="mw-redirect" title="Modern evolutionary synthesis">modern evolutionary synthesis</a> in the 20th century, numerous biologists including <a href="/wiki/W._D._Hamilton" title="W. D. Hamilton">W. D. Hamilton</a>, <a href="/wiki/Alexey_Kondrashov" title="Alexey Kondrashov">Alexey Kondrashov</a>, <a href="/wiki/George_C._Williams_(biologist)" class="mw-redirect" title="George C. Williams (biologist)">George C. Williams</a>, Harris Bernstein, Carol Bernstein, Michael M. Cox, Frederic A. Hopf and Richard E. Michod – have suggested competing explanations for how a vast array of different living species maintain sexual reproduction. </p> <div class="mw-heading mw-heading2"><h2 id="Advantages_of_sex_and_sexual_reproduction">Advantages of sex and sexual reproduction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=2" title="Edit section: Advantages of sex and sexual reproduction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-One_source plainlinks metadata ambox ambox-content ambox-one_source" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>relies largely or entirely upon a <a href="/wiki/Wikipedia:Identifying_reliable_sources" class="mw-redirect" title="Wikipedia:Identifying reliable sources">single source</a></b>.<span class="hide-when-compact"> Relevant discussion may be found on the <a href="/wiki/Talk:Evolution_of_sexual_reproduction##" title="Talk:Evolution of sexual reproduction">talk page</a>. Please help <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Evolution_of_sexual_reproduction&action=edit">improve this article</a> by introducing <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">citations</a> to additional sources at this section.</span> <span class="date-container"><i>(<span class="date">August 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>The concept of sex includes two fundamental phenomena: the sexual process (fusion of genetic information of two individuals) and sexual differentiation (separation of this information into two parts). Depending on the presence or absence of these phenomena, all of the existing forms of reproduction can be classified as asexual, hermaphrodite or dioecious. The sexual process and sexual differentiation are different phenomena, and, in essence, are diametrically opposed. The first creates (increases) diversity of genotypes, and the second decreases it by half. </p><p>Reproductive advantages of the asexual forms are in quantity of the progeny, and the advantages of the hermaphrodite forms are in maximal diversity. Transition from the hermaphrodite to dioecious state leads to a loss of at least half of the diversity. So, the primary challenge is to explain the advantages given by sexual differentiation, i.e. the benefits of two separate sexes compared to hermaphrodites rather than to explain benefits of sexual forms (hermaphrodite + dioecious) over asexual ones. It has already been understood that since sexual reproduction is not associated with any clear reproductive advantages over asexual reproduction, there should be some important advantages in evolution.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template noprint noexcerpt Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:NOTRS" class="mw-redirect" title="Wikipedia:NOTRS"><span title="Does the information in this source still hold up? (August 2021)">better source needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Advantages_due_to_genetic_variation,_DNA_repair_and_genetic_complementation"><span id="Advantages_due_to_genetic_variation.2C_DNA_repair_and_genetic_complementation"></span>Advantages due to genetic variation, DNA repair and genetic complementation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=3" title="Edit section: Advantages due to genetic variation, DNA repair and genetic complementation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-More_citations_needed plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Evolution_of_sexual_reproduction" title="Special:EditPage/Evolution of sexual reproduction">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a> in this section. Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Evolution+of+sexual+reproduction%22">"Evolution of sexual reproduction"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Evolution+of+sexual+reproduction%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Evolution+of+sexual+reproduction%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Evolution+of+sexual+reproduction%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Evolution+of+sexual+reproduction%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Evolution+of+sexual+reproduction%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">April 2023</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table><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">See also: <a href="/wiki/Hill%E2%80%93Robertson_effect" title="Hill–Robertson effect">Hill–Robertson effect</a></div> <p>For the advantage due to genetic variation, there are three possible reasons this might happen. First, sexual reproduction can combine the effects of two beneficial <a href="/wiki/Mutation" title="Mutation">mutations</a> in the same individual (i.e. sex aids in the spread of advantageous traits) without the mutations having to have occurred one after another in a single line of descendants.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Reliable_sources" title="Wikipedia:Reliable sources"><span title="The material near this tag may rely on an unreliable source. (April 2012)">unreliable source?</span></a></i>]</sup> Second, sex acts to bring together currently deleterious mutations to create severely unfit individuals that are then eliminated from the population (i.e. sex aids in the removal of deleterious genes). However, in organisms containing only one set of chromosomes, deleterious mutations would be eliminated immediately, and therefore removal of harmful mutations is an unlikely benefit for sexual reproduction. Lastly, sex creates new gene combinations that may be more fit than previously existing ones, or may simply lead to reduced competition among relatives. </p><p>For the advantage due to <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a>, there is an immediate large benefit of removing DNA damage by recombinational DNA repair during <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> (assuming the initial mutation rate is higher than optimal<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup>), since this removal allows greater survival of progeny with undamaged DNA. The advantage of <a href="/wiki/Complementation_(genetics)" title="Complementation (genetics)">complementation</a> to each sexual partner is avoidance of the bad effects of their deleterious recessive genes in progeny by the masking effect of normal dominant genes contributed by the other partner.<sup id="cite_ref-Bernstein1985_13-0" class="reference"><a href="#cite_note-Bernstein1985-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bernstein1986_14-0" class="reference"><a href="#cite_note-Bernstein1986-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>The classes of hypotheses based on the creation of variation are further broken down below. Any number of these hypotheses may be true in any given species (they are not <a href="/wiki/Mutually_exclusive_events" class="mw-redirect" title="Mutually exclusive events">mutually exclusive</a>), and different hypotheses may apply in different species. However, a research framework based on creation of variation has yet to be found that allows one to determine whether the reason for sex is universal for all sexual species, and, if not, which mechanisms are acting in each species. </p><p>On the other hand, the maintenance of sex based on DNA repair and complementation applies widely to all sexual species. </p> <div class="mw-heading mw-heading3"><h3 id="Protection_from_major_genetic_mutation">Protection from major genetic mutation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=4" title="Edit section: Protection from major genetic mutation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In contrast to the view that sex promotes genetic variation, Heng,<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> and Gorelick and Heng<sup id="cite_ref-Gorelick2011_16-0" class="reference"><a href="#cite_note-Gorelick2011-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> reviewed evidence that sex actually acts as a constraint on genetic variation. They consider that sex acts as a coarse filter, weeding out major genetic changes, such as chromosomal rearrangements, but permitting minor variation, such as changes at the nucleotide or gene level (that are often neutral) to pass through the sexual sieve. </p> <div class="mw-heading mw-heading3"><h3 id="Novel_genotypes">Novel genotypes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=5" title="Edit section: Novel genotypes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-More_citations_needed plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Evolution_of_sexual_reproduction" title="Special:EditPage/Evolution of sexual reproduction">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a> in this section. Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Evolution+of+sexual+reproduction%22">"Evolution of sexual reproduction"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Evolution+of+sexual+reproduction%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Evolution+of+sexual+reproduction%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Evolution+of+sexual+reproduction%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Evolution+of+sexual+reproduction%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Evolution+of+sexual+reproduction%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">April 2023</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Evolsex-dia2a.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Evolsex-dia2a.svg/250px-Evolsex-dia2a.svg.png" decoding="async" width="250" height="173" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Evolsex-dia2a.svg/375px-Evolsex-dia2a.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Evolsex-dia2a.svg/500px-Evolsex-dia2a.svg.png 2x" data-file-width="700" data-file-height="483" /></a><figcaption>This diagram illustrates how sex might create novel genotypes more rapidly. Two advantageous alleles <i>A</i> and <i>B</i> occur at random. The two alleles are recombined rapidly in a sexual population (top), but in an asexual population (bottom) the two alleles must independently arise because of <a href="/wiki/Clonal_interference" title="Clonal interference">clonal interference</a>.</figcaption></figure> <p>Sex could be a method by which novel genotypes are created. Because sex combines genes from two individuals, sexually reproducing populations can more easily combine advantageous genes than can asexual populations. If, in a sexual population, two different advantageous <a href="/wiki/Alleles" class="mw-redirect" title="Alleles">alleles</a> arise at different loci on a chromosome in different members of the population, a chromosome containing the two advantageous alleles can be produced within a few generations by <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a>. However, should the same two alleles arise in different members of an asexual population, the only way that one chromosome can develop the other allele is to independently gain the same mutation, which would take much longer. Several studies have addressed counterarguments, and the question of whether this model is sufficiently robust to explain the predominance of sexual versus asexual reproduction remains.<sup id="cite_ref-Birdsell_17-0" class="reference"><a href="#cite_note-Birdsell-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 73–86">: 73–86 </span></sup><sup id="cite_ref-Gorelick2011_16-1" class="reference"><a href="#cite_note-Gorelick2011-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Ronald_Fisher" title="Ronald Fisher">Ronald Fisher</a> suggested that sex might facilitate the spread of advantageous genes by allowing them to better escape their genetic surroundings, if they should arise on a chromosome with deleterious genes. </p><p>Supporters of these theories respond to the balance argument that the individuals produced by sexual and asexual reproduction may differ in other respects too – which may influence the persistence of sexuality. For example, in the <a href="/wiki/Asexual_reproduction#Alternation_between_sexual_and_asexual_reproduction" title="Asexual reproduction">heterogamous</a> <a href="/wiki/Water_flea" class="mw-redirect" title="Water flea">water fleas</a> of the genus <i>Cladocera</i>, sexual offspring form eggs which are better able to survive the winter versus those the fleas produce asexually. </p> <div class="mw-heading mw-heading3"><h3 id="Increased_resistance_to_parasites">Increased resistance to parasites</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=6" title="Edit section: Increased resistance to parasites"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>One of the most widely discussed theories to explain the persistence of sex is that it is maintained to assist sexual individuals in resisting <a href="/wiki/Parasite" class="mw-redirect" title="Parasite">parasites</a>, also known as the <a href="/wiki/Red_Queen_Hypothesis" class="mw-redirect" title="Red Queen Hypothesis">Red Queen Hypothesis</a>.<sup id="cite_ref-ridley_19-0" class="reference"><a href="#cite_note-ridley-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Birdsell_17-1" class="reference"><a href="#cite_note-Birdsell-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 113–117">: 113–117 </span></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-red_21-0" class="reference"><a href="#cite_note-red-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-parasite_22-0" class="reference"><a href="#cite_note-parasite-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>When an environment changes, previously neutral or deleterious alleles can become favourable. If the environment changed sufficiently rapidly (i.e. between generations), these changes in the environment can make sex advantageous for the individual. Such rapid changes in environment are caused by the co-evolution between hosts and parasites. </p><p>Imagine, for example that there is one gene in parasites with two alleles <i>p</i> and <i>P</i> conferring two types of parasitic ability, and one gene in hosts with two alleles <i>h</i> and <i>H</i>, conferring two types of parasite resistance, such that parasites with allele <i>p</i> can attach themselves to hosts with the allele <i>h</i>, and <i>P</i> to <i>H</i>. Such a situation will lead to cyclic changes in allele frequency – as <i>p</i> increases in frequency, <i>h</i> will be disfavoured. </p><p>In reality, there will be several genes involved in the relationship between hosts and parasites. In an asexual population of hosts, offspring will only have the different parasitic resistance if a mutation arises. In a sexual population of hosts, however, offspring will have a new combination of parasitic resistance alleles. </p><p>In other words, like <a href="/wiki/Lewis_Carroll" title="Lewis Carroll">Lewis Carroll</a>'s Red Queen, sexual hosts are continually "running" (adapting) to "stay in one place" (resist parasites). </p><p>Evidence for this explanation for the evolution of sex is provided by comparison of the rate of <a href="/wiki/Molecular_evolution" title="Molecular evolution">molecular evolution</a> of genes for <a href="/wiki/Kinase" title="Kinase">kinases</a> and <a href="/wiki/Immunoglobulin" class="mw-redirect" title="Immunoglobulin">immunoglobulins</a> in the <a href="/wiki/Immune_system" title="Immune system">immune system</a> with genes coding other <a href="/wiki/Protein" title="Protein">proteins</a>. The genes coding for immune system proteins evolve considerably faster.