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class="vector-toc-list"> </ul> </li> <li id="toc-Opening_and_closing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Opening_and_closing"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Opening and closing</span> </div> </a> <ul id="toc-Opening_and_closing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Inferring_stomatal_behavior_from_gas_exchange" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inferring_stomatal_behavior_from_gas_exchange"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Inferring stomatal behavior from gas exchange</span> </div> </a> <ul id="toc-Inferring_stomatal_behavior_from_gas_exchange-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Development" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Development"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Development</span> </div> </a> <ul id="toc-Development-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Types" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Types"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Types</span> </div> </a> <ul id="toc-Types-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stomatal_crypts" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Stomatal_crypts"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Stomatal crypts</span> </div> </a> <ul id="toc-Stomatal_crypts-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stomata_as_pathogenic_pathways" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Stomata_as_pathogenic_pathways"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Stomata as pathogenic pathways</span> </div> </a> <ul id="toc-Stomata_as_pathogenic_pathways-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stomata_and_climate_change" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Stomata_and_climate_change"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Stomata and climate change</span> </div> </a> <button aria-controls="toc-Stomata_and_climate_change-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Stomata and climate change subsection</span> </button> <ul id="toc-Stomata_and_climate_change-sublist" class="vector-toc-list"> <li id="toc-Response_of_stomata_to_environmental_factors" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Response_of_stomata_to_environmental_factors"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Response of stomata to environmental factors</span> </div> </a> <ul id="toc-Response_of_stomata_to_environmental_factors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Future_adaptations_during_climate_change" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Future_adaptations_during_climate_change"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Future adaptations during climate change</span> </div> </a> <ul id="toc-Future_adaptations_during_climate_change-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Agricultural_implications" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Agricultural_implications"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Agricultural implications</span> </div> </a> <ul id="toc-Agricultural_implications-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Stoma</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 59 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-59" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">59 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AB%D8%BA%D8%B1_(%D9%86%D8%A8%D8%A7%D8%AA)" title="ثغر (نبات) – Arabic" lang="ar" hreflang="ar" data-title="ثغر (نبات)" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Estoma" title="Estoma – Asturian" lang="ast" hreflang="ast" data-title="Estoma" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AA%E0%A6%A4%E0%A7%8D%E0%A6%B0%E0%A6%B0%E0%A6%A8%E0%A7%8D%E0%A6%A7%E0%A7%8D%E0%A6%B0" title="পত্ররন্ধ্র – Bangla" lang="bn" hreflang="bn" data-title="পত্ররন্ধ্র" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A3%D1%81%D1%82%D0%B8%D1%86%D0%B0" title="Устица – Bulgarian" lang="bg" hreflang="bg" data-title="Устица" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Stoma" title="Stoma – Bosnian" lang="bs" hreflang="bs" data-title="Stoma" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Estoma" title="Estoma – Catalan" lang="ca" hreflang="ca" data-title="Estoma" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://cs.wikipedia.org/wiki/Pr%C5%AFduch" title="Průduch – Czech" lang="cs" hreflang="cs" data-title="Průduch" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Spalte%C3%A5bning" title="Spalteåbning – Danish" lang="da" hreflang="da" data-title="Spalteåbning" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Stoma_(Botanik)" title="Stoma (Botanik) – German" lang="de" hreflang="de" data-title="Stoma (Botanik)" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/%C3%95hul%C3%B5he" title="Õhulõhe – Estonian" lang="et" hreflang="et" data-title="Õhulõhe" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%A3%CF%84%CF%8C%CE%BC%CE%B1_(%CE%B2%CE%BF%CF%84%CE%B1%CE%BD%CE%B9%CE%BA%CE%AE)" title="Στόμα (βοτανική) – Greek" lang="el" hreflang="el" data-title="Στόμα (βοτανική)" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Estoma" title="Estoma – Spanish" lang="es" hreflang="es" data-title="Estoma" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Stomo" title="Stomo – Esperanto" lang="eo" hreflang="eo" data-title="Stomo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Estoma" title="Estoma – Basque" lang="eu" hreflang="eu" data-title="Estoma" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B1%D9%88%D8%B2%D9%86%D9%87_%D9%87%D9%88%D8%A7%DB%8C%DB%8C" title="روزنه هوایی – Persian" lang="fa" hreflang="fa" data-title="روزنه هوایی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Stomate" title="Stomate – French" lang="fr" hreflang="fr" data-title="Stomate" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/St%C3%B3ma%C3%AD" title="Stómaí – Irish" lang="ga" hreflang="ga" data-title="Stómaí" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Estoma" title="Estoma – Galician" lang="gl" hreflang="gl" data-title="Estoma" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EA%B8%B0%EA%B3%B5_(%EC%8B%9D%EB%AC%BC%ED%95%99)" title="기공 (식물학) – Korean" lang="ko" hreflang="ko" data-title="기공 (식물학)" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%B0%E0%A4%A8%E0%A5%8D%E0%A4%A7%E0%A5%8D%E0%A4%B0_(%E0%A4%B5%E0%A4%BE%E0%A4%A8%E0%A4%B8%E0%A5%8D%E0%A4%AA%E0%A4%A4%E0%A4%BF%E0%A4%95%E0%A5%80)" title="रन्ध्र (वानस्पतिकी) – Hindi" lang="hi" hreflang="hi" data-title="रन्ध्र (वानस्पतिकी)" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Stoma" title="Stoma – Indonesian" lang="id" hreflang="id" data-title="Stoma" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Stoma" title="Stoma – Italian" lang="it" hreflang="it" data-title="Stoma" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A4%D7%99%D7%95%D7%A0%D7%99%D7%AA" title="פיונית – Hebrew" lang="he" hreflang="he" data-title="פיונית" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Stomata" title="Stomata – Javanese" lang="jv" hreflang="jv" data-title="Stomata" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%AA%E0%B2%A4%E0%B3%8D%E0%B2%B0%E0%B2%B0%E0%B2%82%E0%B2%A7%E0%B3%8D%E0%B2%B0" title="ಪತ್ರರಂಧ್ರ – Kannada" lang="kn" hreflang="kn" data-title="ಪತ್ರರಂಧ್ರ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%91%E1%83%90%E1%83%92%E1%83%94" title="ბაგე – Georgian" lang="ka" hreflang="ka" data-title="ბაგე" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%A2%D1%8B%D0%BD%D1%8B%D1%81_%D1%81%D0%B0%D2%A3%D1%8B%D0%BB%D0%B0%D1%83%D0%BB%D0%B0%D1%80%D1%8B" title="Тыныс саңылаулары – Kazakh" lang="kk" hreflang="kk" data-title="Тыныс саңылаулары" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/Stomata" title="Stomata – Swahili" lang="sw" hreflang="sw" data-title="Stomata" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target"><span>Kiswahili</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Estomat" title="Estomat – Haitian Creole" lang="ht" hreflang="ht" data-title="Estomat" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Stoma" title="Stoma – Latin" lang="la" hreflang="la" data-title="Stoma" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Atv%C4%81rsn%C4%ABtes" title="Atvārsnītes – Latvian" lang="lv" hreflang="lv" data-title="Atvārsnītes" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/%C5%BDiotel%C4%97" title="Žiotelė – Lithuanian" lang="lt" hreflang="lt" data-title="Žiotelė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%86%E0%B4%B8%E0%B5%8D%E0%B4%AF%E0%B4%B0%E0%B4%A8%E0%B5%8D%E0%B4%A7%E0%B5%8D%E0%B4%B0%E0%B4%82" title="ആസ്യരന്ധ്രം – Malayalam" lang="ml" hreflang="ml" data-title="ആസ്യരന്ധ്രം" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%A4%E0%A5%8D%E0%A4%B5%E0%A4%97%E0%A5%8D%E0%A4%B0%E0%A4%82%E0%A4%A7%E0%A5%8D%E0%A4%B0%E0%A5%87" title="त्वग्रंध्रे – Marathi" lang="mr" hreflang="mr" data-title="त्वग्रंध्रे" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target"><span>मराठी</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Stoma" title="Stoma – Malay" lang="ms" hreflang="ms" data-title="Stoma" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%A1%D1%82%D0%BE%D0%BC%D0%B0" title="Стома – Mongolian" lang="mn" hreflang="mn" data-title="Стома" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Huidmondje" title="Huidmondje – Dutch" lang="nl" hreflang="nl" data-title="Huidmondje" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E6%B0%97%E5%AD%94" title="気孔 – Japanese" lang="ja" hreflang="ja" data-title="気孔" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Spalte%C3%A5pning" title="Spalteåpning – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Spalteåpning" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Spalteopning" title="Spalteopning – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Spalteopning" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Aparat_szparkowy" title="Aparat szparkowy – Polish" lang="pl" hreflang="pl" data-title="Aparat szparkowy" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Est%C3%B4mato" title="Estômato – Portuguese" lang="pt" hreflang="pt" data-title="Estômato" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A3%D1%81%D1%82%D1%8C%D0%B8%D1%86%D0%B5" title="Устьице – Russian" lang="ru" hreflang="ru" data-title="Устьице" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Stomata" title="Stomata – Simple English" lang="en-simple" hreflang="en-simple" data-title="Stomata" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Prieduch_(botanika)" title="Prieduch (botanika) – Slovak" lang="sk" hreflang="sk" data-title="Prieduch (botanika)" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Listna_re%C5%BEa" title="Listna reža – Slovenian" lang="sl" hreflang="sl" data-title="Listna reža" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AF%DB%95%D9%85%DB%8C%D9%84%DB%95" title="دەمیلە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="دەمیلە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Stoma" title="Stoma – Serbian" lang="sr" hreflang="sr" data-title="Stoma" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Ilmarako" title="Ilmarako – Finnish" lang="fi" hreflang="fi" data-title="Ilmarako" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Klyv%C3%B6ppning" title="Klyvöppning – Swedish" lang="sv" hreflang="sv" data-title="Klyvöppning" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl badge-Q70894304 mw-list-item" title=""><a href="https://tl.wikipedia.org/wiki/Stoma" title="Stoma – Tagalog" lang="tl" hreflang="tl" data-title="Stoma" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%87%E0%AE%B2%E0%AF%88_%E0%AE%A4%E0%AF%81%E0%AE%B3%E0%AF%88" title="இலை துளை – Tamil" lang="ta" hreflang="ta" data-title="இலை துளை" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%9B%E0%B8%B2%E0%B8%81%E0%B9%83%E0%B8%9A" title="ปากใบ – Thai" lang="th" hreflang="th" data-title="ปากใบ" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Stoma" title="Stoma – Turkish" lang="tr" hreflang="tr" data-title="Stoma" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9F%D1%80%D0%BE%D0%B4%D0%B8%D1%85" title="Продих – Ukrainian" lang="uk" hreflang="uk" data-title="Продих" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Kh%C3%AD_kh%E1%BB%95ng" title="Khí khổng – Vietnamese" lang="vi" hreflang="vi" data-title="Khí khổng" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E6%B0%94%E5%AD%94" title="气孔 – Wu" lang="wuu" hreflang="wuu" data-title="气孔" data-language-autonym="吴语" data-language-local-name="Wu" 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</div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">In plants, a variable pore between paired guard cells</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Stroma_(disambiguation)" class="mw-redirect mw-disambig" title="Stroma (disambiguation)">Stroma</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">For natural and surgically created body openings, see <a href="/wiki/Stoma_(medicine)" title="Stoma (medicine)">Stoma (medicine)</a>.</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Tomato_leaf_stomate_1-color.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tomato_leaf_stomate_1-color.jpg/250px-Tomato_leaf_stomate_1-color.jpg" decoding="async" width="220" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tomato_leaf_stomate_1-color.jpg/330px-Tomato_leaf_stomate_1-color.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tomato_leaf_stomate_1-color.jpg/500px-Tomato_leaf_stomate_1-color.jpg 2x" data-file-width="1024" data-file-height="1049" /></a><figcaption>Stoma in a <a href="/wiki/Tomato" title="Tomato">tomato</a> leaf shown via colorized <a href="/wiki/Scanning_electron_microscope" title="Scanning electron microscope">scanning electron microscope</a> image</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:HPIM0188-ligusterblad.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/02/HPIM0188-ligusterblad.jpg/250px-HPIM0188-ligusterblad.jpg" decoding="async" width="220" height="164" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/02/HPIM0188-ligusterblad.jpg/330px-HPIM0188-ligusterblad.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/02/HPIM0188-ligusterblad.jpg/500px-HPIM0188-ligusterblad.jpg 2x" data-file-width="576" data-file-height="430" /></a><figcaption>A stoma in horizontal cross section</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:LeafUndersideWithStomata.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/LeafUndersideWithStomata.jpg/250px-LeafUndersideWithStomata.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/LeafUndersideWithStomata.jpg/330px-LeafUndersideWithStomata.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e3/LeafUndersideWithStomata.jpg/500px-LeafUndersideWithStomata.jpg 2x" data-file-width="2848" data-file-height="1899" /></a><figcaption>The underside of a leaf. In this species (<i><a href="/wiki/Tradescantia_zebrina" title="Tradescantia zebrina">Tradescantia zebrina</a></i>) the guard cells of the stomata are green because they contain chlorophyll while the epidermal cells are chlorophyll-free and contain red pigments.</figcaption></figure> <p>In <a href="/wiki/Botany" title="Botany">botany</a>, a <b>stoma</b> (<abbr title="plural">pl.</abbr>: <b>stomata</b>, from <a href="/wiki/Greek_language" title="Greek language">Greek</a> <i>στόμα</i>, "mouth"), also called a <b>stomate</b> (<abbr title="plural">pl.</abbr>: <b>stomates</b>), is a pore found in the <a href="/wiki/Epidermis_(botany)" title="Epidermis (botany)">epidermis</a> of leaves, stems, and other organs, that controls the rate of <a href="/wiki/Gas_exchange" title="Gas exchange">gas exchange</a> between the internal air spaces of the leaf and the atmosphere. The pore is bordered by a pair of specialized <a href="/wiki/Ground_tissue#Parenchyma" title="Ground tissue">parenchyma</a> cells known as <a href="/wiki/Guard_cell" title="Guard cell">guard cells</a> that regulate the size of the stomatal opening. </p><p>The term is usually used collectively to refer to the entire stomatal complex, consisting of the paired guard cells and the pore itself, which is referred to as the stomatal aperture.