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id="toc-Evolution_of_life_cycles" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution_of_life_cycles"> <div class="vector-toc-text"> 2 Evolution of life cycles </div> </a> <ul id="toc-Evolution_of_life_cycles-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution_of_plant_anatomy" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution_of_plant_anatomy"> <div class="vector-toc-text"> 3 Evolution of plant anatomy </div> </a> <button aria-controls="toc-Evolution_of_plant_anatomy-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Evolution of plant anatomy subsection </button> <ul id="toc-Evolution_of_plant_anatomy-sublist" class="vector-toc-list"> <li id="toc-Arbuscular_mycorrhizal_symbiosis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Arbuscular_mycorrhizal_symbiosis"> <div class="vector-toc-text"> 3.1 Arbuscular mycorrhizal symbiosis </div> </a> <ul id="toc-Arbuscular_mycorrhizal_symbiosis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cuticle,_stomata_and_intercellular_spaces" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cuticle,_stomata_and_intercellular_spaces"> <div class="vector-toc-text"> 3.2 Cuticle, stomata and intercellular spaces </div> </a> <ul id="toc-Cuticle,_stomata_and_intercellular_spaces-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Xylem" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Xylem"> <div class="vector-toc-text"> 3.3 Xylem </div> </a> <ul id="toc-Xylem-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Endodermis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Endodermis"> <div class="vector-toc-text"> 3.4 Endodermis </div> </a> <ul id="toc-Endodermis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution_of_plant_morphology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution_of_plant_morphology"> <div class="vector-toc-text"> 4 Evolution of plant morphology </div> </a> <button aria-controls="toc-Evolution_of_plant_morphology-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Evolution of plant morphology subsection </button> <ul id="toc-Evolution_of_plant_morphology-sublist" class="vector-toc-list"> <li id="toc-Leaves" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Leaves"> <div class="vector-toc-text"> 4.1 Leaves </div> </a> <ul id="toc-Leaves-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Roots" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Roots"> <div class="vector-toc-text"> 4.2 Roots </div> </a> <ul id="toc-Roots-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tree_form" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Tree_form"> <div class="vector-toc-text"> 4.3 Tree form </div> </a> <ul id="toc-Tree_form-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Seeds" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Seeds"> <div class="vector-toc-text"> 4.4 Seeds </div> </a> <ul id="toc-Seeds-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Flowers" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Flowers"> <div class="vector-toc-text"> 4.5 Flowers </div> </a> <ul id="toc-Flowers-sublist" class="vector-toc-list"> <li id="toc-Origins_of_the_flower" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Origins_of_the_flower"> <div class="vector-toc-text"> 4.5.1 Origins of the flower </div> </a> <ul id="toc-Origins_of_the_flower-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Adaptive_function_of_flowers" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Adaptive_function_of_flowers"> <div class="vector-toc-text"> 4.6 Adaptive function of flowers </div> </a> <ul id="toc-Adaptive_function_of_flowers-sublist" class="vector-toc-list"> <li id="toc-Evolution_of_the_MADS-box_family" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Evolution_of_the_MADS-box_family"> <div class="vector-toc-text"> 4.6.1 Evolution of the MADS-box family </div> </a> <ul id="toc-Evolution_of_the_MADS-box_family-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Factors_influencing_floral_diversity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Factors_influencing_floral_diversity"> <div class="vector-toc-text"> 4.7 Factors influencing floral diversity </div> </a> <ul id="toc-Factors_influencing_floral_diversity-sublist" class="vector-toc-list"> <li id="toc-Flowering_time" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Flowering_time"> <div class="vector-toc-text"> 4.7.1 Flowering time </div> </a> <ul id="toc-Flowering_time-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Theories_of_flower_evolution" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Theories_of_flower_evolution"> <div class="vector-toc-text"> 4.7.2 Theories of flower evolution </div> </a> <ul id="toc-Theories_of_flower_evolution-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mechanisms_and_players_in_evolution_of_plant_morphology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mechanisms_and_players_in_evolution_of_plant_morphology"> <div class="vector-toc-text"> 4.8 Mechanisms and players in evolution of plant morphology </div> </a> <ul id="toc-Mechanisms_and_players_in_evolution_of_plant_morphology-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution_of_photosynthetic_pathways" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution_of_photosynthetic_pathways"> <div class="vector-toc-text"> 5 Evolution of photosynthetic pathways </div> </a> <button aria-controls="toc-Evolution_of_photosynthetic_pathways-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Evolution of photosynthetic pathways subsection </button> <ul id="toc-Evolution_of_photosynthetic_pathways-sublist" class="vector-toc-list"> <li id="toc-Concentrating_carbon" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Concentrating_carbon"> <div class="vector-toc-text"> 5.1 Concentrating carbon </div> </a> <ul id="toc-Concentrating_carbon-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolutionary_record" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolutionary_record"> <div class="vector-toc-text"> 5.2 Evolutionary record </div> </a> <ul id="toc-Evolutionary_record-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Advantage_of_C4" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Advantage_of_C4"> <div class="vector-toc-text"> 5.3 Advantage of C4 </div> </a> <ul id="toc-Advantage_of_C4-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution_of_transcriptional_regulation" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution_of_transcriptional_regulation"> <div class="vector-toc-text"> 6 Evolution of transcriptional regulation </div> </a> <ul id="toc-Evolution_of_transcriptional_regulation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution_of_secondary_metabolism" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution_of_secondary_metabolism"> <div class="vector-toc-text"> 7 Evolution of secondary metabolism </div> </a> <ul id="toc-Evolution_of_secondary_metabolism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution_of_plant-microbe_interactions" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution_of_plant-microbe_interactions"> <div class="vector-toc-text"> 8 Evolution of plant-microbe interactions </div> </a> <ul id="toc-Evolution_of_plant-microbe_interactions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> 9 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 10 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 11 External links </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" > Toggle the table of contents </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading">Evolutionary history of plants</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 23 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-23" aria-hidden="true" > 23 languages </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A7%D9%84%D8%AA%D8%A7%D8%B1%D9%8A%D8%AE_%D8%A7%D9%84%D8%AA%D8%B7%D9%88%D8%B1%D9%8A_%D9%84%D9%84%D9%86%D8%A8%D8%A7%D8%AA%D8%A7%D8%AA" title="التاريخ التطوري للنباتات – Arabic" lang="ar" hreflang="ar" data-title="التاريخ التطوري للنباتات" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Hist%C3%B2ria_evolutiva_de_les_plantes" title="Història evolutiva de les plantes – Catalan" lang="ca" hreflang="ca" data-title="Història evolutiva de les plantes" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Planternes_evolution" title="Planternes evolution – Danish" lang="da" hreflang="da" data-title="Planternes evolution" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Historia_evolutiva_de_las_plantas" title="Historia evolutiva de las plantas – Spanish" lang="es" hreflang="es" data-title="Historia evolutiva de las plantas" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AA%D8%A7%D8%B1%DB%8C%D8%AE_%D9%81%D8%B1%DA%AF%D8%B4%D8%AA%DB%8C_%DA%AF%DB%8C%D8%A7%D9%87%D8%A7%D9%86" title="تاریخ فرگشتی گیاهان – Persian" lang="fa" hreflang="fa" data-title="تاریخ فرگشتی گیاهان" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_v%C3%A9g%C3%A9taux" title="Histoire évolutive des végétaux – French" lang="fr" hreflang="fr" data-title="Histoire évolutive des végétaux" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Historia_evolutiva_das_plantas" title="Historia evolutiva das plantas – Galician" lang="gl" hreflang="gl" data-title="Historia evolutiva das plantas" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%8B%9D%EB%AC%BC%EC%9D%98_%EC%A7%84%ED%99%94" title="식물의 진화 – Korean" lang="ko" hreflang="ko" data-title="식물의 진화" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%AA%E0%A4%BE%E0%A4%A6%E0%A4%AA%E0%A5%8B%E0%A4%82_%E0%A4%95%E0%A4%BE_%E0%A4%B5%E0%A4%BF%E0%A4%95%E0%A4%BE%E0%A4%B8%E0%A4%BE%E0%A4%A4%E0%A5%8D%E0%A4%AE%E0%A4%95_%E0%A4%87%E0%A4%A4%E0%A4%BF%E0%A4%B9%E0%A4%BE%E0%A4%B8" title="पादपों का विकासात्मक इतिहास – Hindi" lang="hi" hreflang="hi" data-title="पादपों का विकासात्मक इतिहास" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%99%D7%A1%D7%98%D7%95%D7%A8%D7%99%D7%94_%D7%90%D7%91%D7%95%D7%9C%D7%95%D7%A6%D7%99%D7%95%D7%A0%D7%99%D7%AA_%D7%A9%D7%9C_%D7%A6%D7%9E%D7%97%D7%99%D7%9D" title="היסטוריה אבולוציונית של צמחים – Hebrew" lang="he" hreflang="he" data-title="היסטוריה אבולוציונית של צמחים" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D3%A8%D1%81%D1%96%D0%BC%D0%B4%D1%96%D0%BA%D1%82%D0%B5%D1%80_%D1%8D%D0%B2%D0%BE%D0%BB%D1%8E%D1%86%D0%B8%D1%8F%D1%81%D1%8B" title="Өсімдіктер эволюциясы – Kazakh" lang="kk" hreflang="kk" data-title="Өсімдіктер эволюциясы" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E6%A4%8D%E7%89%A9%E3%81%AE%E9%80%B2%E5%8C%96" title="植物の進化 – Japanese" lang="ja" hreflang="ja" data-title="植物の進化" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Hist%C3%B3ria_evolutiva_das_plantas" title="História evolutiva das plantas – Portuguese" lang="pt" hreflang="pt" data-title="História evolutiva das plantas" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%AD%D0%B2%D0%BE%D0%BB%D1%8E%D1%86%D0%B8%D1%8F_%D1%80%D0%B0%D1%81%D1%82%D0%B5%D0%BD%D0%B8%D0%B9" title="Эволюция растений – Russian" lang="ru" hreflang="ru" data-title="Эволюция растений" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B7%81%E0%B7%8F%E0%B6%9A_%E0%B7%80%E0%B6%BD_%E0%B6%B4%E0%B6%BB%E0%B7%92%E0%B6%AB%E0%B7%8F%E0%B6%B8%E0%B6%BA%E0%B7%9A_%E0%B6%89%E0%B6%AD%E0%B7%92%E0%B7%84%E0%B7%8F%E0%B7%83%E0%B6%BA" title="ශාක වල පරිණාමයේ ඉතිහාසය – Sinhala" lang="si" hreflang="si" data-title="ශාක වල පරිණාමයේ ඉතිහාසය" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target">සිංහල</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Evolution_of_plants" title="Evolution of plants – Simple English" lang="en-simple" hreflang="en-simple" data-title="Evolution of plants" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D9%85%DB%8E%DA%98%D9%88%D9%88%DB%8C_%D9%BE%DB%95%D8%B1%DB%95%D8%B3%DB%95%D9%86%D8%AF%D9%86%DB%8C_%DA%95%D9%88%DB%95%DA%A9%DB%95%DA%A9%D8%A7%D9%86" title="مێژووی پەرەسەندنی ڕوەکەکان – Central Kurdish" lang="ckb" hreflang="ckb" data-title="مێژووی پەرەسەندنی ڕوەکەکان" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Kasvien_evoluution_aikajana" title="Kasvien evoluution aikajana – Finnish" lang="fi" hreflang="fi" data-title="Kasvien evoluution aikajana" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/V%C3%A4xternas_evolution" title="Växternas evolution – Swedish" lang="sv" hreflang="sv" data-title="Växternas evolution" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Bitkilerin_evrimi" title="Bitkilerin evrimi – Turkish" lang="tr" hreflang="tr" data-title="Bitkilerin evrimi" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%95%D0%B2%D0%BE%D0%BB%D1%8E%D1%86%D1%96%D1%8F_%D1%80%D0%BE%D1%81%D0%BB%D0%B8%D0%BD" title="Еволюція рослин – Ukrainian" lang="uk" hreflang="uk" data-title="Еволюція рослин" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li 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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">Further information: <a href="/wiki/Evolutionary_history_of_life" class="mw-redirect" title="Evolutionary history of life">Evolutionary history of life</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Trilete_spores.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/Trilete_spores.png/220px-Trilete_spores.png" decoding="async" width="220" height="221" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/Trilete_spores.png/330px-Trilete_spores.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c0/Trilete_spores.png/440px-Trilete_spores.png 2x" data-file-width="661" data-file-height="665" /></a><figcaption>A <a href="/wiki/Late_Silurian" class="mw-redirect" title="Late Silurian">late Silurian</a> <a href="/wiki/Sporangium" title="Sporangium">sporangium</a>, artificially colored. Green: A spore tetrad. Blue: A spore bearing a trilete mark – the Y-shaped scar. The spores are about 30–35 μm across.</figcaption></figure> The <a href="/wiki/Evolution" title="Evolution">evolution</a> of <a href="/wiki/Plant" title="Plant">plants</a> has resulted in a wide range of complexity, from the earliest <a href="/wiki/Algal_mat" title="Algal mat">algal mats</a> of unicellular <a href="/wiki/Archaeplastid" class="mw-redirect" title="Archaeplastid">archaeplastids</a> evolved through <a href="/wiki/Endosymbiosis" class="mw-redirect" title="Endosymbiosis">endosymbiosis</a>, through multicellular <a href="/wiki/Marine_habitat" title="Marine habitat">marine</a> and <a href="/wiki/Freshwater" class="mw-redirect" title="Freshwater">freshwater</a> <a href="/wiki/Green_algae" title="Green algae">green algae</a>, to <a href="/wiki/Spore" title="Spore">spore</a>-bearing terrestrial <a href="/wiki/Bryophyte" title="Bryophyte">bryophytes</a>, <a href="/wiki/Lycopod" class="mw-redirect" title="Lycopod">lycopods</a> and <a href="/wiki/Fern" title="Fern">ferns</a>, and eventually to the complex <a href="/wiki/Seed" title="Seed">seed</a>-bearing <a href="/wiki/Gymnosperm" title="Gymnosperm">gymnosperms</a> and <a href="/wiki/Angiosperm" class="mw-redirect" title="Angiosperm">angiosperms</a> (<a href="/wiki/Flowering" class="mw-redirect" title="Flowering">flowering</a> plants) of today. While many of the earliest groups continue to thrive, as exemplified by <a href="/wiki/Red_algae" title="Red algae">red</a> and green algae in marine environments, more recently derived groups have displaced previously ecologically dominant ones; for example, the ascendance of flowering plants over gymnosperms in terrestrial environments.<a href="#cite_note-Stewart-1993-1">[1]</a>: 498  There is evidence that <a href="/wiki/Cyanobacteria" title="Cyanobacteria">cyanobacteria</a> and multicellular <a href="/wiki/Thallus" title="Thallus">thalloid</a> eukaryotes lived in freshwater communities on land as early as 1 billion years ago,<a href="#cite_note-Strother-2011-2">[2]</a> and that communities of complex, multicellular photosynthesizing organisms existed on land in the late <a href="/wiki/Precambrian" title="Precambrian">Precambrian</a>, around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=850">850</a> million years ago.<a href="#cite_note-Knauth-2009-3">[3]</a> Evidence of the emergence of <a href="/wiki/Embryophyte" title="Embryophyte">embryophyte</a> land plants first occurs in the middle <a href="/wiki/Ordovician" title="Ordovician">Ordovician</a> (~<a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=470">470</a> million years ago), and by the middle of the <a href="/wiki/Devonian" title="Devonian">Devonian</a> (~<a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=390">390</a> million years ago), many of the features recognised in land plants today were present, including roots and leaves. By the late Devonian (~<a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=370">370</a> million years ago) some free-sporing plants such as <a href="/wiki/Archaeopteris" title="Archaeopteris">Archaeopteris</a> had <a href="/wiki/Vascular_cambium" title="Vascular cambium">secondary vascular tissue</a> that produced <a href="/wiki/Wood" title="Wood">wood</a> and had formed forests of tall trees. Also by the late Devonian, Elkinsia, an early <a href="/wiki/Pteridospermatophyta" title="Pteridospermatophyta">seed fern</a>, had evolved seeds.<a href="#cite_note-Rothwell-1989-4">[4]</a> Evolutionary innovation continued throughout the rest of the <a href="/wiki/Phanerozoic" title="Phanerozoic">Phanerozoic</a> eon and still continues today. Most plant groups were relatively unscathed by the <a href="/wiki/Permo-Triassic_extinction_event" class="mw-redirect" title="Permo-Triassic extinction event">Permo-Triassic extinction event</a>, although the structures of communities changed. This may have set the scene for the appearance of the flowering plants in the <a href="/wiki/Triassic" title="Triassic">Triassic</a> (~<a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=200">200</a> million years ago), and their later diversification in the Cretaceous and <a href="/wiki/Paleogene" title="Paleogene">Paleogene</a>. The latest major group of plants to evolve were the <a href="/wiki/Poaceae" title="Poaceae">grasses</a>, which became important in the mid-Paleogene, from around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=40">40</a> million years ago. The grasses, as well as many other groups, evolved new mechanisms of metabolism to survive the low CO2 and warm, dry conditions of the <a href="/wiki/Tropics" title="Tropics">tropics</a> over the last <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=10">10</a> million years. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Colonization_of_land">Colonization of land</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=1" title="Edit section: Colonization of land">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Divergence">Divergence</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=2" title="Edit section: Divergence">edit</a>]</div> <a href="/wiki/Land_plant" class="mw-redirect" title="Land plant">Land plants</a> evolved from a group of freshwater <a href="/wiki/Green_algae" title="Green algae">green algae</a>, perhaps as early as 850 mya,<a href="#cite_note-Knauth-2009-3">[3]</a> but algae-like plants might have evolved as early as 1 billion years ago.<a href="#cite_note-Strother-2011-2">[2]</a> The closest living relatives of land plants are the <a href="/wiki/Charophyte" class="mw-redirect" title="Charophyte">charophytes</a>, specifically <a href="/wiki/Charales" title="Charales">Charales</a>; if modern Charales are similar to the distant ancestors they share with land plants, this means that the land plants evolved from a branched, filamentous alga dwelling in shallow fresh water,<a href="#cite_note-Kenrick-1997-5">[5]</a> perhaps at the edge of seasonally desiccating pools.<a href="#cite_note-Raven-2001-6">[6]</a> However, some recent evidence suggests that land plants might have originated from unicellular terrestrial <a href="/wiki/Charophyte" class="mw-redirect" title="Charophyte">charophytes</a> similar to extant <a href="/wiki/Klebsormidiophyceae" class="mw-redirect" title="Klebsormidiophyceae">Klebsormidiophyceae</a>.<a href="#cite_note-Del-Bem-2018-7">[7]</a> The alga would have had a <a href="/wiki/Biological_life_cycle#Haplontic_life_cycle" title="Biological life cycle">haplontic life cycle</a>. It would only very briefly have had paired <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> (the <a href="/wiki/Polyploidy#diploid" title="Polyploidy">diploid</a> condition) when the <a href="/wiki/Ovum" class="mw-redirect" title="Ovum">egg</a> and <a href="/wiki/Sperm" title="Sperm">sperm</a> first fused to form a <a href="/wiki/Zygote" title="Zygote">zygote</a> that would have immediately divided by <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> to produce cells with half the number of unpaired chromosomes (the <a href="/wiki/Polyploidy#haploid" title="Polyploidy">haploid</a> condition). <a href="/wiki/Symbiosis" title="Symbiosis">Co-operative interactions</a> with <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> may have helped early plants adapt to the stresses of the terrestrial realm.<a href="#cite_note-Heckman-2001-8">[8]</a> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Devonianscene-green.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/ba/Devonianscene-green.jpg/200px-Devonianscene-green.jpg" decoding="async" width="200" height="140" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/ba/Devonianscene-green.jpg/300px-Devonianscene-green.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/ba/Devonianscene-green.jpg/400px-Devonianscene-green.jpg 2x" data-file-width="2563" data-file-height="1792" /></a><figcaption>The Devonian marks the beginning of extensive land colonization by plants, which – through their effects on erosion and sedimentation – brought about significant climatic change.