<sup id="cite_ref-kuma_23-0" class="reference"><a href="#cite_note-kuma-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-wolfe_24-0" class="reference"><a href="#cite_note-wolfe-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Further evidence for the Red Queen hypothesis was provided by observing long-term dynamics and parasite coevolution in a "mixed" (sexual and asexual) population of snails (<i><a href="/wiki/Potamopyrgus_antipodarum" class="mw-redirect" title="Potamopyrgus antipodarum">Potamopyrgus antipodarum</a></i>). The number of sexuals, the number asexuals, and the rates of parasite infection for both were monitored. It was found that clones that were plentiful at the beginning of the study became more susceptible to parasites over time. As parasite infections increased, the once plentiful clones dwindled dramatically in number. Some clonal types disappeared entirely. Meanwhile, sexual snail populations remained much more stable over time.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>However, Hanley et al.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> studied mite infestations of a <a href="/wiki/Parthenogenesis" title="Parthenogenesis">parthenogenetic</a> gecko species and its two related sexual ancestral species. Contrary to expectation based on the <a href="/wiki/Red_Queen_hypothesis" title="Red Queen hypothesis">Red Queen hypothesis</a>, they found that the prevalence, abundance and mean intensity of mites in sexual geckos was significantly higher than in asexuals sharing the same habitat. </p><p>In 2011, researchers used the microscopic roundworm <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">Caenorhabditis elegans</a></i> as a host and the pathogenic bacteria <i><a href="/wiki/Serratia_marcescens" title="Serratia marcescens">Serratia marcescens</a></i> to generate a host-parasite coevolutionary system in a controlled environment, allowing them to conduct more than 70 evolution experiments testing the Red Queen Hypothesis. They genetically manipulated the <a href="/wiki/Mating_system" title="Mating system">mating system</a> of <i>C. elegans</i>, causing populations to mate either sexually, by self-fertilization, or a mixture of both within the same population. Then they exposed those populations to the <i>S. marcescens</i> parasite. It was found that the self-fertilizing populations of <i>C. elegans</i> were rapidly driven extinct by the coevolving parasites while sex allowed populations to keep pace with their parasites, a result consistent with the Red Queen Hypothesis.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> In natural populations of <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">C. elegans</a></i>, self-fertilization is the predominant mode of reproduction, but infrequent out-crossing events occur at a rate of about 1%.<sup id="cite_ref-pmid16005289_30-0" class="reference"><a href="#cite_note-pmid16005289-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>Critics of the Red Queen hypothesis question whether the constantly changing environment of hosts and parasites is sufficiently common to explain the evolution of sex. In particular, Otto and Nuismer <sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> presented results showing that species interactions (e.g. host vs parasite interactions) typically select against sex. They concluded that, although the Red Queen hypothesis favors sex under certain circumstances, it alone does not account for the ubiquity of sex. Otto and Gerstein <sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> further stated that "it seems doubtful to us that strong selection per gene is sufficiently commonplace for the Red Queen hypothesis to explain the ubiquity of sex". Parker<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> reviewed numerous genetic studies on plant disease resistance and failed to uncover a single example consistent with the assumptions of the Red Queen hypothesis. </p> <div class="mw-heading mw-heading2"><h2 id="Disadvantages_of_sex_and_sexual_reproduction">Disadvantages of sex and sexual reproduction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=7" title="Edit section: Disadvantages of sex and sexual reproduction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The paradox of the existence of sexual reproduction is that though it is ubiquitous in multicellular organisms, there are ostensibly many inherent disadvantages to reproducing sexually when weighed against the relative advantages of alternative forms of reproduction, such as asexual reproduction. Thus, because sexual reproduction abounds in complex multicellular life, there must be some significant benefit(s) to sex and sexual reproduction that compensates for these fundamental disadvantages. </p> <div class="mw-heading mw-heading3"><h3 id="Population_expansion_cost_of_sex"><span class="anchor" id="Two-fold_cost"></span><span class="anchor" id="xspeed"></span>Population expansion cost of sex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=8" title="Edit section: Population expansion cost of sex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Among the most limiting disadvantages to the evolution of sexual reproduction by natural selection is that an asexual population can grow much more rapidly than a sexual one with each generation. </p><p>For example, assume that the entire population of some theoretical species has 100 total organisms consisting of two sexes (i.e. males and females), with 50:50 male-to-female representation, and that only the females of this species can bear offspring. If all capable members of this population procreated once, a total of 50 offspring would be produced (the <i>F<sub>1</sub></i> generation). Contrast this outcome with an asexual species, in which each and every member of an equally sized 100-organism population is capable of bearing young. If all capable members of this asexual population procreated once, a total of 100 offspring would be produced – twice as many as produced by the sexual population in a single generation. </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Evolsex-dia1a.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/82/Evolsex-dia1a.svg/250px-Evolsex-dia1a.svg.png" decoding="async" width="250" height="116" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/82/Evolsex-dia1a.svg/375px-Evolsex-dia1a.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/82/Evolsex-dia1a.svg/500px-Evolsex-dia1a.svg.png 2x" data-file-width="777" data-file-height="361" /></a><figcaption>This diagram illustrates the <i>two-fold cost of sex</i>. If each individual were to contribute to the same number of offspring (two), <i>(a)</i> the sexual population remains the same size each generation, where the <i>(b)</i> asexual population doubles in size each generation.</figcaption></figure> <p>This idea is sometimes referred to as the <b>two-fold cost</b> of sexual reproduction. It was first described mathematically by <a href="/wiki/John_Maynard_Smith" title="John Maynard Smith">John Maynard Smith</a>.<sup id="cite_ref-maynard_34-0" class="reference"><a href="#cite_note-maynard-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> In his manuscript, Smith further speculated on the impact of an asexual mutant arising in a sexual population, which suppresses <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> and allows eggs to develop into offspring genetically identical to the mother by mitotic division.<sup id="cite_ref-1946-_2005_35-0" class="reference"><a href="#cite_note-1946-_2005-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> The mutant-asexual lineage would double its representation in the population each generation, all else being equal. </p><p>Technically the problem above is not one of sexual reproduction but of having a subset of organisms incapable of bearing offspring. Indeed, some multicellular organisms (<a href="/wiki/Isogamy" title="Isogamy">isogamous</a>) engage in sexual reproduction but all members of the species are capable of bearing offspring.<sup id="cite_ref-Hoekstra_36-0" class="reference"><a href="#cite_note-Hoekstra-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> The two-fold reproductive disadvantage assumes that males contribute only genes to their offspring and sexual females spend half their reproductive potential on sons.<sup id="cite_ref-1946-_2005_35-1" class="reference"><a href="#cite_note-1946-_2005-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> Thus, in this formulation, the principal cost of sex is that males and females must successfully <a href="/wiki/Copulation_(zoology)" title="Copulation (zoology)">copulate</a>, which almost always involves expending energy to come together through time and space. Asexual organisms need not expend the energy necessary to find a mate. </p> <div class="mw-heading mw-heading3"><h3 id="Selfish_cytoplasmic_genes">Selfish cytoplasmic genes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=9" title="Edit section: Selfish cytoplasmic genes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-One_source plainlinks metadata ambox ambox-content ambox-one_source" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>relies largely or entirely upon a <a href="/wiki/Wikipedia:Identifying_reliable_sources" class="mw-redirect" title="Wikipedia:Identifying reliable sources">single source</a></b>.<span class="hide-when-compact"> Relevant discussion may be found on the <a href="/wiki/Talk:Evolution_of_sexual_reproduction##" title="Talk:Evolution of sexual reproduction">talk page</a>. Please help <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Evolution_of_sexual_reproduction&action=edit">improve this article</a> by introducing <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">citations</a> to additional sources at this section.</span> <span class="date-container"><i>(<span class="date">August 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>Sexual reproduction implies that chromosomes and alleles segregate and recombine in every generation, but not all genes are transmitted together to the offspring.<sup id="cite_ref-1946-_2005_35-2" class="reference"><a href="#cite_note-1946-_2005-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> There is a chance of spreading mutants that cause unfair transmission at the expense of their non-mutant colleagues. These mutations are referred to as "selfish" because they promote their own spread at the cost of alternative alleles or of the host organism; they include nuclear meiotic drivers and selfish cytoplasmic genes.<sup id="cite_ref-1946-_2005_35-3" class="reference"><a href="#cite_note-1946-_2005-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> <a href="/wiki/Meiotic_drive" title="Meiotic drive">Meiotic drivers</a> are genes that distort meiosis to produce gametes containing themselves more than the 50% of the time expected by chance. A selfish cytoplasmic gene is a gene located in an organelle, plasmid or intracellular parasite that modifies reproduction to cause its own increase at the expense of the cell or organism that carries it.<sup id="cite_ref-1946-_2005_35-4" class="reference"><a href="#cite_note-1946-_2005-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genetic_heritability_cost_of_sex">Genetic heritability cost of sex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=10" title="Edit section: Genetic heritability cost of sex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A sexually reproducing organism only passes on ~50% of its own genetic material to each L2 offspring. This is a consequence of the fact that gametes from sexually reproducing species are <a href="/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a>. Again, however, this is not applicable to all sexual organisms. There are numerous species which are sexual but do not have a genetic-loss problem because they do not produce males or females. Yeast, for example, are <a href="/wiki/Isogamy" title="Isogamy">isogamous</a> sexual organisms which have two <a href="/wiki/Mating_type" title="Mating type">mating types</a> which fuse and recombine their haploid genomes. Both sexes reproduce during the haploid and diploid stages of their life cycle and have a 100% chance of passing their genes into their offspring.<sup id="cite_ref-Hoekstra_36-1" class="reference"><a href="#cite_note-Hoekstra-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (August 2021)">page needed</span></a></i>]</sup> </p><p>Some species avoid the 50% cost of sexual reproduction, although they have "sex" (in the sense of <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a>). In these species (e.g., <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>, <a href="/wiki/Ciliates" class="mw-redirect" title="Ciliates">ciliates</a>, <a href="/wiki/Dinoflagellate" title="Dinoflagellate">dinoflagellates</a> and <a href="/wiki/Diatom" title="Diatom">diatoms</a>), "sex" and reproduction occur separately.<sup id="cite_ref-Ridley_M_2004_p._314_37-0" class="reference"><a href="#cite_note-Ridley_M_2004_p._314-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="DNA_repair_and_complementation">DNA repair and complementation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=11" title="Edit section: DNA repair and complementation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>As discussed in the earlier part of this article, sexual reproduction is conventionally explained as an adaptation for producing genetic variation through allelic recombination. As acknowledged above, however, serious problems with this explanation have led many biologists to conclude that the benefit of sex is a major unsolved problem in evolutionary biology. </p><p>An alternative "<a href="/wiki/Information_theory" title="Information theory">informational</a>" approach to this problem has led to the view that the two fundamental aspects of sex, <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> and <a href="/wiki/Outcrossing" title="Outcrossing">outcrossing</a>, are adaptive responses to the two major sources of "noise" in transmitting genetic information. Genetic noise can occur as either physical damage to the genome (e.g. chemically altered bases of DNA or breaks in the chromosome) or replication errors (mutations).<sup id="cite_ref-dna-repair_39-0" class="reference"><a href="#cite_note-dna-repair-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bernstein1985_13-1" class="reference"><a href="#cite_note-Bernstein1985-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bernstein1986_14-1" class="reference"><a href="#cite_note-Bernstein1986-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> This alternative view is referred to as the repair and complementation hypothesis, to distinguish it from the traditional variation hypothesis. </p><p>The repair and complementation hypothesis assumes that <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> is fundamentally a DNA repair process, and that when it occurs during <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> it is an adaptation for repairing the genomic DNA which is passed on to progeny. Recombinational repair is the only repair process known which can accurately remove double-strand damages in DNA, and such damages are both common in nature and ordinarily lethal if not repaired. For instance, double-strand breaks in DNA occur about 50 times per cell cycle in human cells (see <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">naturally occurring DNA damage</a>). Recombinational repair is prevalent from the simplest viruses to the most complex multicellular eukaryotes. It is effective against many different types of genomic damage, and in particular is highly efficient at overcoming double-strand damages. Studies of the mechanism of meiotic recombination indicate that meiosis is an adaptation for repairing DNA.