<sup id="cite_ref-Esau_1-0" class="reference"><a href="#cite_note-Esau-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Air, containing <a href="/wiki/Oxygen" title="Oxygen">oxygen</a>, which is used in <a href="/wiki/Cellular_respiration" title="Cellular respiration">respiration</a>, and <a href="/wiki/Carbon_dioxide" title="Carbon dioxide">carbon dioxide</a>, which is used in <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>, passes through stomata by gaseous <a href="/wiki/Diffusion" title="Diffusion">diffusion</a>. <a href="/wiki/Water_vapour" class="mw-redirect" title="Water vapour">Water vapour</a> diffuses through the stomata into the atmosphere as part of a process called <a href="/wiki/Transpiration" title="Transpiration">transpiration</a>. </p><p>Stomata are present in the <a href="/wiki/Sporophyte" title="Sporophyte">sporophyte</a> generation of the vast majority of <a href="/wiki/Land_plants" class="mw-redirect" title="Land plants">land plants</a>, with the exception of <a href="/wiki/Marchantiophyta" title="Marchantiophyta">liverworts</a>, as well as some <a href="/wiki/Mosses" class="mw-redirect" title="Mosses">mosses</a> and <a href="/wiki/Hornworts" class="mw-redirect" title="Hornworts">hornworts</a>. In vascular plants the number, size and distribution of stomata varies widely. <a href="/wiki/Dicotyledon" title="Dicotyledon">Dicotyledons</a> usually have more stomata on the lower surface of the leaves than the upper surface. <a href="/wiki/Monocotyledon" title="Monocotyledon">Monocotyledons</a> such as <a href="/wiki/Onion" title="Onion">onion</a>, <a href="/wiki/Oat" title="Oat">oat</a> and <a href="/wiki/Maize" title="Maize">maize</a> may have about the same number of stomata on both leaf surfaces.<sup id="cite_ref-Weyers_2-0" class="reference"><a href="#cite_note-Weyers-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 5">: 5 </span></sup> In plants with floating leaves, stomata may be found only on the upper epidermis and submerged leaves may lack stomata entirely. Most tree species have stomata only on the lower leaf surface.<sup id="cite_ref-Willmer-1996_3-0" class="reference"><a href="#cite_note-Willmer-1996-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Leaves with stomata on both the upper and lower leaf surfaces are called <i>amphistomatous</i> leaves; leaves with stomata only on the lower surface are <i>hypostomatous</i>, and leaves with stomata only on the upper surface are epistomatous or <i>hyperstomatous</i>.<sup id="cite_ref-Willmer-1996_3-1" class="reference"><a href="#cite_note-Willmer-1996-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Size varies across species, with end-to-end lengths ranging from 10 to 80 <a href="/wiki/Micrometre" title="Micrometre">μm</a> and width ranging from a few to 50 μm.<sup id="cite_ref-Fricker_4-0" class="reference"><a href="#cite_note-Fricker-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=1" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Stoma_with_Accompanying_Guard_Cells.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Stoma_with_Accompanying_Guard_Cells.jpg/240px-Stoma_with_Accompanying_Guard_Cells.jpg" decoding="async" width="240" height="221" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Stoma_with_Accompanying_Guard_Cells.jpg/360px-Stoma_with_Accompanying_Guard_Cells.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/22/Stoma_with_Accompanying_Guard_Cells.jpg/480px-Stoma_with_Accompanying_Guard_Cells.jpg 2x" data-file-width="512" data-file-height="471" /></a><figcaption>Electron micrograph of a stoma from a <a href="/wiki/Bok_choy" title="Bok choy">bok choy</a> (<i>Brassica chinensis</i>) leaf</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="CO2_gain_and_water_loss">CO<sub>2</sub> gain and water loss</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=2" title="Edit section: CO2 gain and water loss"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Carbon_dioxide" title="Carbon dioxide">Carbon dioxide</a>, a key reactant in <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>, is present in the atmosphere at a concentration of about 400 ppm. Most plants require the stomata to be open during daytime. The air spaces in the <a href="/wiki/Leaf" title="Leaf">leaf</a> are saturated with <a href="/wiki/Water_vapour" class="mw-redirect" title="Water vapour">water vapour</a>, which exits the leaf through the stomata in a process known as <a href="/wiki/Transpiration" title="Transpiration">transpiration</a>. Therefore, plants cannot gain carbon dioxide without simultaneously losing water vapour.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Alternative_approaches">Alternative approaches</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=3" title="Edit section: Alternative approaches"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ordinarily, carbon dioxide is fixed to <a href="/wiki/Ribulose_1,5-bisphosphate" title="Ribulose 1,5-bisphosphate">ribulose 1,5-bisphosphate</a> (RuBP) by the enzyme <a href="/wiki/RuBisCO" title="RuBisCO">RuBisCO</a> in <a href="/wiki/Mesophyll" class="mw-redirect" title="Mesophyll">mesophyll</a> cells exposed directly to the air spaces inside the leaf. This exacerbates the transpiration problem for two reasons: first, RuBisCo has a relatively low affinity for carbon dioxide, and second, it fixes oxygen to RuBP, wasting energy and carbon in a process called <a href="/wiki/Photorespiration" title="Photorespiration">photorespiration</a>. For both of these reasons, RuBisCo needs high carbon dioxide concentrations, which means wide stomatal apertures and, as a consequence, high water loss. </p><p>Narrower stomatal apertures can be used in conjunction with an intermediary molecule with a high carbon dioxide affinity, <a href="/wiki/Phosphoenolpyruvate_carboxylase" title="Phosphoenolpyruvate carboxylase">phosphoenolpyruvate carboxylase</a> (PEPcase). Retrieving the products of carbon fixation from PEPCase is an energy-intensive process, however. As a result, the PEPCase alternative is preferable only where water is limiting but light is plentiful, or where high temperatures increase the solubility of oxygen relative to that of carbon dioxide, magnifying RuBisCo's oxygenation problem. </p> <div class="mw-heading mw-heading3"><h3 id="C.A.M._plants">C.A.M. plants</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=4" title="Edit section: C.A.M. plants"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Differences_in_Stomata_Opening_Throughout_the_Day_for_C3_plants_and_CAM_plants_(1).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/26/Differences_in_Stomata_Opening_Throughout_the_Day_for_C3_plants_and_CAM_plants_%281%29.svg/220px-Differences_in_Stomata_Opening_Throughout_the_Day_for_C3_plants_and_CAM_plants_%281%29.svg.png" decoding="async" width="220" height="224" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/26/Differences_in_Stomata_Opening_Throughout_the_Day_for_C3_plants_and_CAM_plants_%281%29.svg/330px-Differences_in_Stomata_Opening_Throughout_the_Day_for_C3_plants_and_CAM_plants_%281%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/26/Differences_in_Stomata_Opening_Throughout_the_Day_for_C3_plants_and_CAM_plants_%281%29.svg/440px-Differences_in_Stomata_Opening_Throughout_the_Day_for_C3_plants_and_CAM_plants_%281%29.svg.png 2x" data-file-width="512" data-file-height="522" /></a><figcaption>C3 and C4 plants(1) stomata stay open all day and close at night. CAM plants(2) stomata open during the morning and close slightly at noon and then open again in the evening.</figcaption></figure> <p>A group of mostly desert plants called "C.A.M." plants (<a href="/wiki/Crassulacean_acid_metabolism" title="Crassulacean acid metabolism">crassulacean acid metabolism</a>, after the family Crassulaceae, which includes the species in which the CAM process was first discovered) open their stomata at night (when water evaporates more slowly from leaves for a given degree of stomatal opening), use PEPcase to fix carbon dioxide and store the products in large vacuoles. The following day, they close their stomata and release the carbon dioxide fixed the previous night into the presence of RuBisCO. This saturates RuBisCO with carbon dioxide, allowing minimal photorespiration. This approach, however, is severely limited by the capacity to store fixed carbon in the vacuoles, so it is preferable only when water is severely limited. </p> <div class="mw-heading mw-heading3"><h3 id="Opening_and_closing">Opening and closing</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=5" title="Edit section: Opening and closing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Guard_cell" title="Guard cell">Guard cell</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Opening_and_Closing_of_Stoma.