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Challenges_to_land_colonization">Challenges to land colonization</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=3" title="Edit section: Challenges to land colonization">edit</a>]</div> Plants were not the first photosynthesisers on land. Weathering rates suggest that organisms capable of photosynthesis were already living on the land <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1,200">1,200</a> million years ago,<a href="#cite_note-Raven-2001-6">[6]</a> and microbial fossils have been found in freshwater lake deposits from <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1,000">1,000</a> million years ago,<a href="#cite_note-9">[9]</a> but the <a href="/wiki/Isotopic_signature#Carbon_isotopes" title="Isotopic signature">carbon isotope record</a> suggests that they were too scarce to impact the atmospheric composition until around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=850">850</a> million years ago.<a href="#cite_note-Knauth-2009-3">[3]</a> These organisms, although phylogenetically diverse,<a href="#cite_note-Battistuzzi-2004-10">[10]</a> were probably small and simple, forming little more than an algal scum.<a href="#cite_note-Raven-2001-6">[6]</a> Since <a href="/wiki/Lichen" title="Lichen">lichens</a> initiate the first step in primary <a href="/wiki/Ecological_succession" title="Ecological succession">ecological succession</a> in contemporary contexts, one hypothesis has been that lichens came on land first and facilitated colonization by plants; however, both molecular phylogenies and the fossil record seem to contradict this.<a href="#cite_note-11">[11]</a> There are multiple potential reasons for why it took so long for land plants to emerge. It could be that atmospheric 'poisoning' prevented eukaryotes from colonising the land prior to the emergence of land plants,<a href="#cite_note-Kump-2005-12">[12]</a> or it could simply have taken a great time for the necessary complexity to evolve.<a href="#cite_note-Butterfield-2009-13">[13]</a> A major challenge to land adaptation would have been the absence of appropriate <a href="/wiki/Soil" title="Soil">soil</a>.<a href="#cite_note-14">[14]</a> Throughout the fossil record, soil is preserved, giving information on what early soils were like. Before land plants, the soil on land was poor in resources essential for life like <a href="/wiki/Nitrogen" title="Nitrogen">nitrogen</a> and <a href="/wiki/Phosphorus" title="Phosphorus">phosphorus</a> and had little capacity for holding water. <div class="mw-heading mw-heading3"><h3 id="Adaptations_to_land_colonization">Adaptations to land colonization</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=4" title="Edit section: Adaptations to land colonization">edit</a>]</div> Evidence of the earliest land plants occurs at about <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=470">470</a> million years ago, in lower middle Ordovician rocks from <a href="/wiki/Saudi_Arabia" title="Saudi Arabia">Saudi Arabia</a><a href="#cite_note-Strother-1996-15">[15]</a> and <a href="/wiki/Gondwana" title="Gondwana">Gondwana</a><a href="#cite_note-Rubinstein-2010-16">[16]</a> in the form of spores known as <a href="/wiki/Cryptospores" class="mw-redirect" title="Cryptospores">cryptospores</a>. These spores have walls made of <a href="/wiki/Sporopollenin" title="Sporopollenin">sporopollenin</a>, an extremely decay-resistant material that means they are well-preserved by the fossil record. These spores were produced either singly (monads), in pairs (dyads) or groups of four (tetrads), and their microstructure resembles that of modern <a href="/wiki/Marchantiophyta" title="Marchantiophyta">liverwort</a> spores, suggesting they share an equivalent grade of organisation.<a href="#cite_note-Wellman-2003-17">[17]</a> Their walls contain <a href="/wiki/Sporopollenin" title="Sporopollenin">sporopollenin</a> – further evidence of an embryophytic affinity.<a href="#cite_note-Steemans-2010-18">[18]</a> Trilete spores similar to those of <a href="/wiki/Tracheophytes" class="mw-redirect" title="Tracheophytes">vascular plants</a> appear soon afterwards, in Upper Ordovician rocks about 455 million years ago.<a href="#cite_note-Steemans-2009-19">[19]</a><a href="#cite_note-Rubinstein-2019-20">[20]</a> Depending exactly when the tetrad splits, each of the four spores may bear a "trilete mark", a Y-shape, reflecting the points at which each cell squashed up against its neighbours.<a href="#cite_note-Gray-1985-21">[21]</a> However, this requires that the spore walls be sturdy and resistant at an early stage. This resistance is closely associated with having a desiccation-resistant outer wall—a trait only of use when spores must survive out of water. Indeed, even those <a href="/wiki/Embryophytes" class="mw-redirect" title="Embryophytes">embryophytes</a> that have returned to the water lack a resistant wall, thus don't bear trilete marks.<a href="#cite_note-Gray-1985-21">[21]</a> A close examination of algal spores shows that none have trilete spores, either because their walls are not resistant enough, or, in those rare cases where they are, because the spores disperse before they are compressed enough to develop the mark or do not fit into a tetrahedral tetrad.<a href="#cite_note-Gray-1985-21">[21]</a> The earliest megafossils of land plants were <a href="/wiki/Thallus" title="Thallus">thalloid</a> organisms, which dwelt in fluvial wetlands and are found to have covered most of an <a href="/wiki/Llandovery_epoch" class="mw-redirect" title="Llandovery epoch">early Silurian</a> flood plain. They could only survive when the land was waterlogged.<a href="#cite_note-22">[22]</a> There were also microbial mats.<a href="#cite_note-Tomescu-2008-23">[23]</a> Once plants had reached the land, there were two approaches to dealing with desiccation. Modern bryophytes either avoid it or give in to it, restricting their ranges to moist settings or drying out and putting their metabolism "on hold" until more water arrives, as in the liverwort genus <a href="/wiki/Targionia_(plant)" title="Targionia (plant)">Targionia</a>. <a href="/wiki/Tracheophytes" class="mw-redirect" title="Tracheophytes">Tracheophytes</a> resist desiccation by controlling the rate of water loss. They all bear a waterproof outer <a href="/wiki/Plant_cuticle" title="Plant cuticle">cuticle</a> layer wherever they are exposed to air (as do some bryophytes), to reduce water loss, but since a total covering would cut them off from CO2 in the atmosphere tracheophytes use variable openings, the <a href="/wiki/Stomata" class="mw-redirect" title="Stomata">stomata</a>, to regulate the rate of gas exchange. Tracheophytes also developed vascular tissue to aid in the movement of water within the organisms (see <a href="#xylem">below</a>), and moved away from a gametophyte dominated life cycle (see <a href="#Changing_life_cycles">below</a>). Vascular tissue ultimately also facilitated upright growth without the support of water and paved the way for the evolution of larger plants on land.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Consequences">Consequences</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=5" title="Edit section: Consequences">edit</a>]</div> A global <a href="/wiki/Ice_age" title="Ice age">glaciation event</a> called <a href="/wiki/Snowball_Earth" title="Snowball Earth">Snowball Earth</a>, from around 720-635 mya in the <a href="/wiki/Cryogenian" title="Cryogenian">Cryogenian</a> period, is believed to have been at least partially caused by early photosynthetic organisms, which reduced the concentration of carbon dioxide and decreased the <a href="/wiki/Greenhouse_effect" title="Greenhouse effect">greenhouse effect</a> in the atmosphere,<a href="#cite_note-24">[24]</a> leading to an <a href="/wiki/Icehouse_Earth" class="mw-redirect" title="Icehouse Earth">icehouse climate</a>. Based on <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clock</a> studies of the previous decade or so, a 2022 study observed that the estimated time for the origin of the multicellular <a href="/wiki/Streptophyta" title="Streptophyta">streptophytes</a> (all except the unicellular basal clade <a href="/wiki/Mesostigmatophyceae" title="Mesostigmatophyceae">Mesostigmatophyceae</a>) fell in the cool Cryogenian while that of the subsequent separation of streptophytes fell in the warm <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a>, which they interpreted as an indication of <a href="/wiki/Selective_pressure" class="mw-redirect" title="Selective pressure">selective pressure</a> by the glacial period to the photosynthesizing organisms, a group of which succeeded in surviving in relatively warmer environments that remained habitable, subsequently flourishing in the later Ediacaran and <a href="/wiki/Phanerozoic" title="Phanerozoic">Phanerozoic</a> on land as embryophytes. The study also theorized that the unicellular morphology and other unique features of the <a href="/wiki/Zygnematophyceae" title="Zygnematophyceae">Zygnematophyceae</a> may reflect further adaptations to a <a href="/wiki/Cryophilic" class="mw-redirect" title="Cryophilic">cold loving</a> life style.<a href="#cite_note-Žárský-2022-25">[25]</a> The establishment of a land-based flora increased the rate of accumulation of oxygen in the atmosphere, as the land plants produced oxygen as a waste product. When this concentration rose above 13%, around 0.45 billion years ago,<a href="#cite_note-26">[26]</a> wildfires became possible, evident from charcoal in the fossil record.<a href="#cite_note-27">[27]</a> Apart from a controversial gap in the Late Devonian, charcoal has been present ever since. Charcoalification is an important <a href="/wiki/Taphonomy" title="Taphonomy">taphonomic</a> mode. Wildfire or burial in hot volcanic ash drives off the volatile compounds, leaving only a residue of pure carbon. This is not a viable food source for fungi, herbivores or detritovores, so it is prone to preservation. It is also robust and can withstand pressure, displaying exquisite, sometimes sub-cellular, detail in remains.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] In addition to the advent of charcoal in the rock record, the terrestrialization of plants has made significant contributions to changes in geology and landscapes. The Ordovician and Silurian show a 1.4 times greater proportion of mudrock in the geologic record than the previous 90% of earth's history and this increase in mudrock is considered to be a result of land plants retaining muds in a terrestrial setting. <a href="#cite_note-28">[28]</a> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_life_cycles">Evolution of life cycles</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=6" title="Edit section: Evolution of life cycles">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Alternation_of_generations" title="Alternation of generations">Alternation of generations</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Angiosperm_life_cycle_diagram-en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Angiosperm_life_cycle_diagram-en.svg/330px-Angiosperm_life_cycle_diagram-en.svg.png" decoding="async" width="330" height="380" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Angiosperm_life_cycle_diagram-en.svg/495px-Angiosperm_life_cycle_diagram-en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/28/Angiosperm_life_cycle_diagram-en.svg/660px-Angiosperm_life_cycle_diagram-en.svg.png 2x" data-file-width="500" data-file-height="575" /></a><figcaption><a href="/wiki/Angiosperm" class="mw-redirect" title="Angiosperm">Angiosperm</a> life cycle</figcaption></figure> All multicellular plants have a life cycle comprising two generations or phases. The <a href="/wiki/Gametophyte" title="Gametophyte">gametophyte</a> phase has a single set of chromosomes (denoted 1n) and produces gametes (sperm and eggs). The <a href="/wiki/Sporophyte" title="Sporophyte">sporophyte</a> phase has paired chromosomes (denoted 2n) and produces spores. The gametophyte and sporophyte phases may be homomorphic, appearing identical in some algae, such as <a href="/wiki/Ulva_lactuca#Life_history" title="Ulva lactuca">Ulva lactuca</a>, but are very different in all modern land plants, a condition known as <a href="/wiki/Gametophyte#Heterospory" title="Gametophyte">heteromorphy</a>.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The pattern in plant evolution has been a shift from homomorphy to heteromorphy. The algal ancestors of land plants were almost certainly <a href="https://en.wiktionary.org/wiki/haplobiontic" class="extiw" title="wikt:haplobiontic">haplobiontic</a>, being haploid for all their life cycles, with a unicellular zygote providing the 2N stage. All land plants (i.e. <a href="/wiki/Embryophytes" class="mw-redirect" title="Embryophytes">embryophytes</a>) are <a href="https://en.wiktionary.org/wiki/diplobiontic" class="extiw" title="wikt:diplobiontic">diplobiontic</a> – that is, both the haploid and diploid stages are multicellular.<a href="#cite_note-Stewart-1993-1">[1]</a> Two trends are apparent: <a href="/wiki/Bryophyte" title="Bryophyte">bryophytes</a> (<a href="/wiki/Marchantiophyta" title="Marchantiophyta">liverworts</a>, <a href="/wiki/Moss" title="Moss">mosses</a> and <a href="/wiki/Hornwort" title="Hornwort">hornworts</a>) have developed the gametophyte as the dominant phase of the life cycle, with the sporophyte becoming almost entirely dependent on it; <a href="/wiki/Vascular_plant" title="Vascular plant">vascular plants</a> have developed the sporophyte as the dominant phase, with the gametophytes being particularly reduced in the <a href="/wiki/Seed_plant" title="Seed plant">seed plants</a>.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] It has been proposed as the basis for the emergence of the diploid phase of the life cycle as the dominant phase that diploidy allows masking of the expression of deleterious mutations through <a href="/wiki/Complementation_(genetics)" title="Complementation (genetics)">genetic complementation</a>.<a href="#cite_note-29">[29]</a><a href="#cite_note-30">[30]</a> Thus if one of the parental genomes in the diploid cells contains <a href="/wiki/Mutation" title="Mutation">mutations</a> leading to defects in one or more <a href="/wiki/Gene_product" title="Gene product">gene products</a>, these deficiencies could be compensated for by the other parental genome (which nevertheless may have its own defects in other genes). As the diploid phase was becoming predominant, the masking effect likely allowed <a href="/wiki/Genome_size" title="Genome size">genome size</a>, and hence information content, to increase without the constraint of having to improve accuracy of replication. The opportunity to increase information content at low cost is advantageous because it permits new adaptations to be encoded. This view has been challenged, with evidence showing that selection is no more effective in the haploid than in the diploid phases of the lifecycle of mosses and angiosperms.<a href="#cite_note-Szövényi-2013-31">[31]</a> There are two competing theories to explain the appearance of a diplobiontic lifecycle.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The <a href="/wiki/Interpolation_theory" title="Interpolation theory">interpolation theory</a> (also known as the antithetic or intercalary theory)<a href="#cite_note-Boyce-2008-32">[32]</a> holds that the interpolation of a multicellular sporophyte phase between two successive gametophyte generations was an innovation caused by preceding meiosis in a freshly germinated zygote with one or more rounds of mitotic division, thereby producing some diploid multicellular tissue before finally meiosis produced spores. This theory implies that the first sporophytes bore a very different and simpler morphology to the gametophyte they depended on.<a href="#cite_note-Boyce-2008-32">[32]</a> This seems to fit well with what is known of the bryophytes, in which a vegetative thalloid gametophyte nurtures a simple sporophyte, which consists of little more than an unbranched sporangium on a stalk. Increasing complexity of the ancestrally simple sporophyte, including the eventual acquisition of photosynthetic cells, would free it from its dependence on a gametophyte, as seen in some hornworts (<a href="/wiki/Anthoceros" title="Anthoceros">Anthoceros</a>), and eventually result in the sporophyte developing organs and vascular tissue, and becoming the dominant phase, as in the tracheophytes (vascular plants).<a href="#cite_note-Stewart-1993-1">[1]</a> This theory may be supported by observations that smaller <a href="/wiki/Cooksonia" title="Cooksonia">Cooksonia</a> individuals must have been supported by a gametophyte generation. The observed appearance of larger axial sizes, with room for photosynthetic tissue and thus self-sustainability, provides a possible route for the development of a self-sufficient sporophyte phase.<a href="#cite_note-Boyce-2008-32">[32]</a> The alternative hypothesis, called the transformation theory (or homologous theory), posits that the sporophyte might have appeared suddenly by delaying the occurrence of meiosis until a fully developed multicellular sporophyte had formed. Since the same genetic material would be employed by both the haploid and diploid phases, they would look the same. This explains the behaviour of some algae, such as Ulva lactuca, which produce alternating phases of identical sporophytes and gametophytes. Subsequent adaption to the desiccating land environment, which makes sexual reproduction difficult, might have resulted in the simplification of the sexually active gametophyte, and elaboration of the sporophyte phase to better disperse the waterproof spores.<a href="#cite_note-Stewart-1993-1">[1]</a> The tissue of sporophytes and gametophytes of vascular plants such as <a href="/wiki/Rhynia" title="Rhynia">Rhynia</a> preserved in the <a href="/wiki/Rhynie_chert" title="Rhynie chert">Rhynie chert</a> is of similar complexity, which is taken to support this hypothesis.<a href="#cite_note-Boyce-2008-32">[32]</a><a href="#cite_note-Kerp-2004-33">[33]</a><a href="#cite_note-Taylor-2005b-34">[34]</a> By contrast, modern vascular plants, with the exception of Psilotum, have heteromorphic sporophytes and gametophytes in which the gametophytes rarely have any vascular tissue.<a href="#cite_note-Holloway-1939-35">[35]</a> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_plant_anatomy">Evolution of plant anatomy</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=7" title="Edit section: Evolution of plant anatomy">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Arbuscular_mycorrhizal_symbiosis">Arbuscular mycorrhizal symbiosis</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=8" title="Edit section: Arbuscular mycorrhizal symbiosis">edit</a>]</div> There is no evidence that early land plants of the Silurian and early Devonian had roots, although fossil evidence of rhizoids occurs for several species, such as <a href="/wiki/Horneophyton" title="Horneophyton">Horneophyton</a>. The earliest land plants did not have vascular systems for transport of water and nutrients either. <a href="/wiki/Aglaophyton" title="Aglaophyton">Aglaophyton</a>, a rootless vascular plant known from Devonian fossils in the <a href="/wiki/Rhynie_chert" title="Rhynie chert">Rhynie chert</a><a href="#cite_note-Kidston-1920-36">[36]</a> was the first land plant discovered to have had a <a href="/wiki/Symbiotic" class="mw-redirect" title="Symbiotic">symbiotic</a> relationship with fungi <a href="#cite_note-Remy-1994-37">[37]</a> which formed <a href="/wiki/Arbuscular_mycorrhiza" title="Arbuscular mycorrhiza">arbuscular mycorrhizas</a>, literally "tree-like fungal roots", in a well-defined cylinder of cells (ring in cross section) in the cortex of its stems. The fungi fed on the plant's sugars, in exchange for nutrients generated or extracted from the soil (especially <a href="/wiki/Phosphate" title="Phosphate">phosphate</a>), to which the plant would otherwise have had no access. Like other rootless land plants of the Silurian and early Devonian Aglaophyton may have relied on arbuscular mycorrhizal fungi for acquisition of water and nutrients from the soil.<a href="#cite_note-38">[38]</a> The fungi were of the phylum <a href="/wiki/Glomeromycota" title="Glomeromycota">Glomeromycota</a>,<a href="#cite_note-Schussler-2001-39">[39]</a> a group that probably first appeared 1 billion years ago and still forms arbuscular mycorrhizal associations today with all major land plant groups from bryophytes to pteridophytes, gymnosperms and angiosperms and with more than 80% of vascular plants.<a href="#cite_note-Smith-2008-40">[40]</a> Evidence from DNA sequence analysis indicates that the arbuscular mycorrhizal mutualism arose in the common ancestor of these land plant groups during their transition to land<a href="#cite_note-Simon-1993b-41">[41]</a> and it may even have been the critical step that enabled them to colonise the land.<a href="#cite_note-Simon-1993a-42">[42]</a> Appearing as they did before these plants had evolved roots, mycorrhizal fungi would have assisted plants in the acquisition of water and mineral nutrients such as <a href="/wiki/Phosphorus" title="Phosphorus">phosphorus</a>, in exchange for organic compounds which they could not synthesize themselves.<a href="#cite_note-Smith-2008-40">[40]</a> Such fungi increase the productivity even of simple plants such as liverworts.<a href="#cite_note-Humphreys-2010-43">[43]</a><a href="#cite_note-Brundrett-2002-44">[44]</a> <div class="mw-heading mw-heading3"><h3 id="Cuticle,_stomata_and_intercellular_spaces">Cuticle, stomata and intercellular spaces</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=9" title="Edit section: Cuticle, stomata and intercellular spaces">edit</a>]</div> To photosynthesise, plants must absorb CO2 from the atmosphere. However, making the tissues available for CO2 to enter allows water to evaporate, so this comes at a price.