<sup id="cite_ref-replication-help_40-0" class="reference"><a href="#cite_note-replication-help-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> These considerations form the basis for the first part of the repair and complementation hypothesis. </p><p>In some lines of descent from the earliest organisms, the <a href="/wiki/Ploidy" title="Ploidy">diploid</a> stage of the sexual cycle, which was at first transient, became the predominant stage, because it allowed <a href="/wiki/Complementation_(genetics)" title="Complementation (genetics)">complementation</a> — the masking of deleterious recessive mutations (i.e. <a href="/wiki/Heterosis" title="Heterosis">hybrid vigor or heterosis</a>). <a href="/wiki/Outcrossing" title="Outcrossing">Outcrossing</a>, the second fundamental aspect of sex, is maintained by the advantage of masking mutations and the disadvantage of <a href="/wiki/Inbreeding" title="Inbreeding">inbreeding</a> (mating with a close relative) which allows expression of recessive mutations (commonly observed as <a href="/wiki/Inbreeding_depression" title="Inbreeding depression">inbreeding depression</a>). This is in accord with <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a>,<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> who concluded that the adaptive advantage of sex is hybrid vigor; or as he put it, "the offspring of two individuals, especially if their progenitors have been subjected to very different conditions, have a great advantage in height, weight, constitutional vigor and fertility over the self fertilised offspring from either one of the same parents." </p><p>However, outcrossing may be abandoned in favor of parthenogenesis or selfing (which retain the advantage of meiotic recombinational repair) under conditions in which the costs of mating are very high. For instance, costs of mating are high when individuals are rare in a geographic area, such as when there has been a forest fire and the individuals entering the burned area are the initial ones to arrive. At such times mates are hard to find, and this favors parthenogenic species. </p><p>In the view of the repair and complementation hypothesis, the removal of DNA damage by recombinational repair produces a new, less deleterious form of informational noise, allelic recombination, as a by-product. This lesser informational noise generates genetic variation, viewed by some as the major effect of sex, as discussed in the earlier parts of this article. </p> <div class="mw-heading mw-heading2"><h2 id="Deleterious_mutation_clearance">Deleterious mutation clearance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=12" title="Edit section: Deleterious mutation clearance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Mutation" title="Mutation">Mutations</a> can have many different effects upon an organism. It is generally believed that the majority of non-neutral mutations are deleterious, which means that they will cause a decrease in the organism's overall fitness.<sup id="cite_ref-Griffiths_42-0" class="reference"><a href="#cite_note-Griffiths-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation cites a broad range of pages. Page ranges should be limited to one or two pages when possible. (August 2021)"><span class="nowrap">page range too broad</span></span></a></i>]</sup> If a mutation has a deleterious effect, it will then usually be removed from the population by the process of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>. Sexual reproduction is believed to be more efficient than asexual reproduction in removing those mutations from the genome.<sup id="cite_ref-kondrashov_43-0" class="reference"><a href="#cite_note-kondrashov-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p><p>There are two main hypotheses which explain how sex may act to remove deleterious <a href="/wiki/Gene" title="Gene">genes</a> from the genome. </p> <div class="mw-heading mw-heading3"><h3 id="Evading_harmful_mutation_build-up">Evading harmful mutation build-up</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=13" title="Edit section: Evading harmful mutation build-up"><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/Muller%27s_ratchet" title="Muller's ratchet">Muller's ratchet</a></div> <p>While <a href="/wiki/DNA" title="DNA">DNA</a> is able to recombine to modify <a href="/wiki/Alleles" class="mw-redirect" title="Alleles">alleles</a>, DNA is also susceptible to mutations within the sequence that can affect an organism in a negative manner. Asexual organisms do not have the ability to recombine their genetic information to form new and differing alleles. Once a <a href="/wiki/Mutation" title="Mutation">mutation</a> occurs in the DNA or other genetic carrying sequence, there is no way for the mutation to be removed from the population until another mutation occurs that ultimately deletes the primary mutation. This is rare among organisms. </p><p><a href="/wiki/Hermann_Joseph_Muller" title="Hermann Joseph Muller">Hermann Joseph Muller</a> introduced the idea that mutations build up in asexual reproducing organisms. Muller described this occurrence by comparing the mutations that accumulate as a <a href="/wiki/Muller%27s_ratchet" title="Muller's ratchet">ratchet</a>. Each mutation that arises in asexually reproducing organisms turns the ratchet once. The ratchet is unable to be rotated backwards, only forwards. The next mutation that occurs turns the ratchet once more. Additional mutations in a population continually turn the ratchet and the mutations, mostly deleterious, continually accumulate without recombination.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> These mutations are passed onto the next generation because the offspring are exact genetic <a href="/wiki/Cloning" title="Cloning">clones</a> of their parents. The genetic load of organisms and their populations will increase due to the addition of multiple deleterious mutations and decrease the overall reproductive success and fitness. </p><p>For sexually reproducing populations, studies have shown that single-celled bottlenecks are beneficial for resisting mutation build-up<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2022)">citation needed</span></a></i>]</sup>. Passaging a population through a single-celled bottleneck involves the fertilization event occurring with <a href="/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a> sets of DNA, forming one fertilized cell. For example, humans undergo a single-celled bottleneck in that the haploid sperm fertilizes the haploid egg, forming the diploid <a href="/wiki/Zygote" title="Zygote">zygote</a>, which is unicellular. This passage through a single cell is beneficial in that it lowers the chance of mutations from being passed on through multiple individuals. Instead, the mutation is only passed onto one individual.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> Further studies using <i><a href="/wiki/Dictyostelium_discoideum" title="Dictyostelium discoideum">Dictyostelium discoideum</a></i> suggest that this unicellular initial stage is important for resisting mutations due to the importance of high relatedness. Highly related individuals are more closely related, and more clonal, whereas less related individuals are less so, increasing the likelihood that an individual in a population of low relatedness may have a detrimental mutation. Highly related populations also tend to thrive better than lowly related because the cost of sacrificing an individual is greatly offset by the benefit gained by its relatives and in turn, its genes, according to <a href="/wiki/Kin_selection" title="Kin selection">kin selection</a>. The studies with <i>D. discoideum</i> showed that conditions of high relatedness resisted mutant individuals more effectively than those of low relatedness, suggesting the importance of high relatedness to resist mutations from proliferating.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Removal_of_deleterious_genes">Removal of deleterious genes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=14" title="Edit section: Removal of deleterious genes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Synergistic_versus_antagonistic_epistasis.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Synergistic_versus_antagonistic_epistasis.svg/200px-Synergistic_versus_antagonistic_epistasis.svg.png" decoding="async" width="200" height="157" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Synergistic_versus_antagonistic_epistasis.svg/300px-Synergistic_versus_antagonistic_epistasis.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Synergistic_versus_antagonistic_epistasis.svg/400px-Synergistic_versus_antagonistic_epistasis.svg.png 2x" data-file-width="724" data-file-height="570" /></a><figcaption>Diagram illustrating different relationships between numbers of mutations and fitness. Kondrashov's model requires <i>synergistic epistasis</i>, which is represented by the red line<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> – each subsequent mutation has a disproportionately large effect on the organism's fitness.</figcaption></figure> <p>This hypothesis was proposed by <a href="/wiki/Alexey_Kondrashov" title="Alexey Kondrashov">Alexey Kondrashov</a>, and is sometimes known as the <i>deterministic mutation hypothesis</i>.<sup id="cite_ref-kondrashov_43-1" class="reference"><a href="#cite_note-kondrashov-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> It assumes that the majority of deleterious mutations are only slightly deleterious, and affect the individual such that the introduction of each additional mutation has an increasingly large effect on the fitness of the organism. This relationship between number of mutations and fitness is known as <a href="/wiki/Epistasis" title="Epistasis"><i>synergistic epistasis</i></a>. </p><p>By way of <a href="/wiki/Analogy" title="Analogy">analogy</a>, think of a <a href="/wiki/Automobile" class="mw-redirect" title="Automobile">car</a> with several minor faults. Each is not sufficient alone to prevent the car from running, but in combination, the faults combine to prevent the car from functioning. </p><p>Similarly, an organism may be able to cope with a few defects, but the presence of many mutations could overwhelm its backup mechanisms. </p><p>Kondrashov argues that the slightly deleterious nature of mutations means that the population will tend to be composed of individuals with a small number of mutations. Sex will act to <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombine</a> these genotypes, creating some individuals with fewer deleterious mutations, and some with more. Because there is a major selective disadvantage to individuals with more mutations, these individuals die out. In essence, sex compartmentalises the deleterious mutations. </p><p>There has been much criticism of Kondrashov's theory, since it relies on two key restrictive conditions. The first requires that the rate of deleterious mutation should exceed one per genome per generation in order to provide a substantial advantage for sex. While there is some empirical evidence for it (for example in <a href="/wiki/Drosophila" title="Drosophila">Drosophila</a><sup id="cite_ref-Drosophila_49-0" class="reference"><a href="#cite_note-Drosophila-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/E._coli" class="mw-redirect" title="E. coli">E. coli</a><sup id="cite_ref-E._coli_50-0" class="reference"><a href="#cite_note-E._coli-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup>), there is also strong evidence against it. Thus, for instance, for the sexual species <i>Saccharomyces cerevisiae</i> (yeast) and <i>Neurospora crassa</i> (fungus), the mutation rate per genome per replication are 0.0027 and 0.0030 respectively. For the nematode worm <i>Caenorhabditis elegans</i>, the mutation rate per effective genome per sexual generation is 0.036.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> Secondly, there should be strong interactions among loci (synergistic epistasis), a mutation-fitness relation for which there is only limited evidence.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> Conversely, there is also the same amount of evidence that mutations show no epistasis (purely additive model) or antagonistic interactions (each additional mutation has a disproportionally <i>small</i> effect). </p> <div class="mw-heading mw-heading2"><h2 id="Other_explanations">Other explanations</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=15" title="Edit section: Other explanations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Geodakyan's_evolutionary_theory_of_sex"><span id="Geodakyan.27s_evolutionary_theory_of_sex"></span>Geodakyan's evolutionary theory of sex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=16" title="Edit section: Geodakyan's evolutionary theory of sex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Geodakyan suggested that sexual dimorphism provides a partitioning of a species' phenotypes into at least two functional partitions: a female partition that secures beneficial features of the species and a male partition that emerged in species with more variable and unpredictable environments. The male partition is suggested to be an "experimental" part of the species that allows the species to expand their ecological niche, and to have alternative configurations. This theory underlines the higher variability and higher mortality in males, in comparison to females. This functional partitioning also explains the higher susceptibility to disease in males, in comparison to females and therefore includes the idea of "protection against parasites" as another functionality of male sex. Geodakyan's evolutionary theory of sex was developed in Russia in 1960–1980 and was not known to the West till the era of the Internet. Trofimova, who analysed psychological sex differences, hypothesised that the male sex might also provide a "redundancy pruning" function.<sup id="cite_ref-Trofimova_54-0" class="reference"><a href="#cite_note-Trofimova-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Speed_of_evolution">Speed of evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=17" title="Edit section: Speed of evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ilan Eshel suggested that sex prevents rapid evolution. He suggests that <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a> breaks up favourable gene combinations more often than it creates them, and sex is maintained because it ensures selection is longer-term than in asexual populations – so the population is less affected by short-term changes.<sup id="cite_ref-Birdsell_17-2" class="reference"><a href="#cite_note-Birdsell-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 85–86">: 85–86 </span></sup><sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> This explanation is not widely accepted, as its assumptions are very restrictive. </p><p>It has recently been shown in experiments with <i><a href="/wiki/Chlamydomonas" title="Chlamydomonas">Chlamydomonas</a></i> algae that sex can remove the speed limit<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (July 2013)">clarification needed</span></a></i>]</sup> on evolution.<sup id="cite_ref-speed_56-0" class="reference"><a href="#cite_note-speed-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </p><p>An information theoretic analysis using a simplified but useful model shows that in asexual reproduction, the information gain per generation of a species is limited to 1 bit per generation, while in sexual reproduction, the information gain is bounded by <span class="nowrap"><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\sqrt {G}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <msqrt> <mi>G</mi> </msqrt> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\sqrt {G}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/7110a12895f8c41e8d18284799da998cbc02037b" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:3.763ex; height:3.009ex;" alt="{\displaystyle {\sqrt {G}}}"></span>,</span> where <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle G}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>G</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle G}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/f5f3c8921a3b352de45446a6789b104458c9f90b" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:1.827ex; height:2.176ex;" alt="{\displaystyle G}"></span> is the size of the genome in bits.