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Opening_and_Closing_of_Stoma.svg/500px-Opening_and_Closing_of_Stoma.svg.png" decoding="async" width="420" height="249" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Opening_and_Closing_of_Stoma.svg/960px-Opening_and_Closing_of_Stoma.svg.png 1.5x" data-file-width="1297" data-file-height="769" /></a><figcaption>Opening and closing of stoma</figcaption></figure> <p>However, most plants do not have CAM and must therefore open and close their stomata during the daytime, in response to changing conditions, such as light intensity, humidity, and carbon dioxide concentration. When conditions are conducive to stomatal opening (e.g., high light intensity and high humidity), a <a href="/wiki/Proton_pump" title="Proton pump">proton pump</a> drives <a href="/wiki/Proton" title="Proton">protons</a> (H<sup>+</sup>) from the guard cells. This means that the cells' <a href="/wiki/Electrical_potential" class="mw-redirect" title="Electrical potential">electrical potential</a> becomes increasingly negative. The negative potential opens potassium voltage-gated channels and so an uptake of <a href="/wiki/Potassium" title="Potassium">potassium</a> ions (K<sup>+</sup>) occurs. To maintain this internal negative voltage so that entry of potassium ions does not stop, negative ions balance the influx of potassium. In some cases, chloride ions enter, while in other plants the organic ion <a href="/wiki/Malate" class="mw-redirect" title="Malate">malate</a> is produced in guard cells. This increase in solute concentration lowers the <a href="/wiki/Water_potential" title="Water potential">water potential</a> inside the cell, which results in the diffusion of water into the cell through <a href="/wiki/Osmosis" title="Osmosis">osmosis</a>. This increases the cell's volume and <a href="/wiki/Osmotic_pressure" title="Osmotic pressure">turgor pressure</a>. Then, because of rings of cellulose <a href="/wiki/Microfibrils" class="mw-redirect" title="Microfibrils">microfibrils</a> that prevent the width of the guard cells from swelling, and thus only allow the extra turgor pressure to elongate the guard cells, whose ends are held firmly in place by surrounding <a href="/wiki/Epidermis_(botany)" title="Epidermis (botany)">epidermal</a> cells, the two guard cells lengthen by bowing apart from one another, creating an open pore through which gas can diffuse.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>When the roots begin to sense a water shortage in the soil, <a href="/wiki/Abscisic_acid" title="Abscisic acid">abscisic acid</a> (ABA) is released.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> ABA binds to receptor proteins in the guard cells' plasma membrane and cytosol, which first raises the pH of the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a> of the cells and cause the concentration of free Ca<sup>2+</sup> to increase in the cytosol due to influx from outside the cell and release of Ca<sup>2+</sup> from internal stores such as the endoplasmic reticulum and vacuoles.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> This causes the chloride (Cl<sup>−</sup>) and organic ions to exit the cells. Second, this stops the uptake of any further K<sup>+</sup> into the cells and, subsequently, the loss of K<sup>+</sup>. The loss of these solutes causes an increase in <a href="/wiki/Water_potential" title="Water potential">water potential</a>, which results in the diffusion of water back out of the cell by <a href="/wiki/Osmosis" title="Osmosis">osmosis</a>. This makes the cell <a href="/wiki/Plasmolysed" class="mw-redirect" title="Plasmolysed">plasmolysed</a>, which results in the closing of the stomatal pores. </p><p>Guard cells have more chloroplasts than the other epidermal cells from which guard cells are derived. Their function is controversial.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Inferring_stomatal_behavior_from_gas_exchange">Inferring stomatal behavior from gas exchange</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=6" title="Edit section: Inferring stomatal behavior from gas exchange"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The degree of stomatal resistance can be determined by measuring leaf gas exchange of a leaf. The <a href="/wiki/Transpiration" title="Transpiration">transpiration</a> rate is dependent on the <a href="/wiki/Diffusion" title="Diffusion">diffusion</a> resistance provided by the stomatal pores and also on the <a href="/wiki/Humidity" title="Humidity">humidity</a> gradient between the leaf's internal air spaces and the outside air. Stomatal resistance (or its inverse, <a href="/wiki/Stomatal_conductance" title="Stomatal conductance">stomatal conductance</a>) can therefore be calculated from the transpiration rate and humidity gradient. This allows scientists to investigate how stomata respond to changes in environmental conditions, such as light intensity and concentrations of gases such as water vapor, carbon dioxide, and <a href="/wiki/Ozone" title="Ozone">ozone</a>.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Evaporation (<i>E</i>) can be calculated as<sup id="cite_ref-calculations_12-0" class="reference"><a href="#cite_note-calculations-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle E={\frac {e_{\text{i}}-e_{\text{a}}}{Pr}},}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>E</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <msub> <mi>e</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>i</mtext> </mrow> </msub> <mo>−</mo> <msub> <mi>e</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>a</mtext> </mrow> </msub> </mrow> <mrow> <mi>P</mi> <mi>r</mi> </mrow> </mfrac> </mrow> <mo>,</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle E={\frac {e_{\text{i}}-e_{\text{a}}}{Pr}},}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/3253e105ed8c26c094a1c70e19f5f7cd88e49e88" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.838ex; width:13.108ex; height:5.009ex;" alt="{\displaystyle E={\frac {e_{\text{i}}-e_{\text{a}}}{Pr}},}" /></span></dd></dl> <p>where <i>e</i><sub>i</sub> and <i>e</i><sub>a</sub> are the partial pressures of water in the leaf and in the ambient air respectively, <i>P</i> is atmospheric pressure, and <i>r</i> is stomatal resistance. The inverse of <i>r</i> is conductance to water vapor (<i>g</i>), so the equation can be rearranged to<sup id="cite_ref-calculations_12-1" class="reference"><a href="#cite_note-calculations-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle E=(e_{\text{i}}-e_{\text{a}})g/P}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>E</mi> <mo>=</mo> <mo stretchy="false">(</mo> <msub> <mi>e</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>i</mtext> </mrow> </msub> <mo>−</mo> <msub> <mi>e</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>a</mtext> </mrow> </msub> <mo stretchy="false">)</mo> <mi>g</mi> <mrow class="MJX-TeXAtom-ORD"> <mo>/</mo> </mrow> <mi>P</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle E=(e_{\text{i}}-e_{\text{a}})g/P}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/52b8051f5e4be60c6bf192de4622275027ddcf20" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:17.459ex; height:2.843ex;" alt="{\displaystyle E=(e_{\text{i}}-e_{\text{a}})g/P}" /></span></dd></dl> <p>and solved for <i>g</i>:<sup id="cite_ref-calculations_12-2" class="reference"><a href="#cite_note-calculations-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle g={\frac {EP}{e_{\text{i}}-e_{\text{a}}}}.}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>g</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mi>E</mi> <mi>P</mi> </mrow> <mrow> <msub> <mi>e</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>i</mtext> </mrow> </msub> <mo>−</mo> <msub> <mi>e</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>a</mtext> </mrow> </msub> </mrow> </mfrac> </mrow> <mo>.</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle g={\frac {EP}{e_{\text{i}}-e_{\text{a}}}}.}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/e4668c4f64b9c48bdfd5f75a716934246cf0cdb6" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.171ex; width:12.449ex; height:5.509ex;" alt="{\displaystyle g={\frac {EP}{e_{\text{i}}-e_{\text{a}}}}.