<a href="#cite_note-Sperry-2003-45">[45]</a> Water is lost much faster than CO2 is absorbed, so plants need to replace it. Early land plants transported water <a href="/wiki/Apoplast" title="Apoplast">apoplastically</a>, within the porous walls of their cells. Later, they evolved three anatomical features that provided the ability to control the inevitable water loss that accompanied  CO2 acquisition.  First, a waterproof outer covering or <a href="/wiki/Plant_cuticle" title="Plant cuticle">cuticle</a> evolved that reduced water loss. Secondly, variable apertures, the <a href="/wiki/Stomata" class="mw-redirect" title="Stomata">stomata</a> that could open and close to regulate the amount of water lost by evaporation during CO2 uptake and thirdly intercellular space between photosynthetic parenchyma cells that allowed improved internal distribution of the CO2 to the chloroplasts. This three-part system provided improved homoiohydry, the regulation of water content of the tissues, providing a particular advantage when water supply is not constant.<a href="#cite_note-Raven-1977-46">[46]</a> The high CO2 concentrations of the Silurian and early Devonian, when plants were first colonising land, meant that they used water relatively efficiently. As CO2 was withdrawn from the atmosphere by plants, more water was lost in its capture, and more elegant water acquisition and transport mechanisms evolved.<a href="#cite_note-Sperry-2003-45">[45]</a> Plants growing upwards into the air needed a system for transporting water from the soil to all the different parts of the above-soil plant, especially to photosynthesising parts. By the end of the <a href="/wiki/Carboniferous" title="Carboniferous">Carboniferous</a>, when CO2 concentrations had been reduced to something approaching that of today, around 17 times more water was lost per unit of CO2 uptake.<a href="#cite_note-Sperry-2003-45">[45]</a> However, even in the "easy" early days, water was always at a premium, and had to be transported to parts of the plant from the wet soil to avoid desiccation.<a href="#cite_note-Raven-1977-46">[46]</a> Water can be wicked by <a href="/wiki/Capillary_action" title="Capillary action">capillary action</a> along a fabric with small spaces. In narrow columns of water, such as those within the plant cell walls or in tracheids, when molecules evaporate from one end, they pull the molecules behind them along the channels. Therefore, evaporation alone provides the driving force for water transport in plants.<a href="#cite_note-Sperry-2003-45">[45]</a>  However, without specialized transport vessels, this <a href="/wiki/Xylem#Cohesion-tension_theory" title="Xylem">cohesion-tension</a> mechanism can cause negative pressures sufficient to collapse water conducting cells, limiting the transport water to no more than a few cm, and therefore limiting the size of the earliest plants.<a href="#cite_note-Sperry-2003-45">[45]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Banded_tube.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Banded_tube.jpg/220px-Banded_tube.jpg" decoding="async" width="220" height="89" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Banded_tube.jpg/330px-Banded_tube.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/96/Banded_tube.jpg/440px-Banded_tube.jpg 2x" data-file-width="805" data-file-height="325" /></a><figcaption>A <a href="/wiki/Banded_tube" title="Banded tube">banded tube</a> from the Late Silurian/Early Devonian. The bands are difficult to see on this specimen, as an opaque carbonaceous coating conceals much of the tube. Bands are just visible in places on the left half of the image. Scale bar: 20 μm</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Xylem">Xylem</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=10" title="Edit section: Xylem">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Xylem" title="Xylem">Xylem</a></div> To be free from the constraints of small size and constant moisture that the <a href="/wiki/Ground_tissue#Parenchyma" title="Ground tissue">parenchymatic</a> transport system inflicted, plants needed a more efficient water transport system.  As plants grew upwards, specialised water transport <a href="/wiki/Vascular_tissue" title="Vascular tissue">vascular tissues</a> evolved, first in the form of simple <a href="/wiki/Hydroid_(botany)" title="Hydroid (botany)">hydroids</a> of the type found in the setae of moss sporophytes. These simple elongated cells were dead and water-filled at maturity, providing a channel for water transport, but their thin, unreinforced walls would collapse under modest water tension, limiting the plant height.  <a href="/wiki/Xylem" title="Xylem">Xylem</a> <a href="/wiki/Tracheid" title="Tracheid">tracheids</a>, wider cells with <a href="/wiki/Lignin" title="Lignin">lignin</a>-reinforced cell walls that were more resistant to collapse under the tension caused by water stress, occur  in more than one plant group by mid-Silurian, and may have a single evolutionary origin, possibly within the hornworts,<a href="#cite_note-Qiu-2006-47">[47]</a> uniting all tracheophytes.  Alternatively,  they may have evolved more than once.<a href="#cite_note-Sperry-2003-45">[45]</a>  Much later, in the Cretaceous, tracheids were followed by <a href="/wiki/Vessel_element" title="Vessel element">vessels</a> in <a href="/wiki/Flowering_plant" title="Flowering plant">flowering plants</a>.<a href="#cite_note-Sperry-2003-45">[45]</a> As water transport mechanisms and waterproof cuticles evolved, plants could survive without being continually covered by a film of water. This transition from <a href="/wiki/Poikilohydry" title="Poikilohydry">poikilohydry</a> to <a href="/wiki/Homoiohydry" title="Homoiohydry">homoiohydry</a> opened up new potential for colonisation.<a href="#cite_note-Sperry-2003-45">[45]</a><a href="#cite_note-Raven-1977-46">[46]</a> The early Devonian pretracheophytes <a href="/wiki/Aglaophyton" title="Aglaophyton">Aglaophyton</a> and <a href="/wiki/Horneophyton" title="Horneophyton">Horneophyton</a> have unreinforced water transport tubes with wall structures very similar to moss hydroids, but they grew alongside several species of <a href="/wiki/Tracheophytes" class="mw-redirect" title="Tracheophytes">tracheophytes</a>, such as <a href="/wiki/Rhynia" title="Rhynia">Rhynia</a> gwynne-vaughanii that had xylem tracheids that were well reinforced by bands of lignin. The earliest macrofossils known to have xylem tracheids are small, mid-Silurian plants of the genus <a href="/wiki/Cooksonia" title="Cooksonia">Cooksonia</a>.<a href="#cite_note-Edwards-1992-48">[48]</a> However, thickened bands on the walls of isolated tube fragments are apparent from the early Silurian onwards.<a href="#cite_note-Niklas-1983-49">[49]</a> Plants continued to innovate ways of reducing the resistance to flow within their cells, progressively increasing the efficiency of their water transport and to increase the resistance of the tracheids to collapse under tension.<a href="#cite_note-Niklas-1985-50">[50]</a><a href="#cite_note-Niklas-1980-51">[51]</a> During the early Devonian, maximum tracheid diameter increased with time, but may have plateaued in the <a href="/wiki/Zosterophyll" title="Zosterophyll">zosterophylls</a> by mid-Devonian.<a href="#cite_note-Niklas-1985-50">[50]</a>  Overall transport rate also depends on the overall cross-sectional area of the xylem bundle itself, and some mid-Devonian plants, such as the Trimerophytes, had much larger steles than their early ancestors.<a href="#cite_note-Niklas-1985-50">[50]</a> While wider tracheids provided higher rates of water transport, they increased the risk of cavitation, the formation of air bubbles resulting from the breakage of the water column under tension.<a href="#cite_note-Sperry-2003-45">[45]</a> Small pits in tracheid walls allow water to by-pass a defective tracheid while preventing air bubbles from passing through<a href="#cite_note-Sperry-2003-45">[45]</a> but at the cost of restricted flow rates. By the Carboniferous, Gymnosperms had developed <a href="/wiki/Pit_(botany)#Types_of_pits" title="Pit (botany)">bordered pits</a>,<a href="#cite_note-Pitterman-2005-52">[52]</a><a href="#cite_note-53">[53]</a> valve-like structures that allow high-conductivity  pits to seal when one side of a tracheid is depressurized. Tracheids have non-perforated end walls with pits, which impose a great deal of resistance on water flow,<a href="#cite_note-Niklas-1985-50">[50]</a> but may have had the advantage of isolating air embolisms caused by cavitation or freezing. Vessels first evolved during the dry, low CO2 periods of the Late Permian, in the horsetails, ferns and <a href="/wiki/Selaginellales" class="mw-redirect" title="Selaginellales">Selaginellales</a> independently, and later appeared in the mid Cretaceous in gnetophytes and angiosperms.<a href="#cite_note-Sperry-2003-45">[45]</a>  Vessel members are open tubes with no end walls, and are arranged end to end to operate as if they were one continuous vessel.<a href="#cite_note-Niklas-1985-50">[50]</a> Vessels allowed the same cross-sectional area of wood to transport much more water than tracheids.<a href="#cite_note-Sperry-2003-45">[45]</a> This allowed plants to fill more of their stems with structural <a href="/wiki/Ground_tissue#Fibres" title="Ground tissue">fibres</a> and also opened a new niche to vines, which could transport water without being as thick as the tree they grew on.<a href="#cite_note-Sperry-2003-45">[45]</a> Despite these advantages, tracheid-based wood is a lot lighter, thus cheaper to make, as vessels need to be much more reinforced to avoid cavitation.<a href="#cite_note-Sperry-2003-45">[45]</a> Once plants had evolved this level of control over water evaporation and water transport, they were truly <a href="/wiki/Homoiohydry" title="Homoiohydry">homoiohydric</a>, able to extract water from their environment through root-like organs rather than relying on a film of surface moisture, enabling them to grow to much greater size<a href="#cite_note-Raven-1977-46">[46]</a><a href="#cite_note-Sperry-2003-45">[45]</a> but as a result of their increased independence from their surroundings, most vascular plants lost their ability to survive desiccation - a costly trait to lose.<a href="#cite_note-Sperry-2003-45">[45]</a> In early land plants, support was mainly provided by turgor pressure, particularly of the outer layer of cells known as the sterome tracheids, and not by the xylem, which was too small, too weak and in too central a position to provide much structural support.<a href="#cite_note-Sperry-2003-45">[45]</a>  Plants with <a href="/wiki/Secondary_xylem" class="mw-redirect" title="Secondary xylem">secondary xylem</a> that  had appeared by mid-Devonian, such as the <a href="/wiki/Trimerophytes" class="mw-redirect" title="Trimerophytes">Trimerophytes</a> and <a href="/wiki/Progymnosperms" class="mw-redirect" title="Progymnosperms">Progymnosperms</a> had much larger vascular cross sections producing strong woody tissue.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Endodermis">Endodermis</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=11" title="Edit section: Endodermis">edit</a>]</div> An <a href="/wiki/Endodermis" title="Endodermis">endodermis</a> may have evolved in the earliest plant roots during the Devonian, but the first fossil evidence for such a structure is Carboniferous.<a href="#cite_note-Sperry-2003-45">[45]</a> The endodermis in the roots surrounds the water transport tissue and regulates ion exchange between the groundwater and the tissues and prevents unwanted pathogens etc. from entering the water transport system. The endodermis can also provide an upwards pressure, forcing water out of the roots when transpiration is not enough of a driver. <div class="mw-heading mw-heading2"><h2 id="Evolution_of_plant_morphology">Evolution of plant morphology</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=12" title="Edit section: Evolution of plant morphology">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Leaves">Leaves</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=13" title="Edit section: Leaves">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Illustration_Isoetes_lacustris0.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Illustration_Isoetes_lacustris0.jpg/220px-Illustration_Isoetes_lacustris0.jpg" decoding="async" width="220" height="369" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Illustration_Isoetes_lacustris0.jpg/330px-Illustration_Isoetes_lacustris0.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Illustration_Isoetes_lacustris0.jpg/440px-Illustration_Isoetes_lacustris0.jpg 2x" data-file-width="1385" data-file-height="2325" /></a><figcaption>The lycopod <a href="/wiki/Isoetes" title="Isoetes">Isoetes</a> bears <a href="/wiki/Microphyll" class="mw-redirect" title="Microphyll">microphylls</a> (leaves with a single vascular trace).</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Vein_sceleton_hydrangea_ies.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/27/Vein_sceleton_hydrangea_ies.jpg/220px-Vein_sceleton_hydrangea_ies.jpg" decoding="async" width="220" height="193" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/27/Vein_sceleton_hydrangea_ies.jpg/330px-Vein_sceleton_hydrangea_ies.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/27/Vein_sceleton_hydrangea_ies.jpg/440px-Vein_sceleton_hydrangea_ies.jpg 2x" data-file-width="800" data-file-height="700" /></a><figcaption>The branching pattern of megaphyll veins may indicate their origin as webbed, dichotomising branches.</figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Leaf_1_web.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Leaf_1_web.jpg/250px-Leaf_1_web.jpg" decoding="async" width="250" height="188" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Leaf_1_web.jpg/375px-Leaf_1_web.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Leaf_1_web.jpg/500px-Leaf_1_web.jpg 2x" data-file-width="1024" data-file-height="768" /></a><figcaption>Leaf lamina. The megaphyllous <a href="/wiki/Leaf" title="Leaf">leaf</a> architecture arose multiple times in different plant lineages</figcaption></figure> Leaves are the primary <a href="/wiki/Photosynthetic" class="mw-redirect" title="Photosynthetic">photosynthetic</a> organs of a modern plant. The origin of <a href="/wiki/Leaves" class="mw-redirect" title="Leaves">leaves</a> was almost certainly triggered by falling concentrations of atmospheric CO2 during the Devonian period, increasing the efficiency with which carbon dioxide could be captured for <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>.<a href="#cite_note-Kenrick-2001-54">[54]</a><a href="#cite_note-Beerling-2001-55">[55]</a> Leaves evolved more than once. Based on their structure, they are classified into two types: <a href="/wiki/Microphylls" class="mw-redirect" title="Microphylls">microphylls</a>, which lack complex venation and may have originated as spiny outgrowths known as enations, and <a href="/wiki/Megaphylls" class="mw-redirect" title="Megaphylls">megaphylls</a>, which are large and have complex <a href="/wiki/Leaf#Venation" title="Leaf">venation</a> that may have arisen from the modification of groups of branches. It has been proposed that these structures arose independently.<a href="#cite_note-Crane-1997-56">[56]</a> Megaphylls, according to Walter Zimmerman's telome theory,<a href="#cite_note-Zimmermann-1959-57">[57]</a> have evolved from plants that showed a three-dimensional branching architecture, through three transformations—overtopping, which led to the lateral position typical of leaves, planation, which involved formation of a planar architecture, webbing or fusion, which united the planar branches, thus leading to the formation of a proper <a href="/wiki/Leaf" title="Leaf">leaf</a> lamina. All three steps happened multiple times in the evolution of today's leaves.<a href="#cite_note-58">[58]</a> It is widely believed that the telome theory is well supported by fossil evidence. However, Wolfgang Hagemann questioned it for morphological and ecological reasons and proposed an alternative theory.<a href="#cite_note-59">[59]</a><a href="#cite_note-60">[60]</a> Whereas according to the telome theory the most primitive land plants have a three-dimensional branching system of radially symmetrical axes (telomes), according to Hagemann's alternative the opposite is proposed: the most primitive land plants that gave rise to vascular plants were flat, thalloid, leaf-like, without axes, somewhat like a liverwort or fern prothallus. Axes such as stems and roots evolved later as new organs. Rolf Sattler proposed an overarching process-oriented view that leaves some limited room for both the telome theory and Hagemann's alternative and in addition takes into consideration the whole continuum between dorsiventral (flat) and radial (cylindrical) structures that can be found in fossil and living land plants.<a href="#cite_note-61">[61]</a><a href="#cite_note-62">[62]</a> This view is supported by research in molecular genetics. Thus, James (2009)<a href="#cite_note-63">[63]</a> concluded that "it is now widely accepted that... radiality [characteristic of axes such as stems] and dorsiventrality [characteristic of leaves] are but extremes of a continuous spectrum. In fact, it is simply the timing of the KNOX gene expression". Before the evolution of <a href="/wiki/Leaves" class="mw-redirect" title="Leaves">leaves</a>, plants had the <a href="/wiki/Photosynthetic" class="mw-redirect" title="Photosynthetic">photosynthetic</a> apparatus on the stems, which they retain albeit leaves have largely assumed that job. Today's megaphyll leaves probably became commonplace some 360mya, about 40my after the simple leafless plants had colonized the land in the <a href="/wiki/Early_Devonian" title="Early Devonian">Early Devonian</a>. This spread has been linked to the fall in the atmospheric <a href="/wiki/Carbon_dioxide" title="Carbon dioxide">carbon dioxide</a> concentrations in the Late <a href="/wiki/Paleozoic" title="Paleozoic">Paleozoic</a> era associated with a rise in density of <a href="/wiki/Stomata" class="mw-redirect" title="Stomata">stomata</a> on leaf surface.<a href="#cite_note-Kenrick-2001-54">[54]</a> This would have resulted in greater <a href="/wiki/Transpiration" title="Transpiration">transpiration</a> rates and gas exchange, but especially at high CO2 concentrations, large leaves with fewer stomata would have heated to lethal temperatures in full sunlight. Increasing the stomatal density allowed for a better-cooled leaf, thus making its spread feasible, but increased <a href="/wiki/Carbon_dioxide" title="Carbon dioxide">CO2</a> uptake at the expense of decreased water use efficiency.<a href="#cite_note-Beerling-2001-55">[55]</a><a href="#cite_note-64">[64]</a> The rhyniophytes of the Rhynie chert consisted only of slender, unornamented axes. The early to middle Devonian <a href="/wiki/Trimerophyte" class="mw-redirect" title="Trimerophyte">trimerophytes</a> may be considered leafy. This group of vascular plants are recognisable by their masses of terminal sporangia, which adorn the ends of axes which may bifurcate or trifurcate.<a href="#cite_note-Stewart-1993-1">[1]</a> Some organisms, such as <a href="/wiki/Psilophyton" title="Psilophyton">Psilophyton</a>, bore <a href="https://en.wiktionary.org/wiki/enation" class="extiw" title="wikt:enation">enations</a>. These are small, spiny outgrowths of the stem, lacking their own vascular supply.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The <a href="/wiki/Zosterophyll" title="Zosterophyll">zosterophylls</a> were already important in the late Silurian, much earlier than any rhyniophytes of comparable complexity.<a href="#cite_note-Kotyk-2002-65">[65]</a> This group, recognisable by their kidney-shaped sporangia which grew on short lateral branches close to the main axes, sometimes branched in a distinctive H-shape.<a href="#cite_note-Stewart-1993-1">[1]</a> Many zosterophylls bore enations (small tissue outgrowths on the surface with variable morphologies) on their axes but none of these had a vascular trace. The first evidence of vascularised enations occurs in a fossil clubmoss known as <a href="/wiki/Baragwanathia" title="Baragwanathia">Baragwanathia</a> that had already appeared in the fossil record in the Late Silurian.<a href="#cite_note-Rickards-2000-66">[66]</a> In this organism, these leaf traces continue into the leaf to form their mid-vein.<a href="#cite_note-Kaplan-2001-67">[67]</a> One theory, the "enation theory", holds that the microphyllous leaves of clubmosses developed by outgrowths of the protostele connecting with existing enations<a href="#cite_note-Stewart-1993-1">[1]</a> The leaves of the Rhynie genus <a href="/wiki/Asteroxylon" title="Asteroxylon">Asteroxylon</a>, which was preserved in the Rhynie chert almost 20 million years later than Baragwanathia, had a primitive vascular supply – in the form of <a href="/w/index.php?title=Leaf_trace&action=edit&redlink=1" class="new" title="Leaf trace (page does not exist)">leaf traces</a> departing from the central protostele towards each individual "leaf".