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Libertine_bubble_theory">Libertine bubble theory</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=18" title="Edit section: Libertine bubble theory"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The evolution of sex can alternatively be described as a kind of <a href="/wiki/Gene" title="Gene">gene</a> exchange that is independent from reproduction.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> According to the <a href="/wiki/Thierry_Lod%C3%A9" title="Thierry Lodé">Thierry Lodé</a>'s "libertine bubble theory", sex originated from an archaic gene transfer process among <a href="/wiki/Prebiotic_(chemistry)" class="mw-redirect" title="Prebiotic (chemistry)">prebiotic</a> bubbles.<sup id="cite_ref-sexisnot_59-0" class="reference"><a href="#cite_note-sexisnot-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> The contact among the pre-biotic bubbles could, through simple food or parasitic reactions, promote the transfer of genetic material from one bubble to another. That interactions between two organisms be in balance appear to be a sufficient condition to make these interactions evolutionarily efficient, i.e. to select bubbles that tolerate these interactions ("libertine" bubbles) through a blind evolutionary process of self-reinforcing gene correlations and compatibility.<sup id="cite_ref-LODÉ_T_2012_61-0" class="reference"><a href="#cite_note-LODÉ_T_2012-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p><p>The "libertine bubble theory" proposes that <a href="/wiki/Meiosis" title="Meiosis">meiotic</a> sex evolved in proto-<a href="/wiki/Eukaryotes" class="mw-redirect" title="Eukaryotes">eukaryotes</a> to solve a problem that bacteria did not have, namely a large amount of <a href="/wiki/DNA" title="DNA">DNA</a> material, occurring in an archaic step of <a href="/wiki/Proto-cell" class="mw-redirect" title="Proto-cell">proto-cell</a> formation and genetic exchanges. So that, rather than providing selective advantages through reproduction, sex could be thought of as a series of separate events which combines step-by-step some very weak benefits of <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a>, meiosis, gametogenesis and <a href="/wiki/Syngamy" class="mw-redirect" title="Syngamy">syngamy</a>.<sup id="cite_ref-ReferenceA_62-0" class="reference"><a href="#cite_note-ReferenceA-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Therefore, current sexual species could be descendants of primitive organisms that practiced more stable exchanges in the long term, while asexual species have emerged, much more recently in evolutionary history, from the conflict of interest resulting from <a href="/wiki/Anisogamy" title="Anisogamy">anisogamy</a>.<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="how? (December 2016)">clarification needed</span></a></i>]</sup> </p><p><b><big>Parasites and Muller's ratchet</big></b> </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-Unreferenced_section plainlinks metadata ambox ambox-content ambox-Unreferenced" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>does not <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">cite</a> any <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">sources</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Evolution_of_sexual_reproduction" title="Special:EditPage/Evolution of sexual reproduction">improve this section</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and <a href="/wiki/Wikipedia:Verifiability#Burden_of_evidence" title="Wikipedia:Verifiability">removed</a>.</span> <span class="date-container"><i>(<span class="date">August 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>R. Stephen Howard and <a href="/w/index.php?title=Curtis_Lively&action=edit&redlink=1" class="new" title="Curtis Lively (page does not exist)">Curtis Lively</a> were the first to suggest that the combined effects of parasitism and <a href="/wiki/Mutation" title="Mutation">mutation</a> accumulation can lead to an increased advantage to sex under conditions not otherwise predicted (Nature, 1994). Using computer simulations, they showed that when the two mechanisms act simultaneously the advantage to sex over <a href="/wiki/Asexual_reproduction" title="Asexual reproduction">asexual reproduction</a> is larger than for either factor operating alone. </p> <div class="mw-heading mw-heading2"><h2 id="Origin_of_sexual_reproduction">Origin of sexual reproduction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=19" title="Edit section: Origin of sexual reproduction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table id="Container" role="presentation" class="nomobile toccolours searchaux" style="text-align:left;padding:0 0.5em;border-style:solid;float:right;clear:right;margin:0.3em 0 0.8em 1.4em;overflow:hidden;min-width:29.95em"><tbody><tr><td colspan="3" id="Title" style="padding:1em 1em 0 1em"><div style="background-color:#77bb77;padding:0 0.2em 0 0.2em;font-weight:bold;text-align:center"><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life"><span style="color:#ffffff;">Life timeline</span></a></div></td></tr><tr><td colspan="3" id="Navbox" style="padding:0;margin:0 1em 0 0;text-align:right"><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"><span>This box: </span><ul><li class="nv-view"><a href="/wiki/Template:Life_timeline" title="Template:Life timeline"><span title="View this template">view</span></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Life_timeline" title="Template talk:Life timeline"><span title="Discuss this template">talk</span></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Life_timeline" title="Special:EditPage/Template:Life timeline"><span title="Edit this template">edit</span></a></li></ul></div></td></tr><tr style="vertical-align:top"><td id="Scale" style="padding:0;margin:0.7em 0 0.7em 0;width:4.2em;position:relative;float:left;font-size:100%;height:50em;border-right:1px solid #242020"><div style="float:right;position:absolute;right:-1px;top:49.56em;transform:translateY(-50%)"><span style="font-size:90%">−4500 —</span></div><div style="float:right;position:absolute;right:-1px;top:48.18em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:46.81em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:45.43em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:44.05em;transform:translateY(-50%)"><span style="font-size:90%">−4000 —</span></div><div 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style="float:right;position:absolute;right:-1px;top:23.40em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:22.03em;transform:translateY(-50%)"><span style="font-size:90%">−2000 —</span></div><div style="float:right;position:absolute;right:-1px;top:20.65em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:19.27em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:17.90em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:16.52em;transform:translateY(-50%)"><span style="font-size:90%">−1500 —</span></div><div style="float:right;position:absolute;right:-1px;top:15.14em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:13.77em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:12.39em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:11.01em;transform:translateY(-50%)"><span style="font-size:90%">−1000 —</span></div><div style="float:right;position:absolute;right:-1px;top:9.64em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:8.26em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:6.88em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:5.51em;transform:translateY(-50%)"><span style="font-size:90%">−500 —</span></div><div style="float:right;position:absolute;right:-1px;top:4.13em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:2.75em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:1.38em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:0.00em;transform:translateY(-50%)"><span style="font-size:90%">0 —</span></div></td><td id="Timeline" class="toccolours notheme" style="padding:0px;margin:0.7em 0 0.7em 0;position:relative;font-size:100%;width:13em;height:50em;float:left;border:none"><div class="notheme" style="font-size:100%;background-color:#e5cc7f;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:50.000em;left:0.910em;width:12.090em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-25.000em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#aebfd1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:44.053em;height:4.537em;left:1.040em;width:11.960em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.269em;transform:translateY(-50%);z-index:5"><a href="/wiki/Origin_of_water_on_Earth" title="Origin of water on Earth">Water</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#aebfd1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:44.053em;left:1.040em;width:0.130em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-22.026em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#b3d9ff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:38.546em;height:5.507em;left:1.170em;width:11.830em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.753em;transform:translateY(-50%);z-index:5"><a href="/wiki/Unicellular_organism" title="Unicellular organism">Single-celled life</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#b3d9ff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:38.546em;left:1.170em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-19.273em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#99ffff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:23.128em;height:15.419em;left:1.430em;width:11.570em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-7.709em;transform:translateY(-50%);z-index:5"><a href="/wiki/Evolution_of_photosynthesis" title="Evolution of photosynthesis">Photosynthesis</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#99ffff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:23.128em;left:1.430em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-11.564em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#b3ffea;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:17.621em;height:5.507em;left:1.690em;width:11.310em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.753em;transform:translateY(-50%);z-index:5"><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryotes</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#b3ffea;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:17.621em;left:1.690em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-8.811em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ccffe7;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:5.958em;height:11.663em;left:1.950em;width:11.050em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-5.831em;transform:translateY(-50%);z-index:5"><a href="/wiki/Multicellular_organism" title="Multicellular organism">Multicellular life</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ccffe7;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:5.958em;left:1.950em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.979em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#77dd00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:1.982em;height:7.379em;left:2.210em;width:1.170em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-3.689em;transform:translateY(-50%);z-index:5"><a href="/wiki/Plant" title="Plant"><span style="display:block; text-align:center; line-height: 0.85em;">P<br />l<br />a<br />n<br />t<br />s</span></a></div></div></div><div class="notheme" style="font-size:100%;background-color:#77dd00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:1.982em;left:2.210em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.991em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ccbbaa;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:2.203em;height:3.756em;left:3.380em;width:9.620em"><div style="font-size:100%;position:relative;top:0.5em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-1.878em;transform:translateY(-50%);z-index:5"><a href="/wiki/Arthropod" title="Arthropod">Arthropods</a> <a href="/wiki/Mollusca" title="Mollusca">Molluscs</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ffff00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:1.982em;left:2.470em;width:3.510em"><div style="font-size:100%;position:relative;top:-0.53em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-0.991em;transform:translateY(-50%);z-index:5"><a href="/wiki/Flowering_plant#Evolutionary_history" title="Flowering plant">Flowers</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#e6b880;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.721em;height:2.583em;left:3.380em;width:5.070em"><div style="font-size:100%;position:relative;top:0.5em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-1.291em;transform:translateY(-50%);z-index:5"><a href="/wiki/Dinosaur" title="Dinosaur">Dinosaurs</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ffcc00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:2.203em;left:8.580em;width:4.420em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.101em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ffcc00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.606em;height:1.597em;left:8.450em;width:4.290em"><div style="font-size:100%;position:relative;top:0.15em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-0.798em;transform:translateY(-50%);z-index:5"><a href="/wiki/Mammal" title="Mammal">Mammals</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#e6b880;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:1.333em;left:5.980em;width:2.600em"><div style="font-size:100%;position:relative;top:-0.225em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-0.666em;transform:translateY(-50%);z-index:5"><a href="/wiki/Bird" title="Bird">Birds</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ffbbbb;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:0.606em;left:8.710em;width:4.030em"><div style="font-size:100%;position:relative;top:0.1em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-0.303em;transform:translateY(-50%);z-index:5"><a href="/wiki/Primate" title="Primate">Primates</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#993c00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:44.053em;height:5.947em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:0em;left:-0.05em"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-2.974em;transform:translateY(-50%);z-index:5"><span style="display:block; text-align:center; line-height: 0.80em;"><a href="/wiki/Hadean" title="Hadean"><span style="color:#ffffff;">H<br />a<br />d<br />e<br />a<br />n</span></a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#936a00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:27.533em;height:16.520em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:-1.2em;left:-0.05em"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-8.260em;transform:translateY(-50%);z-index:5"><span style="display:block; text-align:center; line-height: 0.85em;"><a href="/wiki/Archean" title="Archean"><span style="color:#ffffff;"><br /><br /><br />A<br />r<br />c<br />h<br />e<br />a<br />n</span></a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#f1c309;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:5.958em;height:21.575em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-10.787em;transform:translateY(-50%);z-index:5"><span style="display:block; text-align:center; line-height: 0.85em;"><a href="/wiki/Proterozoic" title="Proterozoic"><br /><br /><br /><br /><br /><br /><br />P<br />r<br />o<br />t<br />e<br />r<br />o<br />z<br />o<br />i<br />c</a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#c1e0c1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:5.958em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:2em;left:0em"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-2.979em;transform:translateY(-50%);z-index:5"><span style="display:block; text-align:center; line-height: 0.85em;"><a href="/wiki/Phanerozoic" title="Phanerozoic">P<br />h<br />a<br />n<br />e<br />r<br />o<br />z<br />o<br />i<br />c</a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ccbbaa;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:8.260em;left:12.714em;width:0.286em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-4.130em;transform:translateY(-50%);z-index:5"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ccbbaa;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;top:0.000em;height:2.203em;left:12.714em;width:0.286em"><div style="font-size:100%;position:relative;top:0em;left:0em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.101em;transform:translateY(-50%);z-index:5"> </div></div></div></td><td id="Annotations" style="padding:0;margin:0.7em 0 0.7em 0;float:left;position:relative;width:11.05em;height:50em"><table role="presentation" style="position:absolute;z-index:15;top:50.