}" /></span></dd></dl> <p>Photosynthetic CO<sub>2</sub> assimilation (<i>A</i>) can be calculated from </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle A={\frac {(C_{\text{a}}-C_{\text{i}})g}{1.6P}},}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>A</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mo stretchy="false">(</mo> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>a</mtext> </mrow> </msub> <mo>−</mo> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>i</mtext> </mrow> </msub> <mo stretchy="false">)</mo> <mi>g</mi> </mrow> <mrow> <mn>1.6</mn> <mi>P</mi> </mrow> </mfrac> </mrow> <mo>,</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle A={\frac {(C_{\text{a}}-C_{\text{i}})g}{1.6P}},}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/e6bb4ea1d39760e88658560ca85097e13a9f3afa" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.005ex; width:17.158ex; height:5.843ex;" alt="{\displaystyle A={\frac {(C_{\text{a}}-C_{\text{i}})g}{1.6P}},}" /></span></dd></dl> <p>where <i>C</i><sub>a</sub> and <i>C</i><sub>i</sub> are the atmospheric and sub-stomatal partial pressures of CO<sub>2</sub> respectively<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="what is "1.6"? (March 2023)">clarification needed</span></a></i>]</sup>. The rate of evaporation from a leaf can be determined using a <a href="/wiki/Photosynthesis_system" title="Photosynthesis system">photosynthesis system</a>. These scientific instruments measure the amount of water vapour leaving the leaf and the vapor pressure of the ambient air. Photosynthetic systems may calculate <a href="/wiki/Water_use_efficiency" class="mw-redirect" title="Water use efficiency">water use efficiency</a> (<i>A</i>/<i>E</i>), <i>g</i>, intrinsic water use efficiency (<i>A</i>/<i>g</i>), and <i>C</i><sub>i</sub>. These scientific instruments are commonly used by plant physiologists to measure CO<sub>2</sub> uptake and thus measure photosynthetic rate.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=7" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Tomato_stoma_observed_through_immersion_oil.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/3/3c/Tomato_stoma_observed_through_immersion_oil.gif" decoding="async" width="216" height="235" class="mw-file-element" data-file-width="216" data-file-height="235" /></a><figcaption>Tomato stoma observed through immersion oil</figcaption></figure> <p>There is little evidence of the evolution of stomata in the fossil record, but they had appeared in land plants by the middle of the Silurian period.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> They may have evolved by the modification of <a href="/wiki/Conceptacles" class="mw-redirect" title="Conceptacles">conceptacles</a> from plants' alga-like ancestors.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> However, the evolution of stomata must have happened at the same time as the waxy <a href="/wiki/Plant_cuticle" title="Plant cuticle">cuticle</a> was evolving – these two traits together constituted a major advantage for early terrestrial plants.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2017)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Development">Development</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=8" title="Edit section: Development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are three major epidermal cell types which all ultimately derive from the outermost (L1) tissue layer of the <a href="/wiki/Shoot_apical_meristem" class="mw-redirect" title="Shoot apical meristem">shoot apical meristem</a>, called protodermal cells: <a href="/wiki/Trichome" title="Trichome">trichomes</a>, <a href="/wiki/Pavement_cells" title="Pavement cells">pavement cells</a> and guard cells, all of which are arranged in a non-random fashion. </p><p>An asymmetrical cell division occurs in protodermal cells resulting in one large cell that is fated to become a pavement cell and a smaller cell called a meristemoid that will eventually differentiate into the guard cells that surround a stoma. This meristemoid then divides asymmetrically one to three times before differentiating into a guard mother cell. The guard mother cell then makes one symmetrical division, which forms a pair of guard cells.<sup id="cite_ref-Bergmann-2004_17-0" class="reference"><a href="#cite_note-Bergmann-2004-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Cell division is inhibited in some cells so there is always at least one cell between stomata.<sup id="cite_ref-Pillitteri-2013_18-0" class="reference"><a href="#cite_note-Pillitteri-2013-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>Stomatal patterning is controlled by the interaction of many <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a> components such as <i>EPF</i> (Epidermal Patterning Factor), <i>ERL</i> (ERecta Like) and <i>YODA</i> (a putative <a href="/wiki/MAP_kinase_kinase_kinase" title="MAP kinase kinase kinase">MAP kinase kinase kinase</a>).<sup id="cite_ref-Pillitteri-2013_18-1" class="reference"><a href="#cite_note-Pillitteri-2013-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Mutations in any one of the genes which encode these factors may alter the development of stomata in the epidermis.<sup id="cite_ref-Pillitteri-2013_18-2" class="reference"><a href="#cite_note-Pillitteri-2013-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> For example, a mutation in one gene causes more stomata that are clustered together, hence is called Too Many Mouths (<i>TMM</i>).<sup id="cite_ref-Bergmann-2004_17-1" class="reference"><a href="#cite_note-Bergmann-2004-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Whereas, disruption of the <i>SPCH</i> (SPeecCHless) gene prevents stomatal development all together.<sup id="cite_ref-Pillitteri-2013_18-3" class="reference"><a href="#cite_note-Pillitteri-2013-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup>  Inhibition of stomatal production can occur by the activation of EPF1, which activates TMM/ERL, which together activate YODA. YODA inhibits SPCH, causing SPCH activity to decrease, preventing asymmetrical cell division that initiates stomata formation.<sup id="cite_ref-Pillitteri-2013_18-4" class="reference"><a href="#cite_note-Pillitteri-2013-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Stomatal development is also coordinated by the cellular peptide signal called stomagen, which signals the activation of the SPCH, resulting in increased number of stomata.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>Environmental and hormonal factors can affect stomatal development. Light increases stomatal development in plants; while, plants grown in the dark have a lower amount of stomata. <a href="/wiki/Auxin" title="Auxin">Auxin</a> represses stomatal development by affecting their development at the receptor level like the ERL and TMM receptors. However, a low concentration of auxin allows for equal division of a guard mother cell and increases the chance of producing guard cells.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>Most <a href="/w/index.php?title=Angiosperm_trees&action=edit&redlink=1" class="new" title="Angiosperm trees (page does not exist)">angiosperm trees</a> have stomata only on their lower leaf surface. <a href="/wiki/Populus" title="Populus">Poplars</a> and <a href="/wiki/Willows" class="mw-redirect" title="Willows">willows</a> have them on both surfaces. When leaves develop stomata on both leaf surfaces, the stomata on the lower surface tend to be larger and more numerous, but there can be a great degree of variation in size and frequency about species and genotypes. <a href="/wiki/Fraxinus_americana" title="Fraxinus americana">White ash</a> and <a href="/wiki/White_birch" class="mw-redirect" title="White birch">white birch</a> leaves had fewer stomata but larger in size. On the other hand <a href="/wiki/Sugar_maple" class="mw-redirect" title="Sugar maple">sugar maple</a> and <a href="/wiki/Silver_maple" class="mw-redirect" title="Silver maple">silver maple</a> had small stomata that were more numerous.