<a href="#cite_note-Taylor-2005a-68">[68]</a> Asteroxylon and Baragwanathia are widely regarded as primitive lycopods,<a href="#cite_note-Stewart-1993-1">[1]</a> a group still <a href="https://en.wiktionary.org/wiki/extant" class="extiw" title="wikt:extant">extant</a> today, represented by the <a href="/wiki/Isoetes" title="Isoetes">quillworts</a>, the <a href="/wiki/Selaginella" title="Selaginella">spikemosses</a> and the <a href="/wiki/Club_moss" class="mw-redirect" title="Club moss">club mosses</a>. Lycopods bear distinctive <a href="/wiki/Microphylls" class="mw-redirect" title="Microphylls">microphylls</a>, defined as leaves with a single vascular trace. Microphylls could grow to some size, those of <a href="/wiki/Lepidodendrales" title="Lepidodendrales">Lepidodendrales</a> reaching over a meter in length, but almost all just bear the one vascular bundle. An exception is the rare branching in some <a href="/wiki/Selaginella" title="Selaginella">Selaginella</a> species. The more familiar leaves, <a href="/wiki/Megaphylls" class="mw-redirect" title="Megaphylls">megaphylls</a>, are thought to have originated four times independently: in the ferns, horsetails, progymnosperms and seed plants.<a href="#cite_note-Boyce-2002-69">[69]</a> They appear to have originated by modifying <a href="/w/index.php?title=Dichotomising&action=edit&redlink=1" class="new" title="Dichotomising (page does not exist)">dichotomising</a> branches, which first overlapped (or "overtopped") one another, became flattened or planated and eventually developed "webbing" and evolved gradually into more leaf-like structures.<a href="#cite_note-Kaplan-2001-67">[67]</a> Megaphylls, by <a href="/wiki/Walter_Max_Zimmermann" title="Walter Max Zimmermann">Zimmerman's</a> <a href="/wiki/Microphyll#Leaf_vasculature" class="mw-redirect" title="Microphyll">telome theory</a>, are composed of a group of webbed branches<a href="#cite_note-Kaplan-2001-67">[67]</a> and hence the "leaf gap" left where the leaf's vascular bundle leaves that of the main branch resembles two axes splitting.<a href="#cite_note-Kaplan-2001-67">[67]</a> In each of the four groups to evolve megaphylls, their leaves first evolved during the Late Devonian to Early Carboniferous, diversifying rapidly until the designs settled down in the mid Carboniferous.<a href="#cite_note-Boyce-2002-69">[69]</a> The cessation of further diversification can be attributed to developmental constraints,<a href="#cite_note-Boyce-2002-69">[69]</a> raising the question of why it took so long for leaves to evolve in the first place. Plants had been on land for at least 50 million years before megaphylls became significant. However, small, rare mesophylls are known from the early Devonian genus <a href="/wiki/Eophyllophyton" title="Eophyllophyton">Eophyllophyton</a> – so development could not have been a barrier to their appearance.<a href="#cite_note-Hao-2003-70">[70]</a> The best explanation so far is that atmospheric CO2 was declining rapidly during this time – falling by around 90% during the Devonian.<a href="#cite_note-Berner-2001-71">[71]</a> This required an increase in stomatal density by 100 times to maintain the rate of photosynthesis. When stomata open to allow water to evaporate from leaves it has a cooling effect, resulting from the loss of <a href="/wiki/Latent_heat" title="Latent heat">latent heat</a> of evaporation. It appears that the low stomatal density in the early Devonian meant that evaporation and evaporative cooling were limited, and that leaves would have overheated if they grew to any size. The stomatal density could not increase, as the primitive steles and limited root systems would not be able to supply water quickly enough to match the rate of transpiration.<a href="#cite_note-Beerling-2001-55">[55]</a> Clearly, leaves are not always beneficial, as illustrated by the frequent occurrence of secondary loss of leaves, exemplified by <a href="/wiki/Cacti" class="mw-redirect" title="Cacti">cacti</a> and the "whisk fern" <a href="/wiki/Psilotum" title="Psilotum">Psilotum</a>. Secondary evolution can disguise the true evolutionary origin of some leaves. Some genera of ferns display complex leaves which are attached to the pseudostele by an outgrowth of the vascular bundle, leaving no leaf gap.<a href="#cite_note-Kaplan-2001-67">[67]</a> <a href="/wiki/Deciduous" title="Deciduous">Deciduous</a> trees deal with another disadvantage to having leaves. The popular belief that plants shed their leaves when the days get too short is misguided; evergreens prospered in the <a href="/wiki/Arctic_Circle" title="Arctic Circle">Arctic Circle</a> during the <a href="/wiki/PETM" class="mw-redirect" title="PETM">most recent</a> <a href="/wiki/Greenhouse_earth" class="mw-redirect" title="Greenhouse earth">greenhouse earth</a>.<a href="#cite_note-Shellito-2006-72">[72]</a> The generally accepted reason for shedding leaves during winter is to cope with the weather – the force of wind and weight of snow are much more comfortably weathered without leaves to increase surface area. Seasonal leaf loss has evolved independently several times and is exhibited in the <a href="/wiki/Ginkgoales" title="Ginkgoales">ginkgoales</a>, some <a href="/wiki/Pinophyta" class="mw-redirect" title="Pinophyta">pinophyta</a> and certain angiosperms.<a href="#cite_note-Aerts-1995-73">[73]</a> Leaf loss may also have arisen as a response to pressure from insects; it may have been less costly to lose leaves entirely during the winter or dry season than to continue investing resources in their repair.<a href="#cite_note-Lababdeira-1994-74">[74]</a> <div class="mw-heading mw-heading3"><h3 id="Roots">Roots</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=14" title="Edit section: Roots">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:204px;max-width:204px"><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:Lepidodendron_aculeatum2.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/27/Lepidodendron_aculeatum2.jpg/200px-Lepidodendron_aculeatum2.jpg" decoding="async" width="200" height="133" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/27/Lepidodendron_aculeatum2.jpg/300px-Lepidodendron_aculeatum2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/2/27/Lepidodendron_aculeatum2.jpg 2x" data-file-width="400" data-file-height="265" /></a></div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:Lepido_root_top.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Lepido_root_top.jpg/200px-Lepido_root_top.jpg" decoding="async" width="200" height="140" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Lepido_root_top.jpg/300px-Lepido_root_top.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Lepido_root_top.jpg/400px-Lepido_root_top.jpg 2x" data-file-width="2250" data-file-height="1575" /></a></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">The rhizomes (<a href="/wiki/Stigmaria" title="Stigmaria">Stigmaria</a>) (bottom image) of lycopsids (including the Carboniferous <a href="/wiki/Lepidodendrales" title="Lepidodendrales">Lepidodendrales</a>) are thought to be developmentally equivalent to the stems (top), as the similar appearance of "leaf scars" and "root scars" on these specimens from different species demonstrates.</div></div></div></div> The evolution of roots had consequences on a global scale. By disturbing the soil and promoting its acidification (by taking up nutrients such as nitrate and phosphate<a href="#cite_note-75">[75]</a>), they enabled it to weather more deeply, injecting carbon compounds deeper into soils<a href="#cite_note-Mora-1996-76">[76]</a> with huge implications for climate.<a href="#cite_note-Berner-1994-77">[77]</a> These effects may have been so profound they led to <a href="/wiki/Late_Devonian_extinction" title="Late Devonian extinction">a mass extinction</a>.<a href="#cite_note-Algeo-1995-78">[78]</a> While there are traces of root-like impressions in fossil soils in the Late Silurian,<a href="#cite_note-Retallack-1986-79">[79]</a> body fossils show the earliest plants to be devoid of roots. Many had prostrate branches that sprawled along the ground, with upright axes or <a href="https://en.wiktionary.org/wiki/thalli" class="extiw" title="wikt:thalli">thalli</a> dotted here and there, and some even had non-photosynthetic subterranean branches which lacked stomata. Roots have a <a href="/wiki/Root_cap" title="Root cap">root cap</a>, unlike specialised branches.<a href="#cite_note-Raven-2001-6">[6]</a> So while Siluro-Devonian plants such as <a href="/wiki/Rhynia" title="Rhynia">Rhynia</a> and <a href="/wiki/Horneophyton" title="Horneophyton">Horneophyton</a> possessed the physiological equivalent of roots,<a href="#cite_note-80">[80]</a><a href="#cite_note-81">[81]</a> roots – defined as organs differentiated from stems – did not arrive until later.<a href="#cite_note-Raven-2001-6">[6]</a> Unfortunately, roots are rarely preserved in the fossil record.<a href="#cite_note-Raven-2001-6">[6]</a> Rhizoids – small structures performing the same role as roots, usually a cell in diameter – probably evolved very early, perhaps even before plants colonised the land; they are recognised in the <a href="/wiki/Characeae" title="Characeae">Characeae</a>, an algal sister group to land plants.<a href="#cite_note-Raven-2001-6">[6]</a> That said, rhizoids probably evolved more than once; the <a href="/wiki/Rhizine" title="Rhizine">rhizines</a> of <a href="/wiki/Lichen" title="Lichen">lichens</a>, for example, perform a similar role. Even some animals (<a href="/wiki/Lamellibrachia" title="Lamellibrachia">Lamellibrachia</a>) have root-like structures.<a href="#cite_note-Raven-2001-6">[6]</a> Rhizoids are clearly visible in the Rhynie chert fossils, and were present in most of the earliest vascular plants, and on this basis seem to have presaged true plant roots.<a href="#cite_note-82">[82]</a> More advanced structures are common in the Rhynie chert, and many other fossils of comparable early Devonian age bear structures that look like, and acted like, roots.<a href="#cite_note-Raven-2001-6">[6]</a> The rhyniophytes bore fine rhizoids, and the trimerophytes and herbaceous lycopods of the chert bore root-like structure penetrating a few centimetres into the soil.<a href="#cite_note-Algeo-1998-83">[83]</a> However, none of these fossils display all the features borne by modern roots,<a href="#cite_note-Raven-2001-6">[6]</a> with the exception of Asteroxylon, which has recently been recognized as bearing roots that evolved independently from those of extant vascular plants.<a href="#cite_note-84">[84]</a> Roots and root-like structures became increasingly common and deeper penetrating during the <a href="/wiki/Devonian" title="Devonian">Devonian</a>, with lycopod trees forming roots around 20 cm long during the Eifelian and Givetian. These were joined by progymnosperms, which rooted up to about a metre deep, during the ensuing Frasnian stage.<a href="#cite_note-Algeo-1998-83">[83]</a> True gymnosperms and zygopterid ferns also formed shallow rooting systems during the Famennian.<a href="#cite_note-Algeo-1998-83">[83]</a> The rhizophores of the lycopods provide a slightly different approach to rooting. They were equivalent to stems, with organs equivalent to leaves performing the role of rootlets.<a href="#cite_note-Raven-2001-6">[6]</a> A similar construction is observed in the extant lycopod Isoetes, and this appears to be evidence that roots evolved independently at least twice, in the lycophytes and other plants,<a href="#cite_note-Raven-2001-6">[6]</a> a proposition supported by studies showing that roots are initiated and their growth promoted by different mechanisms in lycophytes and euphyllophytes.<a href="#cite_note-Coudert-2012-85">[85]</a> Early rooted plants are little more advanced than their Silurian forebears, without a dedicated root system; however, the flat-lying axes can be clearly seen to have growths similar to the rhizoids of bryophytes today.<a href="#cite_note-Kenrick-1997b-86">[86]</a> By the Middle to Late Devonian, most groups of plants had independently developed a rooting system of some nature.<a href="#cite_note-Kenrick-1997b-86">[86]</a> As roots became larger, they could support larger trees, and the soil was weathered to a greater depth.<a href="#cite_note-Algeo-1995-78">[78]</a> This deeper weathering had effects not only on the aforementioned drawdown of CO2, but also opened up new habitats for colonisation by fungi and animals.<a href="#cite_note-Algeo-1998-83">[83]</a> The narrowest roots of modern plants are a mere 40 μm in diameter, and could not physically transport water if they were any narrower.<a href="#cite_note-Raven-2001-6">[6]</a> The earliest fossil roots recovered, by contrast, narrowed from 3 mm to under 700 μm in diameter; of course, <a href="/wiki/Taphonomy" title="Taphonomy">taphonomy</a> is the ultimate control of what thickness can be seen.<a href="#cite_note-Raven-2001-6">[6]</a> <div class="mw-heading mw-heading3"><h3 id="Tree_form">Tree form</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=15" title="Edit section: Tree form">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Psaronius_double_section.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4f/Psaronius_double_section.JPG/220px-Psaronius_double_section.JPG" decoding="async" width="220" height="222" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4f/Psaronius_double_section.JPG/330px-Psaronius_double_section.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4f/Psaronius_double_section.JPG/440px-Psaronius_double_section.JPG 2x" data-file-width="1353" data-file-height="1368" /></a><figcaption>The trunk of early tree fern <a href="/wiki/Psaronius" title="Psaronius">Psaronius</a>, showing internal structure. The top of the plant would have been to the left of the image</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:LepidodendronOhio.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/01/LepidodendronOhio.jpg/220px-LepidodendronOhio.jpg" decoding="async" width="220" height="151" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/01/LepidodendronOhio.jpg/330px-LepidodendronOhio.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/01/LepidodendronOhio.jpg/440px-LepidodendronOhio.jpg 2x" data-file-width="1821" data-file-height="1251" /></a><figcaption>External mold of <a href="/wiki/Lepidodendron" title="Lepidodendron">Lepidodendron</a> trunk showing leaf scars from the <a href="/wiki/Upper_Carboniferous" class="mw-redirect" title="Upper Carboniferous">Upper Carboniferous</a> of <a href="/wiki/Ohio" title="Ohio">Ohio</a></figcaption></figure> The early Devonian landscape was devoid of vegetation taller than waist height. Greater height provided a competitive advantage in the harvesting of sunlight for photosynthesis, overshadowing of competitors and in spore distribution, as spores (and later, seeds) could be blown for greater distances if they started higher. An effective vascular system was required in order to achieve greater heights. To attain <a href="https://en.wiktionary.org/wiki/arborescence" class="extiw" title="wikt:arborescence">arborescence</a>, plants had to develop <a href="/wiki/Wood" title="Wood">woody</a> tissue that provided both support and water transport, and thus needed to evolve the capacity for <a href="/wiki/Secondary_growth" title="Secondary growth">secondary growth</a>. The stele of plants undergoing secondary growth is surrounded by a <a href="/wiki/Vascular_cambium" title="Vascular cambium">vascular cambium</a>, a ring of meristematic cells which produces more xylem on the inside and phloem on the outside. Since xylem cells comprise dead, lignified tissue, subsequent rings of xylem are added to those already present, forming wood. Fossils of plants from the early Devonian show that a simple form of wood first appeared at least 400 million years ago, at a time when all land plants were small and herbaceous.<a href="#cite_note-Gerrienne-2011-87">[87]</a> Because wood evolved long before shrubs and trees, it is likely that its original purpose was for water transport, and that it was only used for mechanical support later.<a href="#cite_note-88">[88]</a> The first plants to develop secondary growth and a woody habit, were apparently the ferns, and as early as the Middle Devonian one species, <a href="/wiki/Wattieza" title="Wattieza">Wattieza</a>, had already reached heights of 8 m and a tree-like habit.<a href="#cite_note-Stein-2007-89">[89]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Givetianwood.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/Givetianwood.jpg/220px-Givetianwood.jpg" decoding="async" width="220" height="190" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/Givetianwood.jpg/330px-Givetianwood.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f1/Givetianwood.jpg/440px-Givetianwood.jpg 2x" data-file-width="3500" data-file-height="3024" /></a><figcaption>A piece of fossilized driftwood from the Middle Devonian (<a href="/wiki/Givetian" title="Givetian">Givetian</a>) of New York State.</figcaption></figure> Other clades did not take long to develop a tree-like stature. The Late Devonian <a href="/wiki/Archaeopteris" title="Archaeopteris">Archaeopteris</a>, a <a href="/wiki/Progymnosperm" title="Progymnosperm">precursor</a> to <a href="/wiki/Gymnosperm" title="Gymnosperm">gymnosperms</a> which evolved from the trimerophytes,<a href="#cite_note-Retallak-1985-90">[90]</a> reached 30 m in height. The progymnosperms were the first plants to develop true wood, grown from a bifacial cambium. The first appearance of one of them, <a href="/w/index.php?title=Rellimia&action=edit&redlink=1" class="new" title="Rellimia (page does not exist)">Rellimia</a>, was in the Middle Devonian.<a href="#cite_note-Dannenhoffer-1989-91">[91]</a> True wood is only thought to have evolved once, giving rise to the concept of a "lignophyte" clade.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Archaeopteris forests were soon supplemented by arborescent lycopods, in the form of <a href="/wiki/Lepidodendrales" title="Lepidodendrales">Lepidodendrales</a>, which exceeded 50m in height and 2m across at the base. These arborescent lycopods rose to dominate Late Devonian and Carboniferous forests that gave rise to <a href="/wiki/Coal" title="Coal">coal</a> deposits.<a href="#cite_note-Davis-2004-92">[92]</a> Lepidodendrales differ from modern trees in exhibiting determinate growth: after building up a reserve of nutrients at a lower height, the plants would "bolt" as a single trunk to a genetically determined height, branch at that level, spread their spores and die.<a href="#cite_note-Donoghue-2005-93">[93]</a> They consisted of "cheap" wood to allow their rapid growth, with at least half of their stems comprising a pith-filled cavity.<a href="#cite_note-Stewart-1993-1">[1]</a> Their wood was also generated by a unifacial vascular cambium – it did not produce new phloem, meaning that the trunks could not grow wider over time.[<a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification needed</a>] The <a href="/wiki/Horsetail" class="mw-redirect" title="Horsetail">horsetail</a> <a href="/wiki/Calamites" title="Calamites">Calamites</a> appeared in the <a href="/wiki/Carboniferous" title="Carboniferous">Carboniferous</a>. Unlike the modern horsetail <a href="/wiki/Equisetum" title="Equisetum">Equisetum</a>, <a href="/wiki/Calamites" title="Calamites">Calamites</a> had a unifacial vascular cambium, allowing them to develop wood and grow to heights in excess of 10 m and to branch repeatedly. While the form of early trees was similar to that of today's, the <a href="/wiki/Spermatophytes" class="mw-redirect" title="Spermatophytes">Spermatophytes</a> or seed plants, the group that contain all modern trees, had yet to evolve. The dominant tree groups today are all seed plants, the gymnosperms, which include the coniferous trees, and the angiosperms, which contain all fruiting and flowering trees. No free-sporing trees like Archaeopteris exist in the extant flora. It was long thought that the angiosperms arose from within the gymnosperms, but recent molecular evidence suggests that their living representatives form two distinct groups.<a href="#cite_note-Bowe-2000-94">[94]</a><a href="#cite_note-Chaw-2000-95">[95]</a><a href="#cite_note-Soltis-2002-96">[96]</a> The molecular data has yet to be fully reconciled with morphological data,<a href="#cite_note-Friis-2006-97">[97]</a><a href="#cite_note-Hilton-2006-98">[98]</a><a href="#cite_note-Bateman-2006-99">[99]</a> but it is becoming accepted that the morphological support for paraphyly is not especially strong.<a href="#cite_note-Frohlich-2007-100">[100]</a> This would lead to the conclusion that both groups arose from within the pteridosperms, probably as early as the <a href="/wiki/Permian" title="Permian">Permian</a>.<a href="#cite_note-Frohlich-2007-100">[100]</a> The angiosperms and their ancestors played a very small role until they diversified during the Cretaceous. They started out as small, damp-loving organisms in the understorey, and have been diversifying ever since the Cretaceous,<a href="#cite_note-101">[101]</a> to become the dominant member of non-<a href="https://en.wiktionary.org/wiki/boreal" class="extiw" title="wikt:boreal">boreal</a> forests today. <div class="mw-heading mw-heading3"><h3 id="Seeds">Seeds</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=16" title="Edit section: Seeds">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Trigonocarpus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/77/Trigonocarpus.jpg/220px-Trigonocarpus.jpg" decoding="async" width="220" height="171" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/77/Trigonocarpus.jpg/330px-Trigonocarpus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/77/Trigonocarpus.jpg/440px-Trigonocarpus.jpg 2x" data-file-width="552" data-file-height="429" /></a><figcaption>The fossil seed <a href="/w/index.php?title=Trigonocarpus&action=edit&redlink=1" class="new" title="Trigonocarpus (page does not exist)">Trigonocarpus</a></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Runcaria_megasporangium_and_cupule_drawing.