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Age_of_Earth" title="Age of Earth">Earth formed</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:48.568em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Origin_of_water_on_Earth" title="Origin of water on Earth">Earliest water</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:47.907em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">LUCA</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:44.053em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest fossils</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:42.952em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Late_Heavy_Bombardment" title="Late Heavy Bombardment">LHB meteorites</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:38.546em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Geological_history_of_oxygen" title="Geological history of oxygen">Earliest oxygen</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:31.278em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Pongola_glaciation" title="Pongola glaciation">Pongola glaciation</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:25.330em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Great_Oxidation_Event" title="Great Oxidation Event">Atmospheric oxygen</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:24.780em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Huronian_glaciation" title="Huronian glaciation">Huronian glaciation</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:22.026em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a class="mw-selflink-fragment" href="#Origin_of_sexual_reproduction">Sexual reproduction</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:17.621em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Francevillian_biota" title="Francevillian biota">Earliest multicellular life</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:16.520em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Earliest fungi</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:9.361em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.25em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Earliest plants</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:8.260em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.4em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Caveasphaera" title="Caveasphaera">Earliest animals</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:7.489em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Cryogenian" title="Cryogenian">Cryogenian ice age</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:6.993em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Ediacaran_biota" title="Ediacaran biota">Ediacaran biota</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:5.958em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.15em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Cambrian_explosion" title="Cambrian explosion">Cambrian explosion</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:4.956em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Hirnantian_glaciation" title="Hirnantian glaciation">Hirnantian glaciation</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:4.075em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Tetrapod" title="Tetrapod">Earliest tetrapods</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:3.304em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Late_Paleozoic_icehouse" title="Late Paleozoic icehouse">Karoo ice age</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.275em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.3em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Ape" title="Ape">Earliest apes</a> / <a href="/wiki/Human" title="Human">humans</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.018em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.4em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0"><a href="/wiki/Quaternary_glaciation" title="Quaternary glaciation">Quaternary ice age</a>*</div></div></td></tr></tbody></table></td></tr><tr><td id="Caption" colspan="3" style="padding:0;margin:0 0.2em 0.7em 0.2em"><div id="Caption" class="toccolours" style="margin:0 0.5em;border-style:none;clear:both;text-align:center;width:26.8em"><div style="float:left;font-size:85%;">(<a href="/wiki/Myr" class="mw-redirect" title="Myr">million years ago</a>)</div><div style="float:right;font-size:95%;">*<i><a href="/wiki/Timeline_of_glaciation" title="Timeline of glaciation">Ice Ages</a></i></div></div></td></tr></tbody></table> <p>Many <a href="/wiki/Protist" title="Protist">protists</a> reproduce sexually, as do many multicellular <a href="/wiki/Plant" title="Plant">plants</a>, <a href="/wiki/Animal" title="Animal">animals</a>, and <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a>. In the eukaryotic fossil record, sexual reproduction first appeared about 2.0 billion years ago in the <a href="/wiki/Proterozoic" title="Proterozoic">Proterozoic Eon</a>,<sup id="cite_ref-NAT-2008_63-0" class="reference"><a href="#cite_note-NAT-2008-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-SCI-20090605_64-0" class="reference"><a href="#cite_note-SCI-20090605-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> although a later date, 1.2 billion years ago, has also been presented.<sup id="cite_ref-PB-2000_65-0" class="reference"><a href="#cite_note-PB-2000-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-BBC-20160704_66-0" class="reference"><a href="#cite_note-BBC-20160704-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Nonetheless, all sexually reproducing <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> organisms likely derive from a single-celled common ancestor.<sup id="cite_ref-Letunic_I_and_Bork_P_1-1" class="reference"><a href="#cite_note-Letunic_I_and_Bork_P-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bernstein2010_67-0" class="reference"><a href="#cite_note-Bernstein2010-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-sexisnot_59-1" class="reference"><a href="#cite_note-sexisnot-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> It is probable that the evolution of sex was an integral part of the evolution of the first eukaryotic cell.<sup id="cite_ref-Ploompuu1999_68-0" class="reference"><a href="#cite_note-Ploompuu1999-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid29436502_69-0" class="reference"><a href="#cite_note-pmid29436502-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> There are a few species which have secondarily lost this feature, such as <a href="/wiki/Bdelloidea" title="Bdelloidea">Bdelloidea</a> and some <a href="/wiki/Parthenocarpic" class="mw-redirect" title="Parthenocarpic">parthenocarpic</a> plants. </p> <div class="mw-heading mw-heading3"><h3 id="Diploidy">Diploidy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=20" title="Edit section: Diploidy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Organisms need to replicate their genetic material in an efficient and reliable manner. The necessity to repair genetic damage is one of the leading theories explaining the origin of sexual reproduction. <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">Diploid</a> individuals can repair a damaged section of their DNA via <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombination</a>, since there are two copies of the gene in the cell and if one copy is <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">damaged</a>, the other copy is unlikely to be damaged at the same site. </p><p>A harmful damage in a <a href="/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a> individual, on the other hand, is more likely to become fixed (i.e. permanent), since any <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> mechanism would have no source from which to recover the original undamaged sequence.<sup id="cite_ref-dna-repair_39-1" class="reference"><a href="#cite_note-dna-repair-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> The most primitive form of sex may have been one organism with damaged DNA replicating an undamaged strand from a similar organism in order to repair itself.<sup id="cite_ref-tatiana_70-0" class="reference"><a href="#cite_note-tatiana-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Meiosis">Meiosis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=21" title="Edit section: Meiosis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sexual reproduction appears to have arisen very early in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> evolution, implying that the essential features of <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> were already present in the <a href="/wiki/Last_eukaryotic_common_ancestor" class="mw-redirect" title="Last eukaryotic common ancestor">last eukaryotic common ancestor</a>.<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bernstein2010_67-1" class="reference"><a href="#cite_note-Bernstein2010-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-BernsteinIntech_72-0" class="reference"><a href="#cite_note-BernsteinIntech-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> In extant organisms, proteins with central functions in meiosis are similar to key proteins in <a href="/wiki/Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">natural transformation</a> in bacteria and DNA transfer in <a href="/wiki/Archaea" title="Archaea">archaea</a>.<sup id="cite_ref-BernsteinIntech_72-1" class="reference"><a href="#cite_note-BernsteinIntech-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bernsteinarchaea_73-0" class="reference"><a href="#cite_note-Bernsteinarchaea-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> For example, <a href="/wiki/RecA" title="RecA">recA</a> recombinase, that catalyses the key functions of DNA <a href="/wiki/Homology_(biology)" title="Homology (biology)">homology</a> search and strand exchange in the bacterial sexual process of transformation, has <a href="/wiki/Orthologs" class="mw-redirect" title="Orthologs">orthologs</a> in eukaryotes that perform similar functions in meiotic recombination<sup id="cite_ref-BernsteinIntech_72-2" class="reference"><a href="#cite_note-BernsteinIntech-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> </p><p>Natural transformation in bacteria, DNA transfer in <a href="/wiki/Archaea" title="Archaea">archaea</a>, and <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> in eukaryotic microorganisms are induced by stressful circumstances such as overcrowding, resource depletion, and DNA damaging conditions.<sup id="cite_ref-LODÉ_T_2012_61-1" class="reference"><a href="#cite_note-LODÉ_T_2012-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-BernsteinIntech_72-3" class="reference"><a href="#cite_note-BernsteinIntech-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bernsteinarchaea_73-1" class="reference"><a href="#cite_note-Bernsteinarchaea-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> This suggests that these sexual processes are adaptations for dealing with stress, particularly stress that causes DNA damage. In bacteria, these stresses induce an altered physiologic state, termed competence, that allows active take-up of DNA from a donor bacterium and the integration of this DNA into the recipient genome (see <a href="/wiki/Natural_competence" title="Natural competence">Natural competence</a>) allowing recombinational repair of the recipients' damaged DNA.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> </p><p>If environmental stresses leading to DNA damage were a persistent challenge to the survival of early microorganisms, then selection would likely have been continuous through the prokaryote to eukaryote transition,<sup id="cite_ref-ReferenceA_62-1" class="reference"><a href="#cite_note-ReferenceA-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-BernsteinIntech_72-4" class="reference"><a href="#cite_note-BernsteinIntech-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> and adaptative adjustments would have followed a course in which bacterial transformation or archaeal DNA transfer naturally gave rise to sexual reproduction in eukaryotes. </p> <div class="mw-heading mw-heading3"><h3 id="Virus-like_RNA-based_origin">Virus-like RNA-based origin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=22" title="Edit section: Virus-like RNA-based origin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sex might also have been present even earlier, in the hypothesized <a href="/wiki/RNA_world" title="RNA world">RNA world</a> that preceded DNA cellular life forms.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> One proposed origin of sex in the RNA world was based on the type of sexual interaction that is known to occur in extant single-stranded segmented RNA viruses, such as influenza virus, and in extant double-stranded segmented RNA viruses such as reovirus.<sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p><p>Exposure to conditions that cause RNA damage could have led to blockage of replication and death of these early RNA life forms. Sex would have allowed re-assortment of segments between two individuals with damaged RNA, permitting undamaged combinations of RNA segments to come together, thus allowing survival. Such a regeneration phenomenon, known as multiplicity reactivation, occurs in the influenza virus<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> and reovirus.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Parasitic_DNA_elements">Parasitic DNA elements</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=23" title="Edit section: Parasitic DNA elements"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Another theory is that sexual reproduction originated from <a href="/wiki/Selfish_DNA" class="mw-redirect" title="Selfish DNA">selfish parasitic genetic elements</a> that exchange genetic material (that is: copies of their own genome) for their transmission and propagation. In some organisms, sexual reproduction has been shown to enhance the spread of parasitic genetic elements (e.g. yeast, filamentous fungi).<sup id="cite_ref-hickey_79-0" class="reference"><a href="#cite_note-hickey-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Bacterial_conjugation" title="Bacterial conjugation">Bacterial conjugation</a> is a form of genetic exchange that some sources describe as "sex", but technically is not a form of reproduction, even though it is a form of <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>. However, it does support the "selfish gene" part theory, since the gene itself is propagated through the <a href="/wiki/F-plasmid" title="F-plasmid">F-plasmid</a>.<sup id="cite_ref-tatiana_70-1" class="reference"><a href="#cite_note-tatiana-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> </p><p>A similar origin of sexual reproduction is proposed to have evolved in ancient <a href="/wiki/Haloarchaea" title="Haloarchaea">haloarchaea</a> as a combination of two independent processes: <a href="/wiki/Jumping_genes" class="mw-redirect" title="Jumping genes">jumping genes</a> and <a href="/wiki/Plasmid" title="Plasmid">plasmid</a> swapping.<sup id="cite_ref-80" class="reference"><a href="#cite_note-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Partial_predation">Partial predation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=24" title="Edit section: Partial predation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A third theory is that sex evolved as a form of <a href="/wiki/Cannibalism" title="Cannibalism">cannibalism</a>: One primitive organism ate another one, but instead of completely digesting it, some of the eaten organism's DNA was incorporated into the DNA of the eater.