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=9" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Different classifications of stoma types exist. One that is widely used is based on the types that <a href="/wiki/Julien_Joseph_Vesque" title="Julien Joseph Vesque">Julien Joseph Vesque</a> introduced in 1889, was further developed by Metcalfe and Chalk,<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> and later complemented by other authors. It is based on the size, shape and arrangement of the subsidiary cells that surround the two guard cells.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> They distinguish for <a href="/wiki/Dicotyledon" title="Dicotyledon">dicots</a>: </p> <ul><li><b><style data-mw-deduplicate="TemplateStyles:r1238216509">.mw-parser-output .vanchor>:target~.vanchor-text{background-color:#b1d2ff}@media screen{html.skin-theme-clientpref-night .mw-parser-output .vanchor>:target~.vanchor-text{background-color:#0f4dc9}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .vanchor>:target~.vanchor-text{background-color:#0f4dc9}}</style><span class="vanchor"><span id="actinocytic"></span><span class="vanchor-text">actinocytic</span></span></b> (meaning <i>star-celled</i>) stomata have guard cells that are surrounded by at least five radiating cells forming a star-like circle. This is a rare type that can for instance be found in the family <a href="/wiki/Ebenaceae" title="Ebenaceae">Ebenaceae</a>.</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="anisocytic"></span><span class="vanchor-text">anisocytic</span></span></b> (meaning <i>unequal celled</i>) stomata have guard cells between two larger subsidiary cells and one distinctly smaller one. This type of stomata can be found in more than thirty dicot families, including <a href="/wiki/Brassicaceae" title="Brassicaceae">Brassicaceae</a>, <a href="/wiki/Solanaceae" title="Solanaceae">Solanaceae</a>, and <a href="/wiki/Crassulaceae" title="Crassulaceae">Crassulaceae</a>. It is sometimes called <i>cruciferous type</i>.</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="anomocytic"></span><span class="vanchor-text">anomocytic</span></span></b> (meaning <i>irregular celled</i>) stomata have guard cells that are surrounded by cells that have the same size, shape and arrangement as the rest of the epidermis cells. This type of stomata can be found in more than hundred dicot families such as <a href="/wiki/Apocynaceae" title="Apocynaceae">Apocynaceae</a>, <a href="/wiki/Boraginaceae" title="Boraginaceae">Boraginaceae</a>, <a href="/wiki/Chenopodiaceae" class="mw-redirect" title="Chenopodiaceae">Chenopodiaceae</a>, and <a href="/wiki/Cucurbitaceae" title="Cucurbitaceae">Cucurbitaceae</a>. It is sometimes called <i>ranunculaceous type</i>.</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="diacytic"></span><span class="vanchor-text">diacytic</span></span></b> (meaning <i>cross-celled</i>) stomata have guard cells surrounded by two subsidiary cells, that each encircle one end of the opening and contact each other opposite to the middle of the opening. This type of stomata can be found in more than ten dicot families such as <a href="/wiki/Caryophyllaceae" title="Caryophyllaceae">Caryophyllaceae</a> and <a href="/wiki/Acanthaceae" title="Acanthaceae">Acanthaceae</a>. It is sometimes called <i>caryophyllaceous type</i>.</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="hemiparacytic"></span><span class="vanchor-text">hemiparacytic</span></span></b> stomata are bordered by just one subsidiary cell that differs from the surrounding epidermis cells, its length parallel to the stoma opening. This type occurs for instance in the <a href="/wiki/Molluginaceae" title="Molluginaceae">Molluginaceae</a> and <a href="/wiki/Aizoaceae" title="Aizoaceae">Aizoaceae</a>.</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="paracytic"></span><span class="vanchor-text">paracytic</span></span></b> (meaning <i>parallel celled</i>) stomata have one or more subsidiary cells parallel to the opening between the guard cells. These subsidiary cells may reach beyond the guard cells or not. This type of stomata can be found in more than hundred dicot families such as <a href="/wiki/Rubiaceae" title="Rubiaceae">Rubiaceae</a>, <a href="/wiki/Convolvulaceae" title="Convolvulaceae">Convolvulaceae</a> and <a href="/wiki/Fabaceae" title="Fabaceae">Fabaceae</a>. It is sometimes called <i>rubiaceous type</i>.</li></ul> <p>In <a href="/wiki/Monocotyledon" title="Monocotyledon">monocots</a>, several different types of stomata occur such as: </p> <ul><li><b>gramineous or graminoid</b> (meaning <i>grass-like</i>) stomata have two guard cells surrounded by two lens-shaped subsidiary cells. The guard cells are narrower in the middle and bulbous on each end. This middle section is strongly thickened. The axis of the subsidiary cells are parallel stoma opening. This type can be found in monocot families including <a href="/wiki/Poaceae" title="Poaceae">Poaceae</a> and <a href="/wiki/Cyperaceae" title="Cyperaceae">Cyperaceae</a>.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="hexacytic"></span><span class="vanchor-text">hexacytic</span></span></b> (meaning <i>six-celled</i>) stomata have six subsidiary cells around both guard cells, one at either end of the opening of the stoma, one adjoining each guard cell, and one between that last subsidiary cell and the standard epidermis cells. This type can be found in some monocot families.</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="tetracytic"></span><span class="vanchor-text">tetracytic</span></span></b> (meaning <i>four-celled</i>) stomata have four subsidiary cells, one on either end of the opening, and one next to each guard cell. This type occurs in many monocot families, but also can be found in some dicots, such as <i><a href="/wiki/Tilia" title="Tilia">Tilia</a></i> and several <a href="/wiki/Asclepiadaceae" class="mw-redirect" title="Asclepiadaceae">Asclepiadaceae</a>.</li></ul> <p>In <a href="/wiki/Fern" title="Fern">ferns</a>, four different types are distinguished: </p> <ul><li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="hypocytic"></span><span class="vanchor-text">hypocytic</span></span></b> stomata have two guard cells in one layer with only ordinary epidermis cells, but with two subsidiary cells on the outer surface of the epidermis, arranged parallel to the guard cells, with a pore between them, overlying the stoma opening.</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="pericytic"></span><span class="vanchor-text">pericytic</span></span></b> stomata have two guard cells that are entirely encircled by one continuous subsidiary cell (like a donut).</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="desmocytic"></span><span class="vanchor-text">desmocytic</span></span></b> stomata have two guard cells that are entirely encircled by one subsidiary cell that has not merged its ends (like a sausage).</li> <li><b><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509" /><span class="vanchor"><span id="polocytic"></span><span class="vanchor-text">polocytic</span></span></b> stomata have two guard cells that are largely encircled by one subsidiary cell, but also contact ordinary epidermis cells (like a U or horseshoe).</li></ul> <p>A catalogue of leaf epidermis prints showing stomata from a wide range of species can be found in Wikimedia commons <a class="external free" href="https://commons.wikimedia.org/wiki/Category:Leaf_epidermis_and_stomata_prints">https://commons.wikimedia.org/wiki/Category:Leaf_epidermis_and_stomata_prints</a> </p> <div class="mw-heading mw-heading2"><h2 id="Stomatal_crypts">Stomatal crypts</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=10" title="Edit section: Stomatal crypts"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stomatal crypts are sunken areas of the leaf epidermis which form a chamber-like structure that contains one or more stomata and sometimes trichomes or accumulations of <a href="/wiki/Epicuticular_wax" title="Epicuticular wax">wax</a>. Stomatal crypts can be an adaption to drought and dry climate conditions when the stomatal crypts are very pronounced. However, dry climates are not the only places where they can be found. The following plants are examples of species with stomatal crypts or antechambers: <a href="/wiki/Nerium" title="Nerium"><i>Nerium oleander</i></a>, conifers, <i>Hakea</i><sup id="cite_ref-Jordan-2008_26-0" class="reference"><a href="#cite_note-Jordan-2008-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> and <i><a href="/wiki/Drimys_winteri" title="Drimys winteri">Drimys winteri</a></i> which is a species of plant found in the <a href="/wiki/Cloud_forest" title="Cloud forest">cloud forest</a>.