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/04/Runcaria_megasporangium_and_cupule_drawing.jpg/90px-Runcaria_megasporangium_and_cupule_drawing.jpg" decoding="async" width="90" height="248" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/04/Runcaria_megasporangium_and_cupule_drawing.jpg/135px-Runcaria_megasporangium_and_cupule_drawing.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/04/Runcaria_megasporangium_and_cupule_drawing.jpg/180px-Runcaria_megasporangium_and_cupule_drawing.jpg 2x" data-file-width="458" data-file-height="1262" /></a><figcaption>The transitional fossil <a href="/wiki/Runcaria" title="Runcaria">Runcaria</a></figcaption></figure> Early land plants reproduced in the fashion of ferns: spores germinated into small gametophytes, which produced eggs and/or sperm. These sperm would swim across moist soils to find the female organs (archegonia) on the same or another gametophyte, where they would fuse with an egg to produce an embryo, which would germinate into a sporophyte.<a href="#cite_note-Algeo-1998-83">[83]</a> Heterosporic plants, as their name suggests, bear spores of two sizes – microspores and megaspores. These would germinate to form microgametophytes and megagametophytes, respectively. This system paved the way for ovules and seeds: taken to the extreme, the megasporangia could bear only a single megaspore tetrad, and to complete the transition to true ovules, three of the megaspores in the original tetrad could be aborted, leaving one megaspore per megasporangium.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The transition to ovules continued with this megaspore being "boxed in" to its sporangium while it germinated. Then, the megagametophyte was contained within a waterproof integument, which enclosed the seed. The pollen grain, which contained a microgametophyte germinated from a microspore , was employed for dispersal of the male gamete, only releasing its desiccation-prone flagellate sperm when it reached a receptive megagametophyte.<a href="#cite_note-Stewart-1993-1">[1]</a> Lycopods and sphenopsids got a fair way down the path to the seed habit without ever crossing the threshold. Fossil lycopod megaspores reaching 1 cm in diameter, and surrounded by vegetative tissue, are known (Lepidocarpon, Achlamydocarpon);– these even germinated into a megagametophyte in situ. However, they fell short of being ovules, since the nucellus, an inner spore-covering layer, does not completely enclose the spore. A very small slit (micropyle) remains, meaning that the megasporangium is still exposed to the atmosphere. This has two consequences – firstly, it means it is not fully resistant to desiccation, and secondly, sperm do not have to "burrow" to access the archegonia of the megaspore.<a href="#cite_note-Stewart-1993-1">[1]</a> A Middle <a href="/wiki/Devonian" title="Devonian">Devonian</a> <a href="/wiki/Transitional_fossil" title="Transitional fossil">precursor</a> to seed plants from Belgium has been identified predating the earliest <a href="/wiki/Seed_plants" class="mw-redirect" title="Seed plants">seed plants</a> by about 20 million years. <a href="/wiki/Runcaria" title="Runcaria">Runcaria</a>, small and radially symmetrical, is an integumented <a href="/wiki/Megasporangium" class="mw-redirect" title="Megasporangium">megasporangium</a> surrounded by a cupule. The <a href="/wiki/Megasporangium" class="mw-redirect" title="Megasporangium">megasporangium</a> bears an unopened distal extension protruding above the multilobed <a href="/wiki/Integument" title="Integument">integument</a>. It is suspected that the extension was involved in anemophilous <a href="/wiki/Pollination" title="Pollination">pollination</a>. Runcaria sheds new light on the sequence of character acquisition leading to the seed. Runcaria has all of the qualities of seed plants except for a solid <a href="/wiki/Seed_coat" class="mw-redirect" title="Seed coat">seed coat</a> and a system to guide the pollen to the ovule.<a href="#cite_note-102">[102]</a> The first spermatophytes (literally: "seed plants") – that is, the first plants to bear true seeds – are called <a href="/wiki/Pteridosperm" class="mw-redirect" title="Pteridosperm">pteridosperms</a>: literally, "seed ferns", so called because their foliage consisted of fern-like fronds, although they were not closely related to ferns. The oldest fossil evidence of seed plants is of Late Devonian age, and they appear to have evolved out of an earlier group known as the <a href="/wiki/Progymnosperms" class="mw-redirect" title="Progymnosperms">progymnosperms</a>. These early seed plants ranged from trees to small, rambling shrubs; like most early progymnosperms, they were woody plants with fern-like foliage. They all bore ovules, but no cones, fruit or similar. While it is difficult to track the early evolution of seeds, the lineage of the seed ferns may be traced from the simple trimerophytes through homosporous <a href="/w/index.php?title=Aneurophytes&action=edit&redlink=1" class="new" title="Aneurophytes (page does not exist)">Aneurophytes</a>.<a href="#cite_note-Stewart-1993-1">[1]</a> The seed plants underwent their first major evolutionary radiation in the <a href="/wiki/Famennian" title="Famennian">Famennian</a> era.<a href="#cite_note-103">[103]</a> This seed model is shared by basically all gymnosperms (literally: "naked seeds"), most of which encase their seeds in a woody cone or fleshy aril (the <a href="/wiki/Taxus" title="Taxus">yew</a>, for example), but none of which fully enclose their seeds. The angiosperms ("vessel seeds") are the only group to fully enclose the seed, in a carpel.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Fully enclosed seeds opened up a new pathway for plants to follow: that of <a href="/wiki/Seed_dormancy" title="Seed dormancy">seed dormancy</a>. The embryo, completely isolated from the external atmosphere and hence protected from desiccation, could survive some years of drought before germinating. Gymnosperm seeds from the Late Carboniferous have been found to contain embryos, suggesting a lengthy gap between fertilisation and germination.<a href="#cite_note-Mapes-1989-104">[104]</a> This period is associated with the entry into a <a href="/wiki/Greenhouse_earth" class="mw-redirect" title="Greenhouse earth">greenhouse earth</a> period, with an associated increase in aridity. This suggests that dormancy arose as a response to drier climatic conditions, where it became advantageous to wait for a moist period before germinating.<a href="#cite_note-Mapes-1989-104">[104]</a> This evolutionary breakthrough appears to have opened a floodgate: previously inhospitable areas, such as dry mountain slopes, could now be tolerated, and were soon covered by trees.<a href="#cite_note-Mapes-1989-104">[104]</a> Seeds offered further advantages to their bearers: they increased the success rate of fertilised gametophytes, and because a nutrient store could be "packaged" in with the embryo, the seeds could germinate rapidly in inhospitable environments, reaching a size where it could fend for itself more quickly.<a href="#cite_note-Algeo-1998-83">[83]</a> For example, without an endosperm, seedlings growing in arid environments would not have the reserves to grow roots deep enough to reach the water table before they expired from dehydration.<a href="#cite_note-Algeo-1998-83">[83]</a> Likewise, seeds germinating in a gloomy understory require an additional reserve of energy to quickly grow high enough to capture sufficient light for self-sustenance.<a href="#cite_note-Algeo-1998-83">[83]</a> A combination of these advantages gave seed plants the ecological edge over the previously dominant genus Archaeopteris, thus increasing the biodiversity of early forests.<a href="#cite_note-Algeo-1998-83">[83]</a> Despite these advantages, it is common for fertilized ovules to fail to mature as seeds.<a href="#cite_note-105">[105]</a> Also during seed dormancy (often associated with unpredictable and stressful conditions) DNA damage accumulates.<a href="#cite_note-106">[106]</a><a href="#cite_note-107">[107]</a><a href="#cite_note-108">[108]</a> Thus DNA damage appears to be a basic problem for survival of seed plants, just as DNA damage is a <a href="/wiki/DNA_damage_(naturally_occurring)#A_major_problem_for_life" title="DNA damage (naturally occurring)">major problem for life</a> in general.<a href="#cite_note-109">[109]</a> <div class="mw-heading mw-heading3"><h3 id="Flowers">Flowers</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=17" title="Edit section: Flowers">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">For a more ecological discussion on the evolution of flowers and earliest flower, see <a href="/wiki/Flower" title="Flower">Flower</a>.</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Crossotheca_nodule.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Crossotheca_nodule.JPG/220px-Crossotheca_nodule.JPG" decoding="async" width="220" height="139" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Crossotheca_nodule.JPG/330px-Crossotheca_nodule.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Crossotheca_nodule.JPG/440px-Crossotheca_nodule.JPG 2x" data-file-width="1000" data-file-height="634" /></a><figcaption>The pollen bearing organs of the early "flower" <a href="/wiki/Crossotheca" title="Crossotheca">Crossotheca</a></figcaption></figure> Flowers are modified leaves possessed only by the <a href="/wiki/Angiosperm" class="mw-redirect" title="Angiosperm">angiosperms</a>, which are relatively late to appear in the fossil record. The group originated and diversified during the Early Cretaceous and became ecologically significant thereafter.<a href="#cite_note-Feild-2011-110">[110]</a> Flower-like structures first appear in the <a href="/wiki/Fossil" title="Fossil">fossil</a> records some ~130 mya, in the <a href="/wiki/Cretaceous" title="Cretaceous">Cretaceous</a>.<a href="#cite_note-Lawton-Rauh-2000-111">[111]</a> However, in 2018, scientists reported the finding of a fossil <a href="/wiki/Flower" title="Flower">flower</a> from about 180 million years ago, 50 million years earlier than previously thought.<a href="#cite_note-Chinese_Academy_of_Sciences-2018-112">[112]</a> The interpretation has been however highly disputed.<a href="#cite_note-113">[113]</a> Colorful and/or pungent structures surround the cones of plants such as <a href="/wiki/Cycads" class="mw-redirect" title="Cycads">cycads</a> and <a href="/wiki/Gnetales" class="mw-redirect" title="Gnetales">Gnetales</a>, making a strict definition of the term "flower" elusive.<a href="#cite_note-Bateman-2006-99">[99]</a> The main function of a flower is <a href="/wiki/Reproduction" title="Reproduction">reproduction</a>, which, before the evolution of the flower and <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">angiosperms</a>, was the job of microsporophylls and megasporophylls. A flower can be considered a powerful evolutionary <a href="/wiki/Innovation" title="Innovation">innovation</a>, because its presence allowed the plant world to access new means and mechanisms for reproduction.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Syncarp_evolution.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Syncarp_evolution.svg/220px-Syncarp_evolution.svg.png" decoding="async" width="220" height="330" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Syncarp_evolution.svg/330px-Syncarp_evolution.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Syncarp_evolution.svg/440px-Syncarp_evolution.svg.png 2x" data-file-width="170" data-file-height="255" /></a><figcaption>The evolution of syncarps. a: sporangia borne at tips of leaf b: Leaf curls up to protect sporangia c: leaf curls to form enclosed roll d: grouping of three rolls into a syncarp</figcaption></figure> The flowering plants have long been assumed to have evolved from within the <a href="/wiki/Gymnosperms" class="mw-redirect" title="Gymnosperms">gymnosperms</a>; according to the traditional morphological view, they are closely allied to the <a href="/wiki/Gnetales" class="mw-redirect" title="Gnetales">Gnetales</a>. However, as noted above, recent molecular evidence is at odds with this hypothesis,<a href="#cite_note-Chaw-2000-95">[95]</a><a href="#cite_note-Soltis-2002-96">[96]</a> and further suggests that Gnetales are more closely related to some gymnosperm groups than angiosperms,<a href="#cite_note-Bowe-2000-94">[94]</a> and that <a href="https://en.wiktionary.org/wiki/extant" class="extiw" title="wikt:extant">extant</a> gymnosperms form a distinct clade to the angiosperms,<a href="#cite_note-Bowe-2000-94">[94]</a><a href="#cite_note-Chaw-2000-95">[95]</a><a href="#cite_note-Soltis-2002-96">[96]</a> the two clades diverging some <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=300">300</a> million years ago.<a href="#cite_note-Nei-2003-114">[114]</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Gnetophyta#Classification" title="Gnetophyta">Gnetophyta § Classification</a></div> The relationship of <a href="/wiki/Stem_group" class="mw-redirect" title="Stem group">stem groups</a> to the <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">angiosperms</a> is important in determining the evolution of flowers. Stem groups provide an insight into the state of earlier "forks" on the path to the current state. Convergence increases the risk of misidentifying stem groups. Since the protection of the <a href="/wiki/Megagametophyte" class="mw-redirect" title="Megagametophyte">megagametophyte</a> is evolutionarily desirable, probably many separate groups evolved protective encasements independently. In flowers, this protection takes the form of a <a href="/wiki/Carpel" class="mw-redirect" title="Carpel">carpel</a>, evolved from a leaf and recruited into a protective role, shielding the ovules. These ovules are further protected by a double-walled <a href="https://en.wiktionary.org/wiki/integument" class="extiw" title="wikt:integument">integument</a>. Penetration of these protective layers needs something more than a free-floating <a href="/wiki/Microgametophyte" class="mw-redirect" title="Microgametophyte">microgametophyte</a>. <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">Angiosperms</a> have pollen grains comprising just three cells. One cell is responsible for drilling down through the integuments, and creating a conduit for the two sperm cells to flow down. The megagametophyte has just seven cells; of these, one fuses with a sperm cell, forming the nucleus of the egg itself, and another joins with the other sperm, and dedicates itself to forming a nutrient-rich <a href="/wiki/Endosperm" title="Endosperm">endosperm</a>. The other cells take auxiliary roles.[<a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify">clarification needed</a>] This process of "<a href="/wiki/Double_fertilisation" class="mw-redirect" title="Double fertilisation">double fertilisation</a>" is unique and common to all angiosperms. <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Bennettitales-cycadeoidaceae.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Bennettitales-cycadeoidaceae.jpg/220px-Bennettitales-cycadeoidaceae.jpg" decoding="async" width="220" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Bennettitales-cycadeoidaceae.jpg/330px-Bennettitales-cycadeoidaceae.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/46/Bennettitales-cycadeoidaceae.jpg/440px-Bennettitales-cycadeoidaceae.jpg 2x" data-file-width="1473" data-file-height="1422" /></a><figcaption>The inflorescences of the <a href="/wiki/Bennettitales" title="Bennettitales">Bennettitales</a> are strikingly similar to flowers, but evolved independently.</figcaption></figure> In the fossil record, there are three intriguing groups which bore flower-like structures. The first is the <a href="/wiki/Permian" title="Permian">Permian</a> pteridosperm <a href="/wiki/Glossopteris" title="Glossopteris">Glossopteris</a>, which already bore recurved leaves resembling carpels. The <a href="/wiki/Mesozoic" title="Mesozoic">Mesozoic</a> <a href="/wiki/Caytonia" title="Caytonia">Caytonia</a> is more flower-like still, with enclosed ovules – but only a single integument. Further, details of their pollen and stamens set them apart from true flowering plants. The <a href="/wiki/Bennettitales" title="Bennettitales">Bennettitales</a> bore remarkably flower-like organs, protected by whorls of <a href="https://en.wiktionary.org/wiki/bract" class="extiw" title="wikt:bract">bracts</a> which may have played a similar role to the petals and sepals of true flowers; however, these flower-like structures evolved independently, as the Bennettitales are more closely related to <a href="/wiki/Cycad" title="Cycad">cycads</a> and <a href="/wiki/Ginkgo" title="Ginkgo">ginkgos</a> than to the angiosperms.<a href="#cite_note-Crepet-2000-115">[115]</a> However, no true flowers are found in any groups save those extant today. Most morphological and molecular analyses place <a href="/wiki/Amborella" title="Amborella">Amborella</a>, the <a href="/wiki/Nymphaeales" title="Nymphaeales">nymphaeales</a> and <a href="/wiki/Austrobaileya" title="Austrobaileya">Austrobaileyaceae</a> in a basal clade called "ANA". This clade appear to have diverged in the early Cretaceous, around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=130">130</a> million years ago – around the same time as the <a href="/wiki/Archaefructus" title="Archaefructus">earliest fossil angiosperm</a>,<a href="#cite_note-Sun-2002-116">[116]</a><a href="#cite_note-Doyle-2003-117">[117]</a> and just after the <a href="/w/index.php?title=Clavatipollenites&action=edit&redlink=1" class="new" title="Clavatipollenites (page does not exist)">first angiosperm-like pollen</a>, 136 million years ago.<a href="#cite_note-Frohlich-2007-100">[100]</a> The <a href="/wiki/Magnoliids" title="Magnoliids">magnoliids</a> diverged soon after, and a rapid radiation had produced eudicots and monocots by <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=125">125</a> million years ago.<a href="#cite_note-Frohlich-2007-100">[100]</a> By the end of the Cretaceous <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=66">66</a> million years ago, over 50% of today's angiosperm orders had evolved, and the clade accounted for 70% of global species.<a href="#cite_note-Wing-1998-118">[118]</a> It was around this time that flowering trees became dominant over <a href="/wiki/Conifer" title="Conifer">conifers</a>.<a href="#cite_note-Stewart-1993-1">[1]</a>: 498  The features of the basal "ANA" groups suggest that angiosperms originated in dark, damp, frequently disturbed areas.<a href="#cite_note-Feild-2004-119">[119]</a> It appears that the angiosperms remained constrained to such habitats throughout the Cretaceous – occupying the niche of small herbs early in the successional series.<a href="#cite_note-Wing-1998-118">[118]</a> This may have restricted their initial significance, but given them the flexibility that accounted for the rapidity of their later diversifications in other habitats.<a href="#cite_note-Feild-2004-119">[119]</a> <style data-mw-deduplicate="TemplateStyles:r1216315873">.mw-parser-output div.clade-gallery{display:table;margin:0 auto}.mw-parser-output div.clade-gallery>div{line-height:1.5em;padding:0.2em 0.4em;border:1px solid #a2a9b1}.mw-parser-output div.clade-gallery div.main-caption{text-align:center;font-weight:bold;border:none}.mw-parser-output div.clade-gallery div.main-footer{text-align:center}.mw-parser-output div.clade-gallery div.main-content{display:table}.mw-parser-output div.clade-gallery table.gallery-element{float:left!important;border:1px none;border:1px solid #a2a9b1;padding:5px;margin:1px;border-collapse:collapse}.mw-parser-output div.clade-gallery div.gallery-element{float:left;border:1px none;border:1px solid #a2a9b1;padding:5px;margin:1px;border-collapse:collapse}.mw-parser-output div.clade-gallery .no-border{border:none}.mw-parser-output div.clade-gallery table.gallery-element.vertical{float:none}.mw-parser-output div.clade-gallery table.gallery-element caption.caption{text-align:center;font-weight:bold;border:none}.mw-parser-output div.clade-gallery table.gallery-element th.header{text-align:center;font-weight:bold}.mw-parser-output div.clade-gallery table.gallery-element td.cladogram{padding:10px;margin:-1px 0}.mw-parser-output div.clade-gallery table.gallery-element td.footer{border:1px solid #a2a9b1;margin:0}.mw-parser-output div.clade-gallery div.header.shading{background-color:#eaecf0}.mw-parser-output div.clade-gallery div.cladogram.shading{background-color:#f8f9fa}.mw-parser-output div.clade-gallery div.footer.shading{background-color:#f8f9fa}</style><div class="clade-gallery"><div class="main-content no-border"><table class="gallery-element" style=""><tbody><tr class="header"><th>Traditional view</th></tr><tr><td class="cladogram"> <div class="clade"><style data-mw-deduplicate="TemplateStyles:r1258728058">body.skin-vector-2022 .mw-parser-output div.clade,body.skin-minerva .mw-parser-output div.clade{overflow-x:auto;overflow-y:hidden}body.skin-minerva .mw-parser-output div.clade p{font-size:inherit}.mw-parser-output table.clade{border-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto}.mw-parser-output table.clade table.clade{width:100%;line-height:inherit}.mw-parser-output table.clade td.clade-label{min-width:0.2em;width:0.2em;padding:0.1em 0.25em;vertical-align:bottom;text-align:center;border-left:1px solid;border-bottom:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-label::before,.mw-parser-output table.clade td.clade-slabel::before{content:"\2060 "}.mw-parser-output table.clade td.clade-fixed-width{overflow:hidden;text-overflow:ellipsis}.mw-parser-output table.clade td.clade-fixed-width:hover{overflow:visible}.mw-parser-output table.clade td.clade-label.first{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-label.