<sup id="cite_ref-tatiana_70-2" class="reference"><a href="#cite_note-tatiana-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Ploompuu1999_68-1" class="reference"><a href="#cite_note-Ploompuu1999-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Vaccination-like_process">Vaccination-like process</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=25" title="Edit section: Vaccination-like process"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-One_source plainlinks metadata ambox ambox-content ambox-one_source" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>relies largely or entirely upon a <a href="/wiki/Wikipedia:Identifying_reliable_sources" class="mw-redirect" title="Wikipedia:Identifying reliable sources">single source</a></b>.<span class="hide-when-compact"> Relevant discussion may be found on the <a href="/wiki/Talk:Evolution_of_sexual_reproduction##" title="Talk:Evolution of sexual reproduction">talk page</a>. Please help <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Evolution_of_sexual_reproduction&action=edit">improve this article</a> by introducing <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">citations</a> to additional sources at this section.</span> <span class="date-container"><i>(<span class="date">August 2021</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>Sex may also be derived from another prokaryotic process. A comprehensive theory called "origin of sex as vaccination" proposes that eukaryan sex-as-<a href="/wiki/Fertilisation" title="Fertilisation">syngamy</a> (fusion sex) arose from prokaryan unilateral sex-as-infection, when infected hosts began swapping nuclearised genomes containing coevolved, vertically transmitted symbionts that provided protection against horizontal superinfection by other, more virulent symbionts. </p><p>Consequently, sex-as-meiosis (fission sex) would evolve as a host strategy for uncoupling from (and thereby render impotent) the acquired symbiotic/parasitic genes.<sup id="cite_ref-sterrer_81-0" class="reference"><a href="#cite_note-sterrer-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mechanistic_origin_of_sexual_reproduction">Mechanistic origin of sexual reproduction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=26" title="Edit section: Mechanistic origin of sexual reproduction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>While theories positing fitness benefits that led to the origin of sex are often problematic,<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (July 2018)">citation needed</span></a></i>]</sup> several theories addressing the emergence of the mechanisms of sexual reproduction have been proposed. </p> <div class="mw-heading mw-heading3"><h3 id="Viral_eukaryogenesis">Viral eukaryogenesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=27" title="Edit section: Viral eukaryogenesis"><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/Viral_eukaryogenesis" title="Viral eukaryogenesis">Viral eukaryogenesis</a></div> <p>The viral eukaryogenesis (VE) theory proposes that eukaryotic cells arose from a combination of a <a href="/wiki/Lysogenic" class="mw-redirect" title="Lysogenic">lysogenic</a> virus, an <a href="/wiki/Archaea" title="Archaea">archaean</a>, and a <a href="/wiki/Bacterium" class="mw-redirect" title="Bacterium">bacterium</a>. This model suggests that the nucleus originated when the lysogenic virus incorporated genetic material from the archaean and the bacterium and took over the role of information storage for the amalgam. The archaeal host transferred much of its functional genome to the virus during the evolution of cytoplasm, but retained the function of gene translation and general metabolism. The bacterium transferred most of its functional genome to the virus as it transitioned into a <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondrion</a>.<sup id="cite_ref-bell-2001_82-0" class="reference"><a href="#cite_note-bell-2001-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> </p><p>For these transformations to lead to the eukaryotic cell cycle, the VE hypothesis specifies a pox-like virus as the lysogenic virus. A pox-like virus is a likely ancestor because of its fundamental similarities with eukaryotic nuclei. These include a double stranded DNA genome, a linear chromosome with short <a href="/wiki/Telomere" title="Telomere">telomeric</a> repeats, a complex membrane bound capsid, the ability to produce capped mRNA, and the ability to export the capped mRNA across the viral membrane into the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>. The presence of a lysogenic pox-like virus ancestor explains the development of meiotic division, an essential component of sexual reproduction.<sup id="cite_ref-bell-2006_83-0" class="reference"><a href="#cite_note-bell-2006-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Meiotic" class="mw-redirect" title="Meiotic">Meiotic</a> division in the VE hypothesis arose because of the evolutionary pressures placed on the lysogenic virus as a result of its inability to enter into the <a href="/wiki/Lytic_cycle" title="Lytic cycle">lytic cycle</a>. This selective pressure resulted in the development of processes allowing the viruses to spread horizontally throughout the population. The outcome of this selection was cell-to-cell fusion. (This is distinct from the conjugation methods used by bacterial plasmids under evolutionary pressure, with important consequences.)<sup id="cite_ref-bell-2001_82-1" class="reference"><a href="#cite_note-bell-2001-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> The possibility of this kind of fusion is supported by the presence of fusion proteins in the envelopes of the pox viruses that allow them to fuse with host membranes. These proteins could have been transferred to the cell membrane during viral reproduction, enabling cell-to-cell fusion between the virus host and an uninfected cell. The theory proposes meiosis originated from the fusion between two cells infected with related but different viruses which recognised each other as uninfected. After the fusion of the two cells, incompatibilities between the two viruses result in a meiotic-like cell division.<sup id="cite_ref-bell-2006_83-1" class="reference"><a href="#cite_note-bell-2006-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </p><p>The two viruses established in the cell would initiate replication in response to signals from the host cell. A mitosis-like cell cycle would proceed until the viral membranes dissolved, at which point linear chromosomes would be bound together with centromeres. The homologous nature of the two viral centromeres would incite the grouping of both sets into tetrads. It is speculated that this grouping may be the origin of crossing over, characteristic of the first division in modern meiosis. The partitioning apparatus of the mitotic-like cell cycle the cells used to replicate independently would then pull each set of chromosomes to one side of the cell, still bound by centromeres. These centromeres would prevent their replication in subsequent division, resulting in four daughter cells with one copy of one of the two original pox-like viruses. The process resulting from combination of two similar pox viruses within the same host closely mimics meiosis.<sup id="cite_ref-bell-2006_83-2" class="reference"><a href="#cite_note-bell-2006-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Neomuran_revolution">Neomuran revolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=28" title="Edit section: Neomuran revolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An alternative theory, proposed by <a href="/wiki/Thomas_Cavalier-Smith" title="Thomas Cavalier-Smith">Thomas Cavalier-Smith</a>, was labeled the <a href="/wiki/Neomura" title="Neomura">Neomuran revolution</a>. The designation "Neomuran revolution" refers to the appearances of the common ancestors of eukaryotes and archaea. Cavalier-Smith proposes that the first neomurans emerged 850 million years ago. Other molecular biologists assume that this group appeared much earlier, but Cavalier-Smith dismisses these claims because they are based on the "theoretically and empirically" unsound model of <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clocks</a>. Cavalier-Smith's theory of the Neomuran revolution has implications for the evolutionary history of the cellular machinery for recombination and sex. It suggests that this machinery evolved in two distinct bouts separated by a long period of stasis; first the appearance of recombination machinery in a bacterial ancestor which was maintained for 3 Gy(billion years), until the neomuran revolution when the mechanics were adapted to the presence of <a href="/wiki/Nucleosome" title="Nucleosome">nucleosomes</a>. The archaeal products of the revolution maintained recombination machinery that was essentially bacterial, whereas the eukaryotic products broke with this bacterial continuity. They introduced cell fusion and ploidy cycles into cell life histories. Cavalier-Smith argues that both bouts of mechanical evolution were motivated by similar selective forces: the need for accurate DNA replication without loss of viability.<sup id="cite_ref-TCSmith_84-0" class="reference"><a href="#cite_note-TCSmith-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> </p> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Questions">Questions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=29" title="Edit section: Questions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some questions biologists have attempted to answer include: </p> <ul><li>Why does sexual reproduction exist, if in many organisms it has a 50% cost (fitness disadvantage) in relation to asexual reproduction?<sup id="cite_ref-Ridley_M_2004_p._314_37-1" class="reference"><a href="#cite_note-Ridley_M_2004_p._314-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup></li> <li>Did <a href="/wiki/Mating_type" title="Mating type">mating types</a> (types of gametes, according to their compatibility) arise as a result of <a href="/wiki/Anisogamy" title="Anisogamy">anisogamy</a> (gamete dimorphism), or did mating types evolve before anisogamy?<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup></li> <li>Why do most sexual organisms use a binary <a href="/wiki/Mating_system" title="Mating system">mating system</a>? Grouping itself offers a survival advantage. A binary recognition based system is the most simple and effective method in maintaining species grouping.<sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup></li> <li>Why do some organisms have gamete dimorphism?</li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=30" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-Letunic_I_and_Bork_P-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Letunic_I_and_Bork_P_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Letunic_I_and_Bork_P_1-1"><sup><i><b>b</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="CITEREFLetunicBork2006" class="citation web cs1">Letunic, I; Bork, P (2006). <a rel="nofollow" class="external text" href="http://itol.embl.de/">"Interactive Tree of Life"</a><span class="reference-accessdate">. Retrieved <span class="nowrap">23 July</span> 2011</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Interactive+Tree+of+Life&rft.date=2006&rft.aulast=Letunic&rft.aufirst=I&rft.au=Bork%2C+P&rft_id=http%3A%2F%2Fitol.embl.de%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text">m<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLetunicBork2007" class="citation journal cs1">Letunic, I; Bork, P (2007). <a rel="nofollow" class="external text" href="http://itol.embl.de/help/17050570.pdf">"Interactive Tree of Life (iTOL): An online tool for phylogenetic tree display and annotation"</a> <span class="cs1-format">(PDF)</span>. <i><a href="/wiki/Bioinformatics_(journal)" title="Bioinformatics (journal)">Bioinformatics</a></i>. <b>23</b> (1): 127–8. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fbioinformatics%2Fbtl529">10.1093/bioinformatics/btl529</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/17050570">17050570</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Bioinformatics&rft.atitle=Interactive+Tree+of+Life+%28iTOL%29%3A+An+online+tool+for+phylogenetic+tree+display+and+annotation&rft.volume=23&rft.issue=1&rft.pages=127-8&rft.date=2007&rft_id=info%3Adoi%2F10.1093%2Fbioinformatics%2Fbtl529&rft_id=info%3Apmid%2F17050570&rft.aulast=Letunic&rft.aufirst=I&rft.au=Bork%2C+P&rft_id=http%3A%2F%2Fitol.embl.de%2Fhelp%2F17050570.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLetunicBork2011" class="citation journal cs1">Letunic, I; Bork, P (2011). <a rel="nofollow" class="external text" href="http://itol.embl.de/help/gkr201.pdf">"Interactive Tree of Life v2: Online annotation and display of phylogenetic trees made easy"</a> <span class="cs1-format">(PDF)</span>. <i><a href="/wiki/Nucleic_Acids_Research" title="Nucleic Acids Research">Nucleic Acids Research</a></i>. <b>39</b> (Web Server issue): W475–8. <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%2Fnar%2Fgkr201">10.1093/nar/gkr201</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/PMC3125724">3125724</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/21470960">21470960</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nucleic+Acids+Research&rft.atitle=Interactive+Tree+of+Life+v2%3A+Online+annotation+and+display+of+phylogenetic+trees+made+easy&rft.volume=39&rft.issue=Web+Server+issue&rft.pages=W475-8&rft.date=2011&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3125724%23id-name%3DPMC&rft_id=info%3Apmid%2F21470960&rft_id=info%3Adoi%2F10.1093%2Fnar%2Fgkr201&rft.aulast=Letunic&rft.aufirst=I&rft.au=Bork%2C+P&rft_id=http%3A%2F%2Fitol.embl.de%2Fhelp%2Fgkr201.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOtto2014" class="citation web cs1">Otto, Sarah (2014). <a rel="nofollow" class="external text" href="https://www.nature.com/scitable/topicpage/sexual-reproduction-and-the-evolution-of-sex-824">"Sexual Reproduction and the Evolution of Sex"</a>. <i>Scitable</i><span class="reference-accessdate">. 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(August 2001). <a rel="nofollow" class="external text" href="https://dx.doi.org/10.1038/35084593">"Do bacteria have sex?"</a>. <i>Nature Reviews Genetics</i>. <b>2</b> (8): 634–639. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F35084593">10.1038/35084593</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/11483988">11483988</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:5465846">5465846</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews+Genetics&rft.atitle=Do+bacteria+have+sex%3F&rft.volume=2&rft.issue=8&rft.pages=634-639&rft.date=2001-08&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A5465846%23id-name%3DS2CID&rft_id=info%3Apmid%2F11483988&rft_id=info%3Adoi%2F10.1038%2F35084593&rft.aulast=Redfield&rft.aufirst=Rosemary+J.&rft_id=http%3A%2F%2Fdx.doi.org%2F10.1038%2F35084593&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></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="CITEREFGoodenoughHeitman2014" class="citation journal cs1">Goodenough, U.; Heitman, J. (1 March 2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3949356">"Origins of Eukaryotic Sexual Reproduction"</a>. <i>Cold Spring Harbor Perspectives in Biology</i>. <b>6</b> (3): a016154. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1101%2Fcshperspect.a016154">10.1101/cshperspect.a016154</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1943-0264">1943-0264</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/PMC3949356">3949356</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/24591519">24591519</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cold+Spring+Harbor+Perspectives+in+Biology&rft.atitle=Origins+of+Eukaryotic+Sexual+Reproduction&rft.volume=6&rft.issue=3&rft.pages=a016154&rft.date=2014-03-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3949356%23id-name%3DPMC&rft.issn=1943-0264&rft_id=info%3Apmid%2F24591519&rft_id=info%3Adoi%2F10.1101%2Fcshperspect.a016154&rft.aulast=Goodenough&rft.aufirst=U.