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Stomata_as_pathogenic_pathways">Stomata as pathogenic pathways</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=11" title="Edit section: Stomata as pathogenic pathways"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stomata are holes in the leaf by which pathogens can enter unchallenged. However, stomata can sense the presence of some, if not all, pathogens.<sup id="cite_ref-Underwood2006_28-0" class="reference"><a href="#cite_note-Underwood2006-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> However, pathogenic bacteria applied to <a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana"><i>Arabidopsis</i></a> plant leaves can release the chemical <a href="/wiki/Coronatine" title="Coronatine">coronatine</a>, which induce the stomata to reopen. <sup id="cite_ref-SchulzeLefert2006_29-0" class="reference"><a href="#cite_note-SchulzeLefert2006-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Stomata_and_climate_change">Stomata and climate change</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=12" title="Edit section: Stomata and climate change"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Response_of_stomata_to_environmental_factors">Response of stomata to environmental factors</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=13" title="Edit section: Response of stomata to environmental factors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a>, plant water transport (<a href="/wiki/Xylem" title="Xylem">xylem</a>) and gas exchange are regulated by stomatal function which is important in the functioning of plants.<sup id="cite_ref-Rico-2013_30-0" class="reference"><a href="#cite_note-Rico-2013-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>Stomata are responsive to light with <a href="/wiki/Visible_spectrum" title="Visible spectrum">blue light</a> being almost 10 times as effective as <a href="/wiki/Red" title="Red">red light</a> in causing stomatal response. Research suggests this is because the light response of stomata to blue light is independent of other leaf components like <a href="/wiki/Chlorophyll" title="Chlorophyll">chlorophyll</a>. Guard cell <a href="/wiki/Protoplasts" class="mw-redirect" title="Protoplasts">protoplasts</a> swell under blue light provided there is sufficient availability of <a href="/wiki/Potassium" title="Potassium">potassium</a>.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Multiple studies have found support that increasing potassium concentrations may increase stomatal opening in the mornings, before the photosynthesis process starts, but that later in the day <a href="/wiki/Sucrose" title="Sucrose">sucrose</a> plays a larger role in regulating stomatal opening.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Zeaxanthin" title="Zeaxanthin">Zeaxanthin</a> in guard cells acts as a blue light photoreceptor which mediates the stomatal opening.<sup id="cite_ref-KochharKaur2020_33-0" class="reference"><a href="#cite_note-KochharKaur2020-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> The effect of blue light on guard cells is reversed by green light, which isomerizes zeaxanthin.<sup id="cite_ref-KochharKaur2020_33-1" class="reference"><a href="#cite_note-KochharKaur2020-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Stomatal_density" class="mw-redirect" title="Stomatal density">Stomatal density</a> and aperture (length of stomata) varies under a number of environmental factors such as atmospheric CO<sub>2</sub> concentration, light intensity, air temperature and photoperiod (daytime duration). <sup id="cite_ref-Buckley-2013_34-0" class="reference"><a href="#cite_note-Buckley-2013-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Rogiers-2011_35-0" class="reference"><a href="#cite_note-Rogiers-2011-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>Decreasing stomatal density is one way plants have responded to the increase in concentration of atmospheric CO<sub>2</sub> ([CO<sub>2</sub>]<sub>atm</sub>).<sup id="cite_ref-Ceccarelli-2010_36-0" class="reference"><a href="#cite_note-Ceccarelli-2010-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Although changes in [CO<sub>2</sub>]<sub>atm</sub> response is the least understood mechanistically, this stomatal response has begun to plateau where it is soon expected to impact <a href="/wiki/Transpiration" title="Transpiration">transpiration</a> and photosynthesis processes in plants.<sup id="cite_ref-Rico-2013_30-1" class="reference"><a href="#cite_note-Rico-2013-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p><p>Drought inhibits stomatal opening, but research on soybeans suggests moderate drought does not have a significant effect on stomatal closure of its leaves. There are different mechanisms of stomatal closure. Low humidity stresses guard cells causing <a href="/wiki/Turgor" class="mw-redirect" title="Turgor">turgor</a> loss, termed hydropassive closure. Hydroactive closure is contrasted as the whole leaf affected by drought stress, believed to be most likely triggered by <a href="/wiki/Abscisic_acid" title="Abscisic acid">abscisic acid</a>.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Future_adaptations_during_climate_change">Future adaptations during climate change</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=14" title="Edit section: Future adaptations during climate change"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>It is expected that [CO<sub>2</sub>]<sub>atm</sub> will reach 500–1000 ppm by 2100.<sup id="cite_ref-Rico-2013_30-2" class="reference"><a href="#cite_note-Rico-2013-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> 96% of the past 400,000 years experienced below 280 ppm CO<sub>2</sub>. From this figure, it is highly probable that <a href="/wiki/Genotype" title="Genotype">genotypes</a> of today’s plants have diverged from their pre-industrial relatives.<sup id="cite_ref-Rico-2013_30-3" class="reference"><a href="#cite_note-Rico-2013-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>The gene <i>HIC</i> (high carbon dioxide) encodes a negative regulator for the development of stomata in plants.<sup id="cite_ref-Gray-2000_39-0" class="reference"><a href="#cite_note-Gray-2000-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Research into the <i>HIC</i> gene using<i> <a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis thaliana</a></i> found no increase of stomatal development in the dominant <a href="/wiki/Allele" title="Allele">allele</a>, but in the ‘wild type’ <a href="/wiki/Recessive_allele" class="mw-redirect" title="Recessive allele">recessive allele</a> showed a large increase, both in response to rising CO<sub>2</sub> levels in the atmosphere.<sup id="cite_ref-Gray-2000_39-1" class="reference"><a href="#cite_note-Gray-2000-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> These studies imply the plants response to changing CO<sub>2</sub> levels is largely controlled by genetics. </p> <div class="mw-heading mw-heading3"><h3 id="Agricultural_implications">Agricultural implications</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=15" title="Edit section: Agricultural implications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The CO<sub>2 </sub>fertiliser effect has been greatly overestimated during <a href="/wiki/Free-air_concentration_enrichment" title="Free-air concentration enrichment">Free-Air Carbon dioxide Enrichment</a> (FACE) experiments where results show increased CO<sub>2</sub> levels in the atmosphere enhances photosynthesis, reduce transpiration, and increase <a href="/wiki/Water-use_efficiency" title="Water-use efficiency">water use efficiency</a> (WUE).