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-slabel{padding:0.1em 0.25em;vertical-align:top;text-align:center;border-left:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-slabel:hover{overflow:visible}.mw-parser-output table.clade td.clade-slabel.last{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-slabel.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-bar{vertical-align:middle;text-align:left;padding:0 0.5em;position:relative}.mw-parser-output table.clade td.clade-bar.reverse{text-align:right;position:relative}.mw-parser-output table.clade td.clade-leaf{border:0;padding:0;text-align:left}.mw-parser-output table.clade td.clade-leafR{border:0;padding:0;text-align:right}.mw-parser-output table.clade td.clade-leaf.reverse{text-align:right}.mw-parser-output table.clade:hover span.linkA{background-color:yellow}.mw-parser-output table.clade:hover span.linkB{background-color:green}</style> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Cycads </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Ginkgo </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Conifers </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">Anthophytes </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Bennettitales </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> Gnetales </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> Angiosperms </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div></td></tr></tbody></table><table class="gallery-element" style=""><tbody><tr class="header"><th>Modern view</th></tr><tr><td class="cladogram"> <div class="clade"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Angiosperms </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">Gymnosperms </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Cycads </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> Bennettitales </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> Ginkgo </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> Conifers </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> Gnetales </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div></td></tr></tbody></table></div><div class="main-footer">Phylogeny of anthophytes and gymnosperms based on Crepet et al. (2000)<a href="#cite_note-Crepet2000-120">[120]</a></div></div> Some propose that the Angiosperms arose from an unknown Seed Fern, Pteridophyte, and view Cycads as living Seed Ferns with both Seed-Bearing and sterile leaves (Cycas revoluta)<a href="#cite_note-Hilton-2006-98">[98]</a> In August 2017, scientists presented a detailed description and 3D reconstruction of possibly the first <a href="/wiki/Flower" title="Flower">flower</a> that lived about 140 million years ago.<a href="#cite_note-Gabbott-2017-121">[121]</a><a href="#cite_note-Sauquet-2017-122">[122]</a> <div class="mw-heading mw-heading4"><h4 id="Origins_of_the_flower">Origins of the flower</h4>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=18" title="Edit section: Origins of the flower">edit</a>]</div> The family <a href="/wiki/Amborellaceae" class="mw-redirect" title="Amborellaceae">Amborellaceae</a> is regarded as being the sister <a href="/wiki/Cladistics" title="Cladistics">clade</a> to all other living flowering plants. A draft genome of Amborella trichopoda was published in December, 2013.<a href="#cite_note-123">[123]</a> By comparing its genome with those of all other living flowering plants, it will be possible to work out the most likely characteristics of the ancestor of A. trichopoda and all other flowering plants, i.e. the ancestral flowering plant.<a href="#cite_note-124">[124]</a> It seems that on the level of the organ, the <a href="/wiki/Leaf" title="Leaf">leaf</a> may be the ancestor of the flower, or at least some floral organs. When some crucial genes involved in flower development are <a href="/wiki/Mutate" class="mw-redirect" title="Mutate">mutated</a>, clusters of leaf-like structures arise in place of flowers. Thus, sometime in history, the developmental program leading to formation of a leaf must have been altered to generate a flower. There probably also exists an overall robust framework within which the floral diversity has been generated. An example of that is a gene called <a href="/wiki/LEAFY" class="mw-redirect" title="LEAFY">LEAFY</a> (LFY), which is involved in flower development in <a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a>. The <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologs</a> of this gene are found in <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">angiosperms</a> as diverse as <a href="/wiki/Tomato" title="Tomato">tomato</a>, <a href="/wiki/Antirrhinum" title="Antirrhinum">snapdragon</a>, <a href="/wiki/Pea" title="Pea">pea</a>, <a href="/wiki/Maize" title="Maize">maize</a> and even <a href="/wiki/Gymnosperms" class="mw-redirect" title="Gymnosperms">gymnosperms</a>. Expression of Arabidopsis thaliana LFY in distant plants like <a href="/wiki/Populus" title="Populus">poplar</a> and <a href="/wiki/Citrus" title="Citrus">citrus</a> also results in flower-production in these plants. The LFY gene regulates the expression of some genes belonging to the <a href="/wiki/MADS-box" title="MADS-box">MADS-box</a> family. These genes, in turn, act as direct controllers of flower development.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Adaptive_function_of_flowers">Adaptive function of flowers</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=19" title="Edit section: Adaptive function of flowers">edit</a>]</div> <a href="/wiki/Flower" title="Flower">Flowers</a> likely emerged during plant evolution as an adaptation to facilitate cross-<a href="/wiki/Fertilization" class="mw-redirect" title="Fertilization">fertilization</a> (<a href="/wiki/Outcrossing" title="Outcrossing">outcrossing</a>), a process that leads to the masking of recessive deleterious <a href="/wiki/Mutation" title="Mutation">mutations</a> in progeny <a href="/wiki/Genome" title="Genome">genomes</a>. This masking effect of expression of deleterious mutations is referred to as <a href="/wiki/Complementation_(genetics)" title="Complementation (genetics)">genetic complementation</a>.<a href="#cite_note-Bernstein-1985-125">[125]</a> This beneficial masking effect of cross-fertilization is also considered to be the basis of <a href="/wiki/Hybrid_vigour" class="mw-redirect" title="Hybrid vigour">hybrid vigor</a> or <a href="/wiki/Heterosis" title="Heterosis">heterosis</a> in progeny. Once flowers have become established in a lineage based on their adaptive function of promoting cross-fertilization, subsequent switching to inbreeding ordinarily then becomes disadvantageous, mainly because it permits expression of the previously masked deleterious recessive mutations, i.e. <a href="/wiki/Inbreeding_depression" title="Inbreeding depression">inbreeding depression</a>. In addition, <a href="/wiki/Meiosis" title="Meiosis">meiosis</a>, the process by which seed progeny are produced in flowering plants, provides a direct mechanism for <a href="/wiki/DNA_repair" title="DNA repair">repairing DNA</a> through genetic recombination.<a href="#cite_note-Hörandl-2024-126">[126]</a> Thus, in flowering plants, the two fundamental processes of sexual reproduction are cross-fertilization (outcrossing) and meiosis and these two processes appear to be maintained respectively by the advantages of genetic complementation and recombinational repair of DNA.<a href="#cite_note-Bernstein-1985-125">[125]</a> <div class="mw-heading mw-heading4"><h4 id="Evolution_of_the_MADS-box_family">Evolution of the MADS-box family</h4>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=20" title="Edit section: Evolution of the MADS-box family">edit</a>]</div> The members of the <a href="/wiki/MADS-box" title="MADS-box">MADS-box</a> family of transcription factors play a very important and evolutionarily conserved role in flower development. According to the <a href="/wiki/The_ABC_Model_of_Flower_Development" class="mw-redirect" title="The ABC Model of Flower Development">ABC Model of flower development</a>, three zones — A, B and C — are generated within the developing flower primordium, by the action of some <a href="/wiki/Transcription_factors" class="mw-redirect" title="Transcription factors">transcription factors</a>, that are members of the <a href="/wiki/MADS-box" title="MADS-box">MADS-box</a> family. Among these, the functions of the B and C domain genes have been evolutionarily more conserved than the A domain gene. Many of these genes have arisen through <a href="/wiki/Gene_duplication" title="Gene duplication">gene duplications</a> of ancestral members of this family. Quite a few of them show redundant functions.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The evolution of the <a href="/wiki/MADS-box" title="MADS-box">MADS-box</a> family has been extensively studied. These genes are present even in <a href="/wiki/Pteridophytes" class="mw-redirect" title="Pteridophytes">pteridophytes</a>, but the spread and diversity is many times higher in <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">angiosperms</a>.<a href="#cite_note-127">[127]</a> There appears to be quite a bit of pattern into how this family has evolved. Consider the evolution of the C-region gene <a href="/wiki/AGAMOUS" class="mw-redirect" title="AGAMOUS">AGAMOUS</a> (AG). It is expressed in today's flowers in the <a href="/wiki/Stamens" class="mw-redirect" title="Stamens">stamens</a>, and the <a href="/wiki/Carpel" class="mw-redirect" title="Carpel">carpel</a>, which are reproductive organs. Its ancestor in <a href="/wiki/Gymnosperms" class="mw-redirect" title="Gymnosperms">gymnosperms</a> also has the same expression pattern. Here, it is expressed in the <a href="/wiki/Strobilus" title="Strobilus">strobili</a>, an organ that produces <a href="/wiki/Pollen" title="Pollen">pollen</a> or ovules.<a href="#cite_note-128">[128]</a> Similarly, the B-genes' (AP3 and PI) ancestors are expressed only in the male organs in <a href="/wiki/Gymnosperms" class="mw-redirect" title="Gymnosperms">gymnosperms</a>. Their descendants in the modern angiosperms also are expressed only in the <a href="/wiki/Stamens" class="mw-redirect" title="Stamens">stamens</a>, the male reproductive organ. Thus, the same, then-existing components were used by the plants in a novel manner to generate the first flower. This is a recurring pattern in <a href="/wiki/Evolution" title="Evolution">evolution</a>.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Factors_influencing_floral_diversity">Factors influencing floral diversity</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=21" title="Edit section: Factors influencing floral diversity">edit</a>]</div> There is enormous variation in floral structure in plants, typically due to changes in the <a href="/wiki/MADS-box" title="MADS-box">MADS-box</a> genes and their expression pattern. For example, grasses possess unique floral structures. The carpels and stamens are surrounded by scale-like <a href="/wiki/Monocot" class="mw-redirect" title="Monocot">lodicules</a> and two bracts, the lemma and the palea, but genetic evidence and morphology suggest that lodicules are homologous to <a href="/wiki/Eudicot" class="mw-redirect" title="Eudicot">eudicot</a> petals.<a href="#cite_note-Thompson-2009-129">[129]</a> The palea and lemma may be homologous to sepals in other groups, or may be unique grass structures.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Another example is that of <a href="/wiki/Linaria_vulgaris" title="Linaria vulgaris">Linaria vulgaris</a>, which has two kinds of flower symmetries-<a href="/wiki/Symmetry_in_biology#Radial_symmetry" title="Symmetry in biology">radial</a> and <a href="/wiki/Symmetry_in_biology#Bilateral_symmetry" title="Symmetry in biology">bilateral</a>. These symmetries are due to <a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">epigenetic</a> changes in just one gene called CYCLOIDEA.<a href="#cite_note-Lawton-Rauh-2000-111">[111]</a> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Pink_rose_albury_botanical_gardens.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Pink_rose_albury_botanical_gardens.jpg/200px-Pink_rose_albury_botanical_gardens.jpg" decoding="async" width="200" height="133" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Pink_rose_albury_botanical_gardens.jpg/300px-Pink_rose_albury_botanical_gardens.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Pink_rose_albury_botanical_gardens.jpg/400px-Pink_rose_albury_botanical_gardens.jpg 2x" data-file-width="1600" data-file-height="1067" /></a><figcaption>Large number of <a href="/wiki/Petals" class="mw-redirect" title="Petals">petals</a> in <a href="/wiki/Rose" title="Rose">roses</a> is the result of human selection</figcaption></figure> Arabidopsis thaliana has a gene called <a href="/wiki/AGAMOUS" class="mw-redirect" title="AGAMOUS">AGAMOUS</a> that plays an important role in defining how many <a href="/wiki/Petals" class="mw-redirect" title="Petals">petals</a> and <a href="/wiki/Sepals" class="mw-redirect" title="Sepals">sepals</a> and other organs are generated. Mutations in this gene give rise to the floral <a href="/wiki/Meristem" title="Meristem">meristem</a> obtaining an indeterminate fate, and proliferation of floral organs in <a href="/wiki/Double-flowered" title="Double-flowered">double-flowered</a> forms of <a href="/wiki/Rose" title="Rose">roses</a>, <a href="/wiki/Carnations" class="mw-redirect" title="Carnations">carnations</a> and <a href="/wiki/Morning_glory" title="Morning glory">morning glory</a>. These <a href="/wiki/Phenotypes" class="mw-redirect" title="Phenotypes">phenotypes</a> have been selected by horticulturists for their increased number of <a href="/wiki/Petals" class="mw-redirect" title="Petals">petals</a>.<a href="#cite_note-130">[130]</a> Several studies on diverse plants like <a href="/wiki/Petunia" title="Petunia">petunia</a>, <a href="/wiki/Tomato" title="Tomato">tomato</a>, <a href="/wiki/Impatiens" title="Impatiens">Impatiens</a>, <a href="/wiki/Maize" title="Maize">maize</a>, etc. have suggested that the enormous diversity of flowers is a result of small changes in <a href="/wiki/Genes" class="mw-redirect" title="Genes">genes</a> controlling their development.<a href="#cite_note-131">[131]</a> The <a href="/wiki/Floral_Genome_Project" title="Floral Genome Project">Floral Genome Project</a> confirmed that the <a href="/wiki/ABC_Model_of_flower_development" class="mw-redirect" title="ABC Model of flower development">ABC Model of flower development</a> is not conserved across all <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">angiosperms</a>. Sometimes expression domains change, as in the case of many <a href="/wiki/Monocots" class="mw-redirect" title="Monocots">monocots</a>, and also in some basal angiosperms like <a href="/wiki/Amborella" title="Amborella">Amborella</a>. Different models of flower development like the Fading boundaries model, or the Overlapping-boundaries model which propose non-rigid domains of expression, may explain these architectures.<a href="#cite_note-132">[132]</a> There is a possibility that from the basal to the modern angiosperms, the domains of floral architecture have become more and more fixed through evolution.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading4"><h4 id="Flowering_time">Flowering time</h4>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=22" title="Edit section: Flowering time">edit</a>]</div> Another floral feature that has been a subject of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> is flowering time. Some plants flower early in their life cycle, others require a period of <a href="/wiki/Vernalization" title="Vernalization">vernalization</a> before flowering. This outcome is based on factors like <a href="/wiki/Temperature" title="Temperature">temperature</a>, <a href="/wiki/Intensity_(physics)" title="Intensity (physics)">light intensity</a>, presence of <a href="/wiki/Pollinators" class="mw-redirect" title="Pollinators">pollinators</a> and other environmental signals: genes like CONSTANS (CO), <a href="/wiki/Flowering_Locus_C" title="Flowering Locus C">Flowering Locus C</a> (FLC) and FRIGIDA regulate integration of environmental signals into the pathway for flower development. Variations in these loci have been associated with flowering time variations between plants. For example, <a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a> ecotypes that grow in the cold, <a href="/wiki/Temperate" class="mw-redirect" title="Temperate">temperate</a> regions require prolonged vernalization before they flower, while the <a href="/wiki/Tropical" class="mw-redirect" title="Tropical">tropical</a> varieties, and the most common lab strains, don't. This variation is due to mutations in the FLC and FRIGIDA genes, rendering them non-functional.<a href="#cite_note-133">[133]</a> Many of the genes involved in this process are conserved across all the plants studied. Sometimes though, despite genetic conservation, the mechanism of action turns out to be different. For example, <a href="/wiki/Rice" title="Rice">rice</a> is a short-day plant, while <a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a> is a long-day plant. Both plants have the proteins CO and FLOWERING LOCUS T (FT), but, in Arabidopsis thaliana, CO enhances FT production, while in rice, the CO homolog represses FT production, resulting in completely opposite downstream effects.<a href="#cite_note-134">[134]</a> <div class="mw-heading mw-heading4"><h4 id="Theories_of_flower_evolution">Theories of flower evolution</h4>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=23" title="Edit section: Theories of flower evolution">edit</a>]</div> The <a href="/wiki/Anthophyta" title="Anthophyta">Anthophyte theory</a> was based on the observation that a gymnospermic group <a href="/wiki/Gnetae" class="mw-redirect" title="Gnetae">Gnetales</a> has a flower-like <a href="/wiki/Ovule" title="Ovule">ovule</a>. It has partially developed <a href="/wiki/Vessel_element" title="Vessel element">vessels</a> as found in the <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">angiosperms</a>, and the <a href="/wiki/Megasporangium" class="mw-redirect" title="Megasporangium">megasporangium</a> is covered by three envelopes, like the <a href="/wiki/Ovary" title="Ovary">ovary</a> structure of angiosperm flowers. However, many other lines of evidence show that Gnetales is not related to angiosperms.<a href="#cite_note-Crepet-2000-115">[115]</a> The Mostly Male theory has a more genetic basis. Proponents of this theory point out that the gymnosperms have two very similar copies of the gene LFY, while angiosperms just have one. <a href="/wiki/Molecular_clock" title="Molecular clock">Molecular clock</a> analysis has shown that the other LFY paralog was lost in angiosperms around the same time as flower fossils become abundant, suggesting that this event might have led to floral evolution.<a href="#cite_note-135">[135]</a> According to this theory, loss of one of the LFY <a href="/wiki/Paralog" class="mw-redirect" title="Paralog">paralog</a> led to flowers that were more male, with the <a href="/wiki/Ovules" class="mw-redirect" title="Ovules">ovules</a> being expressed ectopically. These ovules initially performed the function of attracting <a href="/wiki/Pollinators" class="mw-redirect" title="Pollinators">pollinators</a>, but sometime later, may have been integrated into the core flower.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Mechanisms_and_players_in_evolution_of_plant_morphology">Mechanisms and players in evolution of plant morphology</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=24" title="Edit section: Mechanisms and players in evolution of plant morphology">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Microrna_secondary_structure.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a7/Microrna_secondary_structure.png/220px-Microrna_secondary_structure.png" decoding="async" width="220" height="138" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a7/Microrna_secondary_structure.png/330px-Microrna_secondary_structure.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a7/Microrna_secondary_structure.png/440px-Microrna_secondary_structure.png 2x" data-file-width="684" data-file-height="429" /></a><figcaption>The <a href="/wiki/Stem-loop" title="Stem-loop">stem-loop</a> <a href="/wiki/Secondary_structure" class="mw-redirect" title="Secondary structure">secondary structure</a> of a pre-microRNA from <a href="/wiki/Brassica_oleracea" title="Brassica oleracea">Brassica oleracea</a></figcaption></figure> While environmental factors are significantly responsible for evolutionary change, they act merely as agents for <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>. Change is inherently brought about via phenomena at the genetic level: <a href="/wiki/Mutations" class="mw-redirect" title="Mutations">mutations</a>, chromosomal rearrangements, and <a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">epigenetic</a> changes. While the general types of <a href="/wiki/Mutations" class="mw-redirect" title="Mutations">mutations</a> hold true across the living world, in plants, some other mechanisms have been implicated as highly significant.