&rft.au=Heitman%2C+J.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3949356&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" 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="CITEREFDarwin1800" class="citation book cs1">Darwin, Erasmus (1800). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=YggpAAAAYAAJ&pg=PA104"><i>Phytologia …</i></a>. Dublin, Ireland: P. Byrne. p. 104.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Phytologia+%E2%80%A6&rft.place=Dublin%2C+Ireland&rft.pages=104&rft.pub=P.+Byrne&rft.date=1800&rft.aulast=Darwin&rft.aufirst=Erasmus&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DYggpAAAAYAAJ%26pg%3DPA104&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span> From p. 104: "As the progeny by lateral generation [i.e., vegetative (asexual) reproduction] so exactly resembles the parent stock, it follows, that though any new variety, or improvement, may be thus continued for a century or two, as in grafted fruit-trees, yet that no new variety or improvements can be obtained by this mode of generation; … " "But from the sexual, or amatorial, generation of plants new varieties, or improvements, are frequently obtained; as many of the young plants from seeds are dissimilar to the parent, and some of them supererior to the parent in the qualities we wish to possess; … " " … another advantage occurs from sexual generation, which is the production of new species of plants, or mules, … "</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">English translation: <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWeismann1889" class="citation book cs1">Weismann, August (1889). Poulton, Edward B.; Schönland, Selmar; Shipley, Arthur E. (eds.). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=iGysxQEACAAJ&pg=PA252"><i>Essays Upon Heredity and Kindred Biological Problems</i></a>. Oxford, England: Clarendon Press. pp. 252–332.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Essays+Upon+Heredity+and+Kindred+Biological+Problems&rft.place=Oxford%2C+England&rft.pages=252-332&rft.pub=Clarendon+Press&rft.date=1889&rft.aulast=Weismann&rft.aufirst=August&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DiGysxQEACAAJ%26pg%3DPA252&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span> Ch. 5: The Significance of Sexual Reproduction in the Theory of Natural Selection (1886) <ul><li>Weismann, Aug. (1885) <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/182986#page/42/mode/1up">"Die Bedeutung der sexuellen Fortpflanzung für die Selektions-Theorie"</a> [The significance of sexual reproduction in the theory of natural selection] Stilling, J. (ed.) <i>Tageblatt der 58. Versammlung Deutscher Naturforscher und Aerzte in Strassburg</i> [Daily News of the 58th Conference of German Natural Scientists and Physicians in Strassburg] (in German) Strassburg, Germany: G. Fischbach pp. 42-56.</li> <li>Revised and expanded in: <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWeismann1886" class="citation book cs1 cs1-prop-foreign-lang-source">Weismann, August (1886). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/18057#page/5/mode/1up"><i>Die Bedeutung der sexuellen Fortpflanzung für die Selektions-Theorie</i></a> (in German). Jena, Germany: Gustav Fischer.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Die+Bedeutung+der+sexuellen+Fortpflanzung+f%C3%BCr+die+Selektions-Theorie&rft.place=Jena%2C+Germany&rft.pub=Gustav+Fischer&rft.date=1886&rft.aulast=Weismann&rft.aufirst=August&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F18057%23page%2F5%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li></ul> </span></li> <li id="cite_note-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-9">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDarwin1876" class="citation book cs1">Darwin, Charles (1876). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/194109#page/478/mode/1up"><i>The Effects of Cross and Self Fertilisation in the Vegetable Kingdom</i></a>. London, England: John Murray. p. 462.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Effects+of+Cross+and+Self+Fertilisation+in+the+Vegetable+Kingdom&rft.place=London%2C+England&rft.pages=462&rft.pub=John+Murray&rft.date=1876&rft.aulast=Darwin&rft.aufirst=Charles&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F194109%23page%2F478%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></span> </li> <li id="cite_note-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-10">^</a></b></span> <span class="reference-text">Crow J.F. (1994). Advantages of Sexual Reproduction, Dev. 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Cambridge University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9780521293020" title="Special:BookSources/9780521293020"><bdi>9780521293020</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+Evolution+of+Sex&rft.pub=Cambridge+University+Press&rft.date=1978&rft.isbn=9780521293020&rft.aulast=Smith&rft.aufirst=J.+Maynard&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fevolutionofsex0000mayn&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></span> </li> <li id="cite_note-1946-_2005-35"><span class="mw-cite-backlink">^ <a href="#cite_ref-1946-_2005_35-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-1946-_2005_35-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-1946-_2005_35-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-1946-_2005_35-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-1946-_2005_35-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFStearns2005" class="citation book cs1">Stearns, S. 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Togashi, P. Cox (Eds.) <i>The Evolution of Anisogamy</i>. Cambridge University Press, Cambridge; 2011, p. 22-29.</span> </li> <li id="cite_note-86"><span class="mw-cite-backlink"><b><a href="#cite_ref-86">^</a></b></span> <span class="reference-text">Beukeboom, L. & Perrin, N. (2014). <i>The Evolution of Sex Determination</i>. Oxford University Press, p. 25 <a rel="nofollow" class="external autonumber" href="https://books.google.com/books?id=d4cLBAAAQBAJ&pg=PA25">[5]</a>. Online resources, <a rel="nofollow" class="external autonumber" href="http://www.oup.co.uk/companion/beukeboom">[6]</a>.</span> </li> <li id="cite_note-87"><span class="mw-cite-backlink"><b><a href="#cite_ref-87">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCzáránHoekstra2006" class="citation journal cs1">Czárán, T.L.; Hoekstra, R.F. (2006). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC524165">"Evolution of sexual asymmetry"</a>. <i>BMC Evolutionary Biology</i>. <b>4</b>: 34–46. <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/10.1186%2F1471-2148-4-34">10.1186/1471-2148-4-34</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC524165">524165</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/15383154">15383154</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BMC+Evolutionary+Biology&rft.atitle=Evolution+of+sexual+asymmetry&rft.volume=4&rft.pages=34-46&rft.date=2006&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC524165%23id-name%3DPMC&rft_id=info%3Apmid%2F15383154&rft_id=info%3Adoi%2F10.1186%2F1471-2148-4-34&rft.aulast=Cz%C3%A1r%C3%A1n&rft.aufirst=T.L.&rft.au=Hoekstra%2C+R.F.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC524165&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=31" 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 id="CITEREFBell1982" class="citation book cs1"><a href="/wiki/Graham_Bell_(biologist)" title="Graham Bell (biologist)">Bell, Graham</a> (1982). <i>The masterpiece of nature: the evolution and genetics of sexuality</i>. Berkeley: University of California Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-520-04583-5" title="Special:BookSources/978-0-520-04583-5"><bdi>978-0-520-04583-5</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+masterpiece+of+nature%3A+the+evolution+and+genetics+of+sexuality&rft.place=Berkeley&rft.pub=University+of+California+Press&rft.date=1982&rft.isbn=978-0-520-04583-5&rft.aulast=Bell&rft.aufirst=Graham&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBernsteinHarris_Bernstein1991" class="citation book cs1">Bernstein, Carol; Harris Bernstein (1991). <i>Aging, sex, and DNA repair</i>. Boston: Academic Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-092860-6" title="Special:BookSources/978-0-12-092860-6"><bdi>978-0-12-092860-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Aging%2C+sex%2C+and+DNA+repair&rft.place=Boston&rft.pub=Academic+Press&rft.date=1991&rft.isbn=978-0-12-092860-6&rft.aulast=Bernstein&rft.aufirst=Carol&rft.au=Harris+Bernstein&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHurstJ.R._Peck1996" class="citation journal cs1">Hurst, L.D.; J.R. Peck (1996). "Recent advances in the understanding of the evolution and maintenance of sex". <i><a href="/wiki/Trends_in_Ecology_and_Evolution" class="mw-redirect" title="Trends in Ecology and Evolution">Trends in Ecology and Evolution</a></i>. <b>11</b> (2): 46–52. <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/1996TEcoE..11...46H">1996TEcoE..11...46H</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2F0169-5347%2896%2981041-X">10.1016/0169-5347(96)81041-X</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/21237760">21237760</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+Ecology+and+Evolution&rft.atitle=Recent+advances+in+the+understanding+of+the+evolution+and+maintenance+of+sex&rft.volume=11&rft.issue=2&rft.pages=46-52&rft.date=1996&rft_id=info%3Apmid%2F21237760&rft_id=info%3Adoi%2F10.1016%2F0169-5347%2896%2981041-X&rft_id=info%3Abibcode%2F1996TEcoE..11...46H&rft.aulast=Hurst&rft.aufirst=L.D.&rft.au=J.R.+Peck&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLevinRichard_E._Michod1988" class="citation book cs1">Levin, Bruce R.; Richard E. Michod (1988). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/evolutionofsexex0000unse"><i>The Evolution of sex: an examination of current ideas</i></a></span>. Sunderland, Mass: Sinauer Associates. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87893-459-1" title="Special:BookSources/978-0-87893-459-1"><bdi>978-0-87893-459-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Evolution+of+sex%3A+an+examination+of+current+ideas&rft.place=Sunderland%2C+Mass&rft.pub=Sinauer+Associates&rft.date=1988&rft.isbn=978-0-87893-459-1&rft.aulast=Levin&rft.aufirst=Bruce+R.&rft.au=Richard+E.+Michod&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fevolutionofsexex0000unse&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMaynard_Smith1978" class="citation book cs1"><a href="/wiki/John_Maynard_Smith" title="John Maynard Smith">Maynard Smith, John</a> (1978). <a rel="nofollow" class="external text" href="https://archive.org/details/evolutionofsex0000mayn"><i>The evolution of sex</i></a>. Cambridge, UK: Cambridge University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-521-21887-0" title="Special:BookSources/978-0-521-21887-0"><bdi>978-0-521-21887-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+evolution+of+sex&rft.place=Cambridge%2C+UK&rft.pub=Cambridge+University+Press&rft.date=1978&rft.isbn=978-0-521-21887-0&rft.aulast=Maynard+Smith&rft.aufirst=John&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fevolutionofsex0000mayn&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMichod1995" class="citation book cs1">Michod, Richard E. (1995). <a rel="nofollow" class="external text" href="https://archive.org/details/erosevolutionnat0000mich"><i>Eros and evolution: a natural philosophy of sex</i></a>. Reading, Mass: Addison-Wesley Pub. Co. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-201-40754-9" title="Special:BookSources/978-0-201-40754-9"><bdi>978-0-201-40754-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Eros+and+evolution%3A+a+natural+philosophy+of+sex&rft.place=Reading%2C+Mass&rft.pub=Addison-Wesley+Pub.+Co&rft.date=1995&rft.isbn=978-0-201-40754-9&rft.aulast=Michod&rft.aufirst=Richard+E.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Ferosevolutionnat0000mich&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation news cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20151219170335/http://www.nbcnews.com/id/27927661/">"Scientists put sex origin mystery to bed, Wild strawberry research provides evidence on when gender emerges"</a>. <i><a href="/wiki/NBC_News" title="NBC News">NBC News</a></i>. Archived from <a rel="nofollow" class="external text" href="http://www.nbcnews.com/id/27927661">the original</a> on 19 December 2015<span class="reference-accessdate">. Retrieved <span class="nowrap">25 November</span> 2008</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=NBC+News&rft.atitle=Scientists+put+sex+origin+mystery+to+bed%2C+Wild+strawberry+research+provides+evidence+on+when+gender+emerges&rft_id=http%3A%2F%2Fwww.nbcnews.com%2Fid%2F27927661&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRidley1993" class="citation book cs1"><a href="/wiki/Mark_Ridley_(zoologist)" title="Mark Ridley (zoologist)">Ridley, Mark</a> (1993). <i>Evolution</i>. Oxford: Blackwell Scientific. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-632-03481-9" title="Special:BookSources/978-0-632-03481-9"><bdi>978-0-632-03481-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution&rft.place=Oxford&rft.pub=Blackwell+Scientific&rft.date=1993&rft.isbn=978-0-632-03481-9&rft.aulast=Ridley&rft.aufirst=Mark&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRidley2000" class="citation book cs1"><a href="/wiki/Mark_Ridley_(zoologist)" title="Mark Ridley (zoologist)">Ridley, Mark</a> (2000). <i>Mendel's demon: gene justice and the complexity of life</i>. London: Weidenfeld & Nicolson. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-297-64634-1" title="Special:BookSources/978-0-297-64634-1"><bdi>978-0-297-64634-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Mendel%27s+demon%3A+gene+justice+and+the+complexity+of+life&rft.place=London&rft.pub=Weidenfeld+%26+Nicolson&rft.date=2000&rft.isbn=978-0-297-64634-1&rft.aulast=Ridley&rft.aufirst=Mark&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRidley1995" class="citation book cs1"><a href="/wiki/Matt_Ridley" title="Matt Ridley">Ridley, Matt</a> (1995). <i>The Red Queen: sex and the evolution of human nature</i>. New York: Penguin Books. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-14-024548-6" title="Special:BookSources/978-0-14-024548-6"><bdi>978-0-14-024548-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Red+Queen%3A+sex+and+the+evolution+of+human+nature&rft.place=New+York&rft.pub=Penguin+Books&rft.date=1995&rft.isbn=978-0-14-024548-6&rft.aulast=Ridley&rft.aufirst=Matt&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSzathmáryJohn_Maynard_Smith1995" class="citation book cs1"><a href="/wiki/E%C3%B6rs_Szathm%C3%A1ry" title="Eörs Szathmáry">Szathmáry, Eörs</a>; <a href="/wiki/John_Maynard_Smith" title="John Maynard Smith">John Maynard Smith</a> (1995). <a href="/wiki/The_Major_Transitions_in_Evolution" title="The Major Transitions in Evolution"><i>The Major Transitions in Evolution</i></a>. Oxford: W.H. Freeman Spektrum. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7167-4525-9" title="Special:BookSources/978-0-7167-4525-9"><bdi>978-0-7167-4525-9</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+Major+Transitions+in+Evolution&rft.place=Oxford&rft.pub=W.H.+Freeman+Spektrum&rft.date=1995&rft.isbn=978-0-7167-4525-9&rft.aulast=Szathm%C3%A1ry&rft.aufirst=E%C3%B6rs&rft.au=John+Maynard+Smith&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTaylor1996" class="citation book cs1">Taylor, Timothy (1996). <a rel="nofollow" class="external text" href="https://archive.org/details/prehistoryofsexf00tay_zn9"><i>The prehistory of sex: four million years of human sexual culture</i></a>. New York: Bantam Books. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-553-09694-1" title="Special:BookSources/978-0-553-09694-1"><bdi>978-0-553-09694-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+prehistory+of+sex%3A+four+million+years+of+human+sexual+culture&rft.place=New+York&rft.pub=Bantam+Books&rft.date=1996&rft.isbn=978-0-553-09694-1&rft.aulast=Taylor&rft.aufirst=Timothy&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fprehistoryofsexf00tay_zn9&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWilliams1975" class="citation book cs1"><a href="/wiki/George_C._Williams_(biologist)" class="mw-redirect" title="George C. Williams (biologist)">Williams, George</a> (1975). <i>Sex and evolution</i>. Princeton, N.J: Princeton University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-691-08147-2" title="Special:BookSources/978-0-691-08147-2"><bdi>978-0-691-08147-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Sex+and+evolution&rft.place=Princeton%2C+N.J&rft.pub=Princeton+University+Press&rft.date=1975&rft.isbn=978-0-691-08147-2&rft.aulast=Williams&rft.aufirst=George&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+sexual+reproduction" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_sexual_reproduction&action=edit&section=32" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://www.livescience.com/health/050330_sex_good.html">Why Sex is Good (2005)</a></li> <li><a rel="nofollow" class="external text" href="http://www.philippwesche.org/old1/es.html">The Evolution of Sexs (2003)</a></li> <li><a rel="nofollow" class="external text" href="http://www.evolocus.com/Textbooks/Geodakian2012.pdf">The Evolutionary Theory of Sex (2012)</a></li> <li><a rel="nofollow" class="external text" href="https://www.nature.com/scitable/topicpage/sexual-reproduction-and-the-evolution-of-sex-824/">Sexual Reproduction and the Evolution of Sex</a> (<a rel="nofollow" class="external text" href="https://archive.today/20231008141323/https://www.nature.com/scitable/topicpage/sexual-reproduction-and-the-evolution-of-sex-824/">Archived (2023)</a>) − <a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature journal</a> (2008)</li></ul> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1236075235">.mw-parser-output .navbox{box-sizing:border-box;border:1px solid #a2a9b1;width:100%;clear:both;font-size:88%;text-align:center;padding:1px;margin:1em auto 0}.mw-parser-output .navbox .navbox{margin-top:0}.mw-parser-output .navbox+.navbox,.mw-parser-output .navbox+.navbox-styles+.navbox{margin-top:-1px}.mw-parser-output .navbox-inner,.mw-parser-output .navbox-subgroup{width:100%}.mw-parser-output .navbox-group,.mw-parser-output .navbox-title,.mw-parser-output .navbox-abovebelow{padding:0.25em 1em;line-height:1.5em;text-align:center}.mw-parser-output .navbox-group{white-space:nowrap;text-align:right}.mw-parser-output .navbox,.mw-parser-output .navbox-subgroup{background-color:#fdfdfd}.mw-parser-output .navbox-list{line-height:1.5em;border-color:#fdfdfd}.mw-parser-output .navbox-list-with-group{text-align:left;border-left-width:2px;border-left-style:solid}.mw-parser-output tr+tr>.navbox-abovebelow,.mw-parser-output tr+tr>.navbox-group,.mw-parser-output tr+tr>.navbox-image,.mw-parser-output tr+tr>.navbox-list{border-top:2px solid #fdfdfd}.mw-parser-output .navbox-title{background-color:#ccf}.mw-parser-output .navbox-abovebelow,.mw-parser-output .navbox-group,.mw-parser-output .navbox-subgroup .navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title 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template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Sex_(biology)" title="Special:EditPage/Template:Sex (biology)"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Sex" style="font-size:114%;margin:0 4em"><a href="/wiki/Sex" title="Sex">Sex</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Biological<br />terms</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/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a> <ul><li><a href="/wiki/Male" title="Male">Male</a></li> <li><a href="/wiki/Female" title="Female">Female</a></li></ul></li> <li><a href="/wiki/Sexual_differentiation" title="Sexual differentiation">Sexual differentiation</a> <ul><li><a href="/wiki/Feminization_(biology)" title="Feminization (biology)">Feminization</a></li> <li><a href="/wiki/Virilization" title="Virilization">Virilization</a></li></ul></li> <li><a href="/wiki/Sex-determination_system" title="Sex-determination system">Sex-determination system</a> <ul><li><a href="/wiki/XY_sex-determination_system" title="XY sex-determination system">XY</a></li> <li><a href="/wiki/ZW_sex-determination_system" title="ZW sex-determination system">ZW</a></li> <li><a href="/wiki/XO_sex-determination_system" title="XO sex-determination system">XO</a></li> <li><a href="/wiki/ZO_sex-determination_system" title="ZO sex-determination system">ZO</a></li> <li><a href="/wiki/Temperature-dependent_sex_determination" title="Temperature-dependent sex determination">Temperature-dependent</a></li> <li><a href="/wiki/Haplodiploidy" title="Haplodiploidy">Haplodiploidy</a></li></ul></li> <li><a href="/wiki/Sex_chromosome" title="Sex chromosome">Sex chromosome</a></li> <li><a href="/wiki/Testis-determining_factor" class="mw-redirect" title="Testis-determining factor">Testis-determining factor</a></li> <li><a href="/wiki/Hermaphrodite" title="Hermaphrodite">Hermaphrodite</a> <ul><li><a href="/wiki/Sequential_hermaphroditism" title="Sequential hermaphroditism">Sequential hermaphroditism</a></li> <li><a href="/wiki/Simultaneous_hermaphroditism" title="Simultaneous hermaphroditism">Simultaneous hermaphroditism</a></li></ul></li> <li><a href="/wiki/Intersex" title="Intersex">Intersex</a></li> <li><a href="/wiki/Parasexual_cycle" title="Parasexual cycle">parasexuality</a></li> <li><a href="/wiki/Sex_as_a_biological_variable" title="Sex as a biological variable">Sex as a biological variable</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sexual_reproduction" title="Sexual reproduction">Sexual<br />reproduction</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 class="mw-selflink selflink">Evolution of sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy" title="Anisogamy">Anisogamy</a></li> <li><a href="/wiki/Isogamy" title="Isogamy">Isogamy</a></li></ul></li> <li><a href="/wiki/Germ_cell" title="Germ cell">Germ cell</a></li> <li><a href="/wiki/Reproductive_system" title="Reproductive system">Reproductive system</a></li> <li><a href="/wiki/Sex_organ" title="Sex organ">Sex organ</a></li> <li><a href="/wiki/Mating" title="Mating">Mating</a></li> <li><a href="/wiki/Meiosis" title="Meiosis">Meiosis</a></li> <li><a href="/wiki/Gametogenesis" title="Gametogenesis">Gametogenesis</a> <ul><li><a href="/wiki/Spermatogenesis" title="Spermatogenesis">Spermatogenesis</a></li> <li><a href="/wiki/Oogenesis" title="Oogenesis">Oogenesis</a></li></ul></li> <li><a href="/wiki/Gamete" title="Gamete">Gamete</a> <ul><li><a href="/wiki/Spermatozoon" title="Spermatozoon">spermatozoon</a></li> <li><a href="/wiki/Egg_cell" title="Egg cell">ovum</a></li></ul></li> <li><a href="/wiki/Fertilisation" title="Fertilisation">Fertilization</a> <ul><li><a href="/wiki/External_fertilization" title="External fertilization">External</a></li> <li><a href="/wiki/Internal_fertilization" title="Internal fertilization">Internal</a></li></ul></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a></li> <li><a href="/wiki/Plant_reproduction" title="Plant reproduction">Plant reproduction</a></li> <li><a href="/wiki/Mating_in_fungi" title="Mating in fungi">Fungal reproduction</a></li> <li><a href="/wiki/Sexual_reproduction_in_animals" class="mw-redirect" title="Sexual reproduction in animals">Sexual reproduction in animals</a> <ul><li><a href="/wiki/Sexual_intercourse" title="Sexual intercourse">Sexual intercourse</a> <ul><li><a href="/wiki/Penile-vaginal_intercourse" title="Penile-vaginal intercourse">Penile-vaginal intercourse</a></li></ul></li> <li><a href="/wiki/Copulation_(zoology)" title="Copulation (zoology)">Copulation</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/Human_reproduction" title="Human reproduction">Human reproduction</a></li> <li><a href="/wiki/Lordosis_behavior" title="Lordosis behavior">Lordosis behavior</a></li> <li><a href="/wiki/Pelvic_thrust" title="Pelvic thrust">Pelvic thrust</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Human_sexuality" title="Human sexuality">Sexuality</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/Plant_reproductive_morphology" title="Plant reproductive morphology">Plant sexuality</a></li> <li><a href="/wiki/Animal_sexual_behaviour" title="Animal sexual behaviour">Animal sexuality</a> <ul><li><a href="/wiki/Human_sexuality" title="Human sexuality">Human sexuality</a> <ul><li><a href="/wiki/Mechanics_of_human_sexuality" title="Mechanics of human sexuality">Mechanics</a></li> <li><a href="/wiki/Sexual_differentiation_in_humans" title="Sexual differentiation in humans">Differentiation</a></li> <li><a href="/wiki/Human_sexual_activity" title="Human sexual activity">Activity</a></li></ul></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Sex" title="Category:Sex">Category</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_biology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a 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 class="mw-selflink selflink">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="The_development_of_phenotype" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><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:Genarch" title="Template:Genarch"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genarch" title="Template talk:Genarch"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genarch" title="Special:EditPage/Template:Genarch"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="The_development_of_phenotype" style="font-size:114%;margin:0 4em">The <a href="/wiki/Morphogenesis" title="Morphogenesis">development</a> of <a href="/wiki/Phenotype" title="Phenotype">phenotype</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key concepts</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Reaction_norm" title="Reaction norm">Reaction norm</a></li> <li><a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">Gene–environment interaction</a></li> <li><a href="/wiki/Gene%E2%80%93environment_correlation" title="Gene–environment correlation">Gene–environment correlation</a></li> <li><a href="/wiki/Operon" title="Operon">Operon</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Heterochrony" title="Heterochrony">Heterochrony</a> <ul><li><a href="/wiki/Neoteny" title="Neoteny">Neoteny</a></li></ul></li> <li><a href="/wiki/Heterotopy" title="Heterotopy">Heterotopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_architecture" title="Genetic architecture">Genetic architecture</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a></li> <li><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a>/<a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">evolutionary landscape</a></li> <li><a href="/wiki/Pleiotropy" title="Pleiotropy">Pleiotropy</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Plasticity</a></li> <li><a href="/wiki/Polygenic_inheritance" class="mw-redirect" title="Polygenic inheritance">Polygenic inheritance</a></li> <li><a href="/wiki/Transgressive_segregation" title="Transgressive segregation">Transgressive segregation</a></li> <li><a href="/wiki/Sequence_space_(evolution)" title="Sequence space (evolution)">Sequence space</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-genetic influences</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Maternal_effect" title="Maternal effect">Maternal effect</a></li> <li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">Genomic imprinting</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Polyphenism" title="Polyphenism">Polyphenism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Developmental architecture</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Morphogenesis" title="Morphogenesis">Morphogenesis</a> <ul><li><a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">Eyespot</a></li> <li><a href="/wiki/Pattern_formation" title="Pattern formation">Pattern formation</a></li></ul></li> <li><a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">Segmentation</a></li> <li><a href="/wiki/Metamerism_(biology)" title="Metamerism (biology)">Metamerism</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution of genetic systems</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolvability" title="Evolvability">Evolvability</a></li> <li><a href="/wiki/Robustness_(evolution)" title="Robustness (evolution)">Robustness</a></li> <li><a href="/wiki/Neutral_network_(evolution)" title="Neutral network (evolution)">Neutral networks</a></li> <li><a class="mw-selflink selflink">Evolution of sexual reproduction</a></li> <li><a href="/wiki/Evolutionary_tinkering" title="Evolutionary tinkering">Tinkering</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Control of development</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Systems</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation of gene expression</a></li> <li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Evo-devo_gene_toolkit" title="Evo-devo gene toolkit">Evo-devo gene toolkit</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Homeobox" title="Homeobox">Homeobox</a></li> <li><a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Elements</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Homeotic_gene" title="Homeotic gene">Homeotic gene</a></li> <li><a href="/wiki/Hox_gene" title="Hox gene">Hox gene</a></li> <li><a href="/wiki/Pax_genes" title="Pax genes">Pax genes</a> <ul><li><a href="/wiki/PAX6" title="PAX6">eyeless gene</a></li></ul></li> <li><a href="/wiki/DLX_gene_family" title="DLX gene family">Distal-less</a></li> <li><a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li> <li><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</a></li> <li><a href="/wiki/Morphogen" title="Morphogen">Morphogen</a></li> <li><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Cell surface receptor</a></li> <li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential figures</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a></li> <li><a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a></li> <li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> + <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> <ul><li><a href="/wiki/Lac_operon" title="Lac operon">Lac operon</a></li></ul></li> <li><a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a></li> <li><a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a></li> <li><a href="/wiki/William_McGinnis" title="William McGinnis">William McGinnis</a></li> <li><a href="/wiki/Michael_Levine_(biologist)" title="Michael Levine (biologist)">Mike Levine</a></li> <li><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Sean B. Carroll</a> <ul><li><i><a href="/wiki/Endless_Forms_Most_Beautiful_(book)" title="Endless Forms Most Beautiful (book)">Endless Forms Most Beautiful</a></i></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Debates</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Morphogenetic_field" title="Morphogenetic field">Morphogenetic field</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</a></div></td></tr></tbody></table></div> <style data-mw-deduplicate="TemplateStyles:r1130092004">.mw-parser-output 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src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/21px-Tree_of_life.svg.png" decoding="async" width="21" height="17" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/32px-Tree_of_life.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/42px-Tree_of_life.svg.png 2x" data-file-width="915" data-file-height="760" /></a></span> </span><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology</a></li><li class="portal-bar-item"><span class="nowrap"><span typeof="mw:File"><a href="/wiki/File:Nuvola_apps_kalzium.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Nuvola_apps_kalzium.svg/19px-Nuvola_apps_kalzium.svg.png" decoding="async" width="19" height="19" class="mw-file-element" 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