<sup id="cite_ref-Ceccarelli-2010_36-1" class="reference"><a href="#cite_note-Ceccarelli-2010-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Increased <a href="/wiki/Biomass" title="Biomass">biomass</a> is one of the effects with simulations from experiments predicting a 5–20% increase in crop yields at 550 ppm of CO<sub>2</sub>.<sup id="cite_ref-Tubiello-2007_40-0" class="reference"><a href="#cite_note-Tubiello-2007-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> Rates of leaf photosynthesis were shown to increase by 30–50% in <a href="/wiki/C3_carbon_fixation" title="C3 carbon fixation">C3</a> plants, and 10–25% in <a href="/wiki/C4_carbon_fixation" title="C4 carbon fixation">C4</a> under doubled CO<sub>2</sub> levels.<sup id="cite_ref-Tubiello-2007_40-1" class="reference"><a href="#cite_note-Tubiello-2007-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> The existence of a <a href="/wiki/Feedback_mechanism" class="mw-redirect" title="Feedback mechanism">feedback mechanism</a> results a <a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">phenotypic plasticity</a> in response to [CO<sub>2</sub>]<sub>atm</sub> that may have been an adaptive trait in the evolution of plant respiration and function.<sup id="cite_ref-Rico-2013_30-4" class="reference"><a href="#cite_note-Rico-2013-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Rogiers-2011_35-1" class="reference"><a href="#cite_note-Rogiers-2011-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>Predicting how stomata perform during adaptation is useful for understanding the productivity of plant systems for both natural and <a href="/wiki/Agricultural_systems" class="mw-redirect" title="Agricultural systems">agricultural systems</a>.<sup id="cite_ref-Buckley-2013_34-1" class="reference"><a href="#cite_note-Buckley-2013-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Plant breeders and farmers are beginning to work together using evolutionary and participatory plant breeding to find the best suited species such as heat and drought resistant crop varieties that could naturally evolve to the change in the face of food security challenges.<sup id="cite_ref-Ceccarelli-2010_36-2" class="reference"><a href="#cite_note-Ceccarelli-2010-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stoma&action=edit&section=16" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-Esau-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Esau_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFEsau1977" class="citation book cs1">Esau, K. (1977). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/anatomyofseedpla00esau"><i>Anatomy of Seed Plants</i></a></span>. Wiley and Sons. p. <a rel="nofollow" class="external text" href="https://archive.org/details/anatomyofseedpla00esau/page/88">88</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-471-24520-9" title="Special:BookSources/978-0-471-24520-9"><bdi>978-0-471-24520-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Anatomy+of+Seed+Plants&rft.pages=88&rft.pub=Wiley+and+Sons&rft.date=1977&rft.isbn=978-0-471-24520-9&rft.aulast=Esau&rft.aufirst=K.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fanatomyofseedpla00esau&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStoma" class="Z3988"></span></span> </li> <li id="cite_note-Weyers-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-Weyers_2-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFWeyersMeidner1990" class="citation book cs1">Weyers, J. D. B.; Meidner, H. (1990). <i>Methods in stomatal research</i>. Longman Group UK Ltd. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0582034839" title="Special:BookSources/978-0582034839"><bdi>978-0582034839</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Methods+in+stomatal+research&rft.pub=Longman+Group+UK+Ltd.&rft.date=1990&rft.isbn=978-0582034839&rft.aulast=Weyers&rft.aufirst=J.+D.+B.&rft.au=Meidner%2C+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStoma" class="Z3988"></span></span> </li> <li id="cite_note-Willmer-1996-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-Willmer-1996_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Willmer-1996_3-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFWillmerFricker1996" class="citation book cs1">Willmer, Colin; Fricker, Mark (1996). <i>Stomata</i>. Springer. p. 16. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-94-011-0579-8">10.1007/978-94-011-0579-8</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-94-010-4256-7" title="Special:BookSources/978-94-010-4256-7"><bdi>978-94-010-4256-7</bdi></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:224833888">224833888</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Stomata&rft.pages=16&rft.pub=Springer&rft.date=1996&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A224833888%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1007%2F978-94-011-0579-8&rft.isbn=978-94-010-4256-7&rft.aulast=Willmer&rft.aufirst=Colin&rft.au=Fricker%2C+Mark&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStoma" class="Z3988"></span></span> </li> <li id="cite_note-Fricker-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-Fricker_4-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFFrickerWillmer2012" class="citation book cs1">Fricker, M.; Willmer, C. (2012). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=9sjoCAAAQBAJ&pg=PA18"><i>Stomata</i></a>. Springer Netherlands. p. 18. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-94-011-0579-8" title="Special:BookSources/978-94-011-0579-8"><bdi>978-94-011-0579-8</bdi></a><span class="reference-accessdate">. Retrieved <span class="nowrap">15 June</span> 2016</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Stomata&rft.pages=18&rft.pub=Springer+Netherlands&rft.date=2012&rft.isbn=978-94-011-0579-8&rft.aulast=Fricker&rft.aufirst=M.&rft.au=Willmer%2C+C.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D9sjoCAAAQBAJ%26pg%3DPA18&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStoma" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text">Debbie Swarthout and C.Michael Hogan. 2010. <a rel="nofollow" class="external text" href="http://www.eoearth.org/article/Stomata"><i>Stomata</i>. Encyclopedia of Earth</a>. 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=The+HIC+signalling+pathway+links+CO%3Csub%3E2%3C%2Fsub%3E+perception+to+stomatal+development&rft.volume=408&rft.issue=6813&rft.pages=%3Cspan+class%3D%22nowrap%22%3E713-%3C%2Fspan%3E716&rft.date=2000&rft_id=info%3Adoi%2F10.1038%2F35047071&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A83843467%23id-name%3DS2CID&rft_id=info%3Apmid%2F11130071&rft_id=info%3Abibcode%2F2000Natur.408..713G&rft.aulast=Gray&rft.aufirst=J&rft.au=Holroyd%2C+G&rft.au=van+der+Lee%2C+F&rft.au=Bahrami%2C+A&rft.au=Sijmons%2C+P&rft.au=Woodward%2C+F&rft.au=Schuch%2C+W&rft.au=Hetherington%2C+A&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStoma" class="Z3988"></span></span> </li> <li id="cite_note-Tubiello-2007-40"><span class="mw-cite-backlink">^ <a href="#cite_ref-Tubiello-2007_40-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Tubiello-2007_40-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFTubielloSoussanaHowden2007" class="citation journal cs1">Tubiello, FN; Soussana, J-F; Howden, SM (2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2148358">"Crop and pasture response to climate change"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>104</b> (50): <span class="nowrap">19686–</span>19690. <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/2007PNAS..10419686T">2007PNAS..10419686T</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.0701728104">10.1073/pnas.0701728104</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2148358">2148358</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18077401">18077401</a>.</cite><span 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.navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Botany" title="Template:Botany"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Botany" title="Template talk:Botany"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Botany" title="Special:EditPage/Template:Botany"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Botany862" style="font-size:114%;margin:0 4em"><a href="/wiki/Botany" title="Botany">Botany</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2" style="background: #C3EEC3;"><div> <ul><li><a href="/wiki/History_of_botany