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <a href="/w/index.php?title=Genome_doubling&action=edit&redlink=1" class="new" title="Genome doubling (page does not exist)">Genome doubling</a> is a relatively common occurrence in plant evolution and results in <a href="/wiki/Polyploidy" title="Polyploidy">polyploidy</a>, which is consequently a common feature in plants. It is estimated that at least half (and probably all) plants have seen genome doubling in their history. Genome doubling entails <a href="/wiki/Gene_duplication" title="Gene duplication">gene duplication</a>, thus generating functional redundancy in most genes. The duplicated genes may attain new function, either by changes in expression pattern or changes in activity. Polyploidy and gene duplication are believed to be among the most powerful forces in evolution of plant form; though it is not known why <a href="/wiki/Genome" title="Genome">genome</a> doubling is such a frequent process in plants. One probable reason is the production of large amounts of <a href="/wiki/Secondary_metabolites" class="mw-redirect" title="Secondary metabolites">secondary metabolites</a> in plant cells. Some of them might interfere in the normal process of chromosomal segregation, causing genome duplication.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Maize-teosinte.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Maize-teosinte.jpg/150px-Maize-teosinte.jpg" decoding="async" width="150" height="93" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Maize-teosinte.jpg/225px-Maize-teosinte.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Maize-teosinte.jpg/300px-Maize-teosinte.jpg 2x" data-file-width="1037" data-file-height="643" /></a><figcaption>Top: <a href="/wiki/Teosinte" class="mw-redirect" title="Teosinte">teosinte</a>, bottom: <a href="/wiki/Maize" title="Maize">maize</a>, middle: maize-teosinte hybrid</figcaption></figure> In recent times, plants have been shown to possess significant <a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a> families, which are conserved across many plant lineages. In comparison to <a href="/wiki/Animal" title="Animal">animals</a>, while the number of plant miRNA families are lesser than animals, the size of each family is much larger. The <a href="/wiki/MiRNA" class="mw-redirect" title="MiRNA">miRNA</a> genes are also much more spread out in the genome than those in animals, where they are more clustered. It has been proposed that these miRNA families have expanded by duplications of chromosomal regions.<a href="#cite_note-136">[136]</a> Many miRNA genes involved in regulation of <a href="/wiki/Plant_development" title="Plant development">plant development</a> have been found to be quite conserved between plants studied.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <a href="/wiki/Domestication" title="Domestication">Domestication</a> of plants like <a href="/wiki/Maize" title="Maize">maize</a>, <a href="/wiki/Rice" title="Rice">rice</a>, <a href="/wiki/Barley" title="Barley">barley</a>, <a href="/wiki/Wheat" title="Wheat">wheat</a> etc. has also been a significant driving force in their evolution. Research concerning the origin of maize has found that it is a domesticated derivative of a wild plant from <a href="/wiki/Mexico" title="Mexico">Mexico</a> called <a href="/wiki/Teosinte" class="mw-redirect" title="Teosinte">teosinte</a>. Teosinte belongs to the <a href="/wiki/Genus" title="Genus">genus</a> Zea, just as maize, but bears very small <a href="/wiki/Inflorescence" title="Inflorescence">inflorescence</a>, 5–10 hard cobs and a highly branched and spread out stem. <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Cauliflower.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/25/Cauliflower.JPG/150px-Cauliflower.JPG" decoding="async" width="150" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/2/25/Cauliflower.JPG 1.5x" data-file-width="172" data-file-height="189" /></a><figcaption><a href="/wiki/Cauliflower" title="Cauliflower">Cauliflower</a> – Brassica oleracea var. botrytis</figcaption></figure> Crosses between a particular teosinte variety and maize yields fertile offspring that are intermediate in <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> between maize and teosinte. <a href="/wiki/QTL" class="mw-redirect" title="QTL">QTL</a> analysis has also revealed some loci that, when mutated in maize, yield a teosinte-like stem or teosinte-like cobs. <a href="/wiki/Molecular_clock" title="Molecular clock">Molecular clock</a> analysis of these genes estimates their origins to some 9,000 years ago, well in accordance with other records of maize domestication. It is believed that a small group of farmers must have selected some maize-like natural mutant of teosinte some 9,000 years ago in Mexico, and subjected it to continuous selection to yield the familiar maize plant of today.<a href="#cite_note-137">[137]</a> The edible cauliflower is a domesticated version of the wild plant <a href="/wiki/Brassica_oleracea" title="Brassica oleracea">Brassica oleracea</a>, which does not possess the dense undifferentiated <a href="/wiki/Inflorescence" title="Inflorescence">inflorescence</a>, called the curd, that cauliflower possesses. Cauliflower possesses a single mutation in a gene called CAL, controlling <a href="/wiki/Meristem" title="Meristem">meristem</a> differentiation into <a href="/wiki/Inflorescence" title="Inflorescence">inflorescence</a>. This causes the cells at the floral meristem to gain an undifferentiated identity and, instead of growing into a <a href="/wiki/Flower" title="Flower">flower</a>, they grow into a dense mass of inflorescence meristem cells in arrested development.<a href="#cite_note-138">[138]</a> This mutation has been selected through domestication since at least the time of the <a href="/wiki/Greeks" title="Greeks">Greek</a> empire. <div class="mw-heading mw-heading2"><h2 id="Evolution_of_photosynthetic_pathways">Evolution of photosynthetic pathways</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=25" title="Edit section: Evolution of photosynthetic pathways">edit</a>]</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:HatchSlackpathway2.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/HatchSlackpathway2.svg/300px-HatchSlackpathway2.svg.png" decoding="async" width="300" height="413" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/HatchSlackpathway2.svg/450px-HatchSlackpathway2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/HatchSlackpathway2.svg/600px-HatchSlackpathway2.svg.png 2x" data-file-width="705" data-file-height="970" /></a><figcaption>The C4 carbon concentrating mechanism</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Evolution_of_photosynthesis" title="Evolution of photosynthesis">Evolution of photosynthesis</a></div> The <a href="/wiki/C4_carbon_fixation" title="C4 carbon fixation">C4 metabolic pathway</a> is a valuable recent evolutionary innovation in plants, involving a complex set of adaptive changes to <a href="/wiki/Physiology" title="Physiology">physiology</a> and <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> patterns.<a href="#cite_note-Williams-2013-139">[139]</a> <a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a> is a complex chemical pathway facilitated by a range of <a href="/wiki/Enzymes" class="mw-redirect" title="Enzymes">enzymes</a> and co-enzymes. The <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> <a href="/wiki/RuBisCO" title="RuBisCO">RuBisCO</a> is responsible for "fixing" CO2 – that is, it attaches it to a carbon-based molecule to form a sugar, which can be used by the plant, releasing an oxygen molecule. However, the enzyme is notoriously inefficient, and, as ambient temperature rises, will increasingly fix oxygen instead of CO2 in a process called <a href="/wiki/Photorespiration" title="Photorespiration">photorespiration</a>. This is energetically costly as the plant has to use energy to turn the products of photorespiration back into a form that can react with CO2.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Concentrating_carbon">Concentrating carbon</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=26" title="Edit section: Concentrating carbon">edit</a>]</div> C4 plants evolved carbon concentrating mechanisms that work by increasing the concentration of CO2 around RuBisCO, and excluding oxygen, thereby increasing the efficiency of photosynthesis by decreasing photorespiration. The process of concentrating CO2 around RuBisCO requires more energy than allowing gases to <a href="/wiki/Diffusion" title="Diffusion">diffuse</a>, but under certain conditions – i.e. warm temperatures (>25 °C), low CO2 concentrations, or high oxygen concentrations – pays off in terms of the decreased loss of sugars through photorespiration. One type of C4 metabolism employs a so-called <a href="/wiki/Kranz_anatomy" class="mw-redirect" title="Kranz anatomy">Kranz anatomy</a>. This transports CO2 through an outer mesophyll layer, via a range of organic molecules, to the central bundle sheath cells, where the CO2 is released. In this way, CO2 is concentrated near the site of RuBisCO operation. Because RuBisCO is operating in an environment with much more CO2 than it otherwise would be, it performs more efficiently. A second mechanism, <a href="/wiki/CAM_photosynthesis" class="mw-redirect" title="CAM photosynthesis">CAM photosynthesis</a>, temporally separates photosynthesis from the action of RuBisCO. RuBisCO only operates during the day, when stomata are sealed and CO2 is provided by the breakdown of the chemical <a href="/wiki/Malate" class="mw-redirect" title="Malate">malate</a>. More CO2 is then harvested from the atmosphere when stomata open, during the cool, moist nights, reducing water loss. <div class="mw-heading mw-heading3"><h3 id="Evolutionary_record">Evolutionary record</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=27" title="Edit section: Evolutionary record">edit</a>]</div> These two pathways, with the same effect on RuBisCO, evolved a number of times independently – indeed, C4 alone arose 62 times in 18 different plant <a href="/wiki/Family_(biology)" title="Family (biology)">families</a>. A number of 'pre-adaptations' seem to have paved the way for C4, leading to its clustering in certain clades: it has most frequently been innovated in plants that already had features such as extensive vascular bundle sheath tissue.<a href="#cite_note-Christin-2012-140">[140]</a> Many potential evolutionary pathways resulting in the C4 <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> are possible and have been characterised using <a href="/wiki/Bayesian_inference" title="Bayesian inference">Bayesian inference</a>,<a href="#cite_note-Williams-2013-139">[139]</a> confirming that non-photosynthetic adaptations often provide evolutionary stepping stones for the further evolution of C4. The C4 construction is used by a subset of grasses, while CAM is employed by many succulents and <a href="/wiki/Cacti" class="mw-redirect" title="Cacti">cacti</a>. The C4 trait appears to have emerged during the <a href="/wiki/Oligocene" title="Oligocene">Oligocene</a>, around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=25–32">25 to 32</a> million years ago;<a href="#cite_note-Osborne-2006-141">[141]</a> however, they did not become ecologically significant until the <a href="/wiki/Miocene" title="Miocene">Miocene</a>, <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=6–7">6 to 7</a> million years ago.<a href="#cite_note-Retallack-1997-142">[142]</a> Remarkably, some charcoalified fossils preserve tissue organised into the Kranz anatomy, with intact bundle sheath cells,<a href="#cite_note-Thomasson-1986-143">[143]</a> allowing the presence C4 metabolism to be identified. Isotopic markers are used to deduce their distribution and significance. C3 plants preferentially use the lighter of two <a href="/wiki/Isotopes" class="mw-redirect" title="Isotopes">isotopes</a> of carbon in the atmosphere, 12C, which is more readily involved in the chemical pathways involved in its fixation. Because C4 metabolism involves a further chemical step, this effect is accentuated. Plant material can be <a href="/wiki/Mass_spectrometry" title="Mass spectrometry">analysed</a> to deduce the ratio of the heavier 13C to 12C. This ratio is denoted δ13 C. C3 plants are on average around 14‰ (parts per thousand) lighter than the atmospheric ratio, while C4 plants are about 28‰ lighter. The δ13 C of CAM plants depends on the percentage of carbon fixed at night relative to what is fixed in the day, being closer to C3 plants if they fix most carbon in the day and closer to C4 plants if they fix all their carbon at night.<a href="#cite_note-O'Leary-1988-144">[144]</a> Original fossil material in sufficient quantity to analyse the grass itself is scarce, but horses provide a good proxy. They were globally widespread in the period of interest, and browsed almost exclusively on grasses. There's an old phrase in isotope paleontology, "you are what you eat (plus a little bit)" – this refers to the fact that organisms reflect the isotopic composition of whatever they eat, plus a small adjustment factor. There is a good record of horse teeth throughout the globe, and their δ13 C record shows a sharp negative inflection around <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=6–7">6 to 7</a> million years ago, during the Messinian that is interpreted as resulting from the rise of C4 plants on a global scale.<a href="#cite_note-Retallack-1997-142">[142]</a> <div class="mw-heading mw-heading3"><h3 id="Advantage_of_C4">Advantage of C4</h3>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=28" title="Edit section: Advantage of C4">edit</a>]</div> While C4 enhances the efficiency of RuBisCO, the concentration of carbon is highly energy intensive. This means that C4 plants only have an advantage over C3 organisms in certain conditions: namely, high temperatures and low rainfall. C4 plants also need high levels of sunlight to thrive.<a href="#cite_note-Osborne-2009-145">[145]</a> Models suggest that, without wildfires removing shade-casting trees and shrubs, there would be no space for C4 plants.<a href="#cite_note-Bond-2005-146">[146]</a> But, wildfires have occurred for 400 million years. The Carboniferous (~<a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=300">300</a> million years ago) had notoriously high oxygen levels – almost enough to allow <a href="/wiki/Spontaneous_combustion" title="Spontaneous combustion">spontaneous combustion</a><a href="#cite_note-147">[147]</a> – and very low CO2, but no C4 isotopic signature has been found. There also does not seem to be a sudden trigger for the Miocene rise.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] During the Miocene, the atmosphere and climate were relatively stable. If anything, CO2 increased gradually from <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=14–9">14 to 9</a> million years ago before settling down to concentrations similar to the Holocene.<a href="#cite_note-Pagani-2005-148">[148]</a> This suggests that it did not have a key role in invoking C4 evolution.<a href="#cite_note-Osborne-2006-141">[141]</a> Grasses themselves (the group which would give rise to the most occurrences of C4) had probably been around for 60 million years or more, so had had plenty of time to evolve C4,<a href="#cite_note-Piperno-2005-149">[149]</a><a href="#cite_note-Prasad-2005-150">[150]</a> which, in any case, is present in a diverse range of groups and thus evolved independently. There is a strong signal of <a href="/wiki/Climate_change" title="Climate change">climate change</a> in South Asia;<a href="#cite_note-Osborne-2006-141">[141]</a> increasing aridity – hence increasing fire frequency and intensity – may have led to an increase in the importance of grasslands.<a href="#cite_note-Keeley-2005-151">[151]</a> However, this is difficult to reconcile with the North American record.<a href="#cite_note-Osborne-2006-141">[141]</a> It is possible that the signal is entirely biological, forced by the fire<a href="#cite_note-Retallack-2001-152">[152]</a> driven acceleration of grass evolution – which, both by increasing weathering and incorporating more carbon into sediments, reduced atmospheric CO2 levels.<a href="#cite_note-Retallack-2001-152">[152]</a> Finally, there is evidence that the onset of C4 from <a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=9–7">9 to 7</a> million years ago is a biased signal, which only holds true for North America, from where most samples originate; emerging evidence suggests that grasslands evolved to a dominant state at least 15Ma earlier in South America.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading2"><h2 id="Evolution_of_transcriptional_regulation">Evolution of transcriptional regulation</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=29" title="Edit section: Evolution of transcriptional regulation">edit</a>]</div> Transcription factors and transcriptional regulatory networks play key roles in plant development and stress responses, as well as their evolution. During plant landing, many novel transcription factor families emerged and are preferentially wired into the networks of multicellular development, reproduction, and organ development, contributing to more complex morphogenesis of land plants.<a href="#cite_note-Jin-2015-153">[153]</a> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_secondary_metabolism">Evolution of secondary metabolism</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=30" title="Edit section: Evolution of secondary metabolism">edit</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Azadirachtin.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Azadirachtin.png/250px-Azadirachtin.png" decoding="async" width="250" height="232" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Azadirachtin.png/375px-Azadirachtin.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Azadirachtin.png/500px-Azadirachtin.png 2x" data-file-width="789" data-file-height="731" /></a><figcaption>Structure of <a href="/wiki/Azadirachtin" title="Azadirachtin">Azadirachtin</a>, a terpenoid produced by the <a href="/wiki/Neem" class="mw-redirect" title="Neem">Neem</a> plant, which helps ward off microbes and insects. Many secondary metabolites have complex structures</figcaption></figure> <a href="/wiki/Secondary_metabolites" class="mw-redirect" title="Secondary metabolites">Secondary metabolites</a> are essentially low <a href="/wiki/Molecular_weight" class="mw-redirect" title="Molecular weight">molecular weight</a> compounds, sometimes having complex structures, that are not essential for the normal processes of <a href="/wiki/Cell_growth" title="Cell growth">growth</a>, <a href="/wiki/Biological_development" class="mw-redirect" title="Biological development">development</a>, or <a href="/wiki/Reproduction" title="Reproduction">reproduction</a>. They function in processes as diverse as <a href="/wiki/Immunity_(medical)" class="mw-redirect" title="Immunity (medical)">immunity</a>, anti-herbivory, <a href="/wiki/Pollinator" title="Pollinator">pollinator</a> attraction, <a href="/wiki/Communication" title="Communication">communication</a> between plants, maintaining <a href="/wiki/Symbiotic" class="mw-redirect" title="Symbiotic">symbiotic</a> associations with soil flora, or enhancing the rate of <a href="/wiki/Fertilization" class="mw-redirect" title="Fertilization">fertilization</a>, and hence are significant from the evo-devo perspective. Secondary metabolites are structurally and functionally diverse, and it is estimated that hundreds of thousands of enzymes might be involved in the process of producing them, with about 15–25% of the genome coding for these enzymes, and every species having its unique arsenal of secondary metabolites.<a href="#cite_note-154">[154]</a> Many of these metabolites, such as <a href="/wiki/Salicylic_acid" title="Salicylic acid">salicylic acid</a> are of medical significance to humans.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] The purpose of producing so many secondary metabolites, with a significant proportion of the <a href="/wiki/Metabolome" title="Metabolome">metabolome</a> devoted to this activity is unclear. It is postulated that most of these chemicals help in generating immunity and, in consequence, the diversity of these metabolites is a result of a constant <a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">arms race</a> between plants and their parasites. Some evidence supports this case. A central question involves the reproductive cost to maintaining such a large inventory of genes devoted to producing secondary metabolites. Various models have been suggested that probe into this aspect of the question, but a consensus on the extent of the cost has yet to be established;<a href="#cite_note-155">[155]</a> as it is still difficult to predict whether a plant with more secondary metabolites increases its survival or reproductive success compared to other plants in its vicinity.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Secondary metabolite production seems to have arisen quite early during evolution. In plants, they seem to have spread out using mechanisms including gene duplications or the evolution of novel genes. Furthermore, research has shown that diversity in some of these compounds may be positively selected for. Although the role of novel gene evolution in the evolution of secondary metabolism is clear, there are several examples where new metabolites have been formed by small changes in the reaction. For example, <a href="/w/index.php?title=Cyanogen_glycosides&action=edit&redlink=1" class="new" title="Cyanogen glycosides (page does not exist)">cyanogen glycosides</a> have been proposed to have evolved multiple times in different plant lineages. There are several such instances of <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergent evolution</a>. For example, enzymes for synthesis of <a href="/wiki/Limonene" title="Limonene">limonene</a> – a <a href="/wiki/Terpene" title="Terpene">terpene</a> – are more similar between angiosperms and gymnosperms than to their own terpene synthesis enzymes. This suggests independent evolution of the limonene biosynthetic pathway in these two lineages.<a href="#cite_note-156">[156]</a> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_plant-microbe_interactions">Evolution of plant-microbe interactions</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=31" title="Edit section: Evolution of plant-microbe interactions">edit</a>]</div> The origin of microbes on Earth, tracing back to the beginning of life more than 3.5 billion years ago, indicates that microbe-microbe interactions have continuously evolved and diversified over time, long before plants started to colonize land 450 million years ago. Therefore, it is likely that both intra- and inter-kingdom intermicrobial interactions represent strong drivers of the establishment of plant-associated <a href="/wiki/Microbial_consortia" class="mw-redirect" title="Microbial consortia">microbial consortia</a> at the soil-root interface. Nonetheless, it remains unclear to what extent these interactions in the rhizosphere/phyllosphere and in endophytic plant compartments (i.e., within the host) shape microbial assemblages in nature and whether microbial adaptation to plant habitats drive habitat-specific microbe-microbe interaction strategies that impact plant fitness. Furthermore, the contribution of competitive and cooperative microbe-microbe interactions to the overall community structure remains difficult to evaluate in nature due to the strong environmental noise.<a href="#cite_note-Hassani-2018-157">[157]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=32" title="Edit section: See also">edit</a>]</div> <ul><li><a href="/wiki/Herbivory#evolution" class="mw-redirect" title="Herbivory">Evolution of herbivory</a></li> <li><a href="/wiki/Evolutionary_history_of_life" class="mw-redirect" title="Evolutionary history of life">Evolutionary history of life</a></li> <li><a href="/wiki/Paleobotany" title="Paleobotany">Paleobotany</a></li> <li><a href="/wiki/Plant_evolutionary_developmental_biology" title="Plant evolutionary developmental biology">Plant evolutionary developmental biology</a></li> <li><a href="/wiki/Timeline_of_plant_evolution" title="Timeline of plant evolution">Timeline of plant evolution</a><div style="clear:both;" class=""></div></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Evolutionary_history_of_plants&action=edit&section=33" title="Edit section: References">edit</a>]</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-Stewart-1993-1">^ <a href="#cite_ref-Stewart-1993_1-0">a</a> <a href="#cite_ref-Stewart-1993_1-1">b</a> <a href="#cite_ref-Stewart-1993_1-2">c</a> <a href="#cite_ref-Stewart-1993_1-3">d</a> <a href="#cite_ref-Stewart-1993_1-4">e</a> <a href="#cite_ref-Stewart-1993_1-5">f</a> <a href="#cite_ref-Stewart-1993_1-6">g</a> <a href="#cite_ref-Stewart-1993_1-7">h</a> <a href="#cite_ref-Stewart-1993_1-8">i</a> <a href="#cite_ref-Stewart-1993_1-9">j</a> <a href="#cite_ref-Stewart-1993_1-10">k</a> <a href="#cite_ref-Stewart-1993_1-11">l</a> <a href="#cite_ref-Stewart-1993_1-12">m</a> <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="CITEREFStewartRothwell1993" class="citation book cs1">Stewart, W.N.; Rothwell, G.W. (1993). Paleobotany and the evolution of plants (2 ed.). 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Science. 296 (5569): 899–904. <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/2002Sci...296..899S">2002Sci...296..899S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.1069439">10.1126/science.1069439</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/11988572">11988572</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:1910388">1910388</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Archaefructaceae%2C+a+New+Basal+Angiosperm+Family&rft.volume=296&rft.issue=5569&rft.pages=899-904&rft.date=2002&rft_id=info%3Adoi%2F10.1126%2Fscience.1069439&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A1910388%23id-name%3DS2CID&rft_id=info%3Apmid%2F11988572&rft_id=info%3Abibcode%2F2002Sci...296..899S&rft.au=Sun%2C+G.&rft.au=Ji%2C+Q.&rft.au=Dilcher%2C+D.L.&rft.au=Zheng%2C+S.&rft.au=Nixon%2C+K.C.&rft.au=Wang%2C+X.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolutionary+history+of+plants" class="Z3988"> </li> <li id="cite_note-Doyle-2003-117"><a href="#cite_ref-Doyle-2003_117-0">^</a> In fact, <a href="/wiki/Archaefructus" title="Archaefructus">Archaefructus</a> probably didn't bear true flowers: see <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFriis,_E.M.Doyle,_J.A.Endress,_P.K.Leng,_Q.2003" class="citation journal cs1">Friis, E.M.; Doyle, J.A.; Endress, P.K.; Leng, Q. 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Microbiome, 6(1): 58. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1186%2Fs40168-018-0445-0">10.1186/s40168-018-0445-0</a>. <a href="/wiki/File:CC_BY_icon.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e1/CC_BY_icon.svg/50px-CC_BY_icon.svg.png" decoding="async" width="50" height="18" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e1/CC_BY_icon.svg/75px-CC_BY_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e1/CC_BY_icon.svg/100px-CC_BY_icon.svg.png 2x" data-file-width="88" data-file-height="31" /></a> Material was copied from this source, which is available under a <a rel="nofollow" class="external text" href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution 4.0 International 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title="Bryology">Bryology</a></li> <li><a href="/wiki/Dendrology" title="Dendrology">Dendrology</a></li> <li><a href="/wiki/Ethnobotany" title="Ethnobotany">Ethnobotany</a></li> <li><a href="/wiki/Paleobotany" title="Paleobotany">Paleobotany</a></li> <li><a href="/wiki/Phycology" title="Phycology">Phycology</a></li> <li><a href="/wiki/Phytochemistry" title="Phytochemistry">Phytochemistry</a></li> <li><a href="/wiki/Phytogeography" title="Phytogeography">Phytogeography</a> <ul><li><a href="/wiki/Geobotanical_prospecting" title="Geobotanical prospecting">Geobotany</a></li></ul></li> <li><a href="/wiki/Plant_anatomy" title="Plant anatomy">Plant anatomy</a></li> <li><a href="/wiki/Plant_ecology" title="Plant ecology">Plant ecology</a></li> <li><a href="/wiki/Plant_pathology" title="Plant pathology">Plant pathology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant" title="Plant">Plant</a> groups</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/Algae" title="Algae">Algae</a></li> <li><a href="/wiki/Archaeplastida" title="Archaeplastida">Archaeplastida</a></li> <li><a href="/wiki/Bryophyte" title="Bryophyte">Bryophyte</a></li> <li><a href="/wiki/Non-vascular_plant" title="Non-vascular plant">Non-vascular plants</a></li> <li><a href="/wiki/Vascular_plant" title="Vascular plant">Vascular plants</a></li> <li><a href="/wiki/Fern" title="Fern">Fern</a></li> <li><a href="/wiki/Lycophyte" title="Lycophyte">Lycophyte</a></li> <li><a href="/wiki/Spermatophyte" class="mw-redirect" title="Spermatophyte">Spermatophytes</a></li> <li><a href="/wiki/Gymnosperm" title="Gymnosperm">Gymnosperm</a></li> <li><a href="/wiki/Flowering_plant" title="Flowering plant">Angiosperm</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_anatomy" title="Plant anatomy">Plant anatomy</a> <div class="hlist"><ul><li><a href="/wiki/Plant_morphology" title="Plant morphology">Plant morphology</a> (<a href="/wiki/Glossary_of_plant_morphology" title="Glossary of plant morphology">glossary</a>)</li></ul></div></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%"><a href="/wiki/Plant_cell" title="Plant cell">Plant cells</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/Cell_wall" title="Cell wall">Cell wall</a></li> <li><a href="/wiki/Phragmoplast" title="Phragmoplast">Phragmoplast</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li> <li><a href="/wiki/Plasmodesma" title="Plasmodesma">Plasmodesma</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%"><a href="/wiki/Tissue_(biology)" title="Tissue (biology)">Tissues</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/Cork_cambium" title="Cork cambium">Cork</a></li> <li><a href="/wiki/Ground_tissue" title="Ground tissue">Ground tissue</a> <ul><li><a href="/wiki/Leaf#Mesophyll" title="Leaf">Mesophyll</a></li></ul></li> <li><a href="/wiki/Meristem" title="Meristem">Meristem</a></li> <li><a href="/wiki/Storage_organ" title="Storage organ">Storage organs</a></li> <li><a href="/wiki/Vascular_tissue" title="Vascular tissue">Vascular tissue</a> <ul><li><a href="/wiki/Vascular_bundle" title="Vascular bundle">Vascular bundle</a></li></ul></li> <li><a href="/wiki/Wood" title="Wood">Wood</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%">Vegetative</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/Bulb" title="Bulb">Bulb</a></li> <li><a href="/wiki/Root" title="Root">Root</a></li> <li><a href="/wiki/Rhizoid" title="Rhizoid">Rhizoid</a></li> <li><a href="/wiki/Rhizome" title="Rhizome">Rhizome</a></li> <li><a href="/wiki/Shoot_(botany)" title="Shoot (botany)">Shoot</a> <ul><li><a href="/wiki/Bud" title="Bud">Bud</a></li> <li><a href="/wiki/Leaf" title="Leaf">Leaf</a> <ul><li><a href="/wiki/Cataphyll" title="Cataphyll">Cataphyll</a></li> <li><a href="/wiki/Petiole_(botany)" title="Petiole (botany)">Petiole</a></li></ul></li> <li><a href="/wiki/Sessility_(botany)" title="Sessility (botany)">Sessility</a></li> <li><a href="/wiki/Plant_stem" title="Plant stem">Stem</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%"><a href="/wiki/Plant_reproductive_morphology" title="Plant reproductive morphology">Reproductive</a> (incl. Flower)</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/Archegonium" title="Archegonium">Archegonium</a></li> <li><a href="/wiki/Antheridium" title="Antheridium">Antheridium</a></li> <li><a href="/wiki/Stamen" title="Stamen">Androecium</a> <ul><li><a href="/wiki/Pollen" title="Pollen">Pollen</a></li> <li><a href="/wiki/Stamen" title="Stamen">Stamen</a> <ul><li><a href="/wiki/Anther" class="mw-redirect" title="Anther">Anther</a></li> <li><a href="/wiki/Stamen#Morphology_and_terminology" title="Stamen">Filament</a></li></ul></li> <li><a href="/wiki/Staminode" title="Staminode">Staminode</a></li> <li><a href="/wiki/Tapetum_(botany)" title="Tapetum (botany)">Tapetum</a></li></ul></li> <li><a href="/wiki/Flower" title="Flower">Flower</a> <ul><li><a href="/wiki/Aestivation_(botany)" title="Aestivation (botany)">Aestivation</a></li> <li><a href="/wiki/ABC_model_of_flower_development" title="ABC model of flower development">Flower development</a></li> <li><a href="/wiki/Floral_diagram" title="Floral diagram">Floral diagram</a></li> <li><a href="/wiki/Floral_formula" title="Floral formula">Floral formula</a></li> <li><a href="/wiki/Floral_symmetry" title="Floral symmetry">Floral symmetry</a></li> <li><a href="/wiki/Whorl_(botany)" title="Whorl (botany)">Whorl</a></li></ul></li> <li><a href="/wiki/Fruit" title="Fruit">Fruit</a> <ul><li><a href="/wiki/Fruit_anatomy" class="mw-redirect" title="Fruit anatomy">Anatomy</a></li> <li><a href="/wiki/Berry_(botany)" title="Berry (botany)">Berry</a></li> <li><a href="/wiki/Capsule_(fruit)" title="Capsule (fruit)">Capsule</a></li> <li><a href="/wiki/Nut_(fruit)" title="Nut (fruit)">Nut</a></li> <li><a href="/wiki/Pyrena" title="Pyrena">Pyrena</a></li> <li><a href="/wiki/Seed" title="Seed">Seed</a> <ul><li><a href="/wiki/Seed_dispersal" title="Seed dispersal">Dispersal</a></li> <li><a href="/wiki/Endosperm" title="Endosperm">Endosperm</a></li></ul></li></ul></li> <li><a href="/wiki/Gametophyte" title="Gametophyte">Gametophyte</a></li> <li><a href="/wiki/Column_(botany)" title="Column (botany)">Gynandrium</a></li> <li><a href="/wiki/Gynoecium" title="Gynoecium">Gynoecium</a> <ul><li><a href="/wiki/Carpel" class="mw-redirect" title="Carpel">Carpel</a> <ul><li><a href="/wiki/Ovary_(botany)" title="Ovary (botany)">Ovary</a> <ul><li><a href="/wiki/Locule" title="Locule">Locule</a></li> <li><a href="/wiki/Ovule" title="Ovule">Ovule</a></li></ul></li> <li><a href="/wiki/Stigma_(botany)" title="Stigma (botany)">Stigma</a></li> <li><a href="/wiki/Style_(botany)" title="Style (botany)">Style</a></li></ul></li></ul></li> <li><a href="/wiki/Hypanthium" title="Hypanthium">Hypanthium (Floral cup)</a></li> <li><a href="/wiki/Inflorescence" title="Inflorescence">Inflorescence</a> <ul><li><a href="/wiki/Bract" title="Bract">Bract</a></li> <li><a href="/wiki/Pedicel_(botany)" title="Pedicel (botany)">Pedicellate</a></li> <li><a href="/wiki/Raceme" title="Raceme">Raceme</a></li> <li><a href="/wiki/Umbel" title="Umbel">Umbel</a></li></ul></li> <li><a href="/wiki/Perianth" title="Perianth">Perianth</a> <ul><li><a href="/wiki/Tepal" title="Tepal">Tepal</a></li> <li><a href="/wiki/Petal" title="Petal">Petal</a></li> <li><a href="/wiki/Sepal" title="Sepal">Sepal</a></li></ul></li> <li><a href="/wiki/Embryo#Plant_embryos" title="Embryo">Plant embryo</a></li> <li><a href="/wiki/Receptacle_(botany)" title="Receptacle (botany)">Receptacle</a></li> <li><a href="/wiki/Sporophyll" title="Sporophyll">Sporophyll</a></li> <li><a href="/wiki/Sporophyte" title="Sporophyte">Sporophyte</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #D4EED4;;width:1%">Surface structures</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_cuticle" title="Plant cuticle">Cuticle</a></li> <li><a href="/wiki/Epicuticular_wax" title="Epicuticular wax">Epicuticular wax</a></li> <li><a href="/wiki/Epidermis_(botany)" title="Epidermis (botany)">Epidermis</a></li> <li><a href="/wiki/Nectar" title="Nectar">Nectar</a></li> <li><a href="/wiki/Stoma" title="Stoma">Stoma</a></li> <li><a href="/wiki/Thorns,_spines,_and_prickles" title="Thorns, spines, and prickles">Thorns, spines, and prickles</a></li> <li><a href="/wiki/Trichome" title="Trichome">Trichome</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_physiology" title="Plant physiology">Plant physiology</a> Materials</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/Aleurone" title="Aleurone">Aleurone</a></li> <li><a href="/wiki/Apical_dominance" title="Apical dominance">Apical dominance</a></li> <li><a href="/wiki/Bulk_movement" title="Bulk movement">Bulk flow</a></li> <li><a href="/wiki/Cellulose" title="Cellulose">Cellulose</a></li> <li><a href="/wiki/Plant_nutrition" title="Plant nutrition">Nutrition</a></li> <li><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a> <ul><li><a href="/wiki/Chlorophyll" title="Chlorophyll">Chlorophyll</a></li></ul></li> <li><a href="/wiki/Phytomelanin" title="Phytomelanin">Phytomelanin</a></li> <li><a href="/wiki/Plant_hormone" title="Plant hormone">Plant hormones</a></li> <li><a href="/wiki/Respiratory_system#Plants" title="Respiratory system">Respiration</a> <ul><li><a href="/wiki/Gas_exchange#Plants" title="Gas exchange">Gas Exchange</a></li> <li><a href="/wiki/Cellular_respiration" title="Cellular respiration">Cellular respiration</a></li></ul></li> <li><a href="/wiki/Sap" title="Sap">Sap</a></li> <li><a href="/wiki/Starch" title="Starch">Starch</a></li> <li><a href="/wiki/Sugar" title="Sugar">Sugar</a></li> <li><a href="/wiki/Transpiration" title="Transpiration">Transpiration</a></li> <li><a href="/wiki/Turgor_pressure" title="Turgor pressure">Turgor pressure</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%">Plant growth and habit</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/Habit_(biology)#Structure" title="Habit (biology)">Habit</a> <ul><li><a href="/wiki/Cushion_plant" title="Cushion plant">Cushion plants</a></li> <li><a href="/wiki/Rosette_(botany)" title="Rosette (botany)">Rosettes</a></li> <li><a href="/wiki/Shrub" title="Shrub">Shrubs</a> <ul><li><a href="/wiki/Prostrate_shrub" title="Prostrate shrub">Prostrate shrubs</a></li> <li><a href="/wiki/Subshrub" title="Subshrub">Subshrubs</a></li></ul></li> <li><a href="/wiki/Succulent_plant" title="Succulent plant">Succulent plants</a></li> <li><a href="/wiki/Tree" title="Tree">Trees</a></li> <li><a href="/wiki/Vine" title="Vine">Vines</a> <ul><li><a href="/wiki/Liana" title="Liana">Lianas</a></li></ul></li></ul></li> <li><a href="/wiki/Herbaceous_plant" title="Herbaceous plant">Herbaceous plants</a></li> <li><a href="/wiki/Secondary_growth" title="Secondary growth">Secondary growth</a></li> <li><a href="/wiki/Woody_plant" title="Woody plant">Woody plants</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_reproduction" title="Plant reproduction">Reproduction</a> <div class="hlist"><ul><li><a href="/wiki/Plant_evolution" title="Plant evolution">Evolution</a></li><li><a href="/wiki/Plant_ecology" title="Plant ecology">Ecology</a></li></ul></div></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/Alternation_of_generations" title="Alternation of generations">Alternation of generations</a></li> <li><a href="/wiki/Double_fertilization" title="Double fertilization">Double fertilization</a></li> <li><a href="/wiki/Plant_evolutionary_developmental_biology" title="Plant evolutionary developmental biology">Evolutionary development</a></li> <li><a class="mw-selflink selflink">Evolutionary history</a> <ul><li><a href="/wiki/Timeline_of_plant_evolution" title="Timeline of plant evolution">timeline</a></li></ul></li> <li><a href="/wiki/Flora" title="Flora">Flora</a></li> <li><a href="/wiki/Germination" title="Germination">Germination</a></li> <li><a href="/wiki/Pollination" title="Pollination">Pollination</a> <ul><li><a href="/wiki/Artificial_pollination" class="mw-redirect" title="Artificial pollination">Artificial</a></li> <li><a href="/wiki/Pollinator" title="Pollinator">Pollinators</a></li> <li><a href="/wiki/Pollen_tube" title="Pollen tube">Pollen tube</a></li> <li><a href="/wiki/Self-pollination" title="Self-pollination">Self</a></li></ul></li> <li><a href="/wiki/Sporangium" title="Sporangium">Sporangium</a> <ul><li><a href="/wiki/Microsporangia" class="mw-redirect" title="Microsporangia">Microsporangia</a> <ul><li><a href="/wiki/Microspore" title="Microspore">Microspore</a></li></ul></li> <li><a href="/wiki/Sporangium" title="Sporangium">Megasporangium</a> <ul><li><a href="/wiki/Megaspore" title="Megaspore">Megaspore</a></li></ul></li> <li><a href="/wiki/Spore" title="Spore">Spore</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Plant_taxonomy" title="Plant taxonomy">Plant taxonomy</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/Taxonomy_(biology)" title="Taxonomy (biology)">Biological classification</a></li> <li><a href="/wiki/Botanical_nomenclature" title="Botanical nomenclature">Botanical nomenclature</a> <ul><li><a href="/wiki/Botanical_name" title="Botanical name">Botanical name</a></li> <li><a href="/wiki/Correct_name" title="Correct name">Correct name</a></li> <li><a href="/wiki/Author_citation_(botany)" title="Author citation (botany)">Author citation</a></li> <li><a href="/wiki/International_Code_of_Nomenclature_for_algae,_fungi,_and_plants" title="International Code of Nomenclature for algae, fungi, and plants">International Code of Nomenclature (ICN)</a></li> <li><a href="/wiki/International_Code_of_Nomenclature_for_Cultivated_Plants" title="International Code of Nomenclature for Cultivated Plants">ICN for Cultivated Plants (ICNCP)</a></li></ul></li> <li><a href="/wiki/Cultivated_plant_taxonomy" title="Cultivated plant taxonomy">Cultivated plant taxonomy</a> <ul><li><a href="/wiki/Citrus_taxonomy" title="Citrus taxonomy">Citrus taxonomy</a></li> <li><a href="/wiki/Cultigen" title="Cultigen">Cultigen</a> <ul><li><a href="/wiki/Cultivar" title="Cultivar">Cultivar</a></li> <li><a href="/wiki/Cultivar_group" title="Cultivar group">Group</a></li> <li><a href="/wiki/Grex_(horticulture)" title="Grex (horticulture)">Grex</a></li></ul></li></ul></li> <li><a href="/wiki/History_of_plant_systematics" title="History of plant systematics">History of plant systematics</a></li> <li><a href="/wiki/Herbarium" title="Herbarium">Herbarium</a></li> <li><a href="/wiki/International_Association_for_Plant_Taxonomy" title="International Association for Plant Taxonomy">International Association for Plant Taxonomy</a> (IAPT)</li> <li><a href="/wiki/List_of_systems_of_plant_taxonomy" title="List of systems of plant taxonomy">Plant taxonomy systems</a></li> <li><a href="/wiki/Taxonomic_rank" title="Taxonomic rank">Taxonomic rank</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%">Practice</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/Agronomy" title="Agronomy">Agronomy</a></li> <li><a href="/wiki/Floriculture" title="Floriculture">Floriculture</a></li> <li><a href="/wiki/Forestry" title="Forestry">Forestry</a></li> <li><a href="/wiki/Horticulture" title="Horticulture">Horticulture</a></li> <li><a href="/wiki/Phytochemical" title="Phytochemical">Phytochemical</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><div class="hlist"><ul><li>Lists</li><li>Related topics</li></ul></div></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Glossary_of_botanical_terms" title="Glossary of botanical terms">Botanical terms</a></li> <li><a href="/wiki/List_of_botanists" title="List of botanists">Botanists</a> <ul><li><a href="/wiki/List_of_botanists_by_author_abbreviation_(W%E2%80%93Z)" title="List of botanists by author abbreviation (W–Z)">by author abbreviation</a></li></ul></li> <li><a href="/wiki/Botanical_expeditions" title="Botanical expeditions">Botanical expeditions</a></li> <li><a href="/wiki/List_of_individual_trees" title="List of individual trees">Individual trees</a></li> <li><a href="/wiki/Lists_of_plants" title="Lists of plants">Plants</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="background: #C3EEC3;"><div> <ul><li><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" /> <a href="/wiki/Category:Botany" title="Category:Botany">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 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 class="mw-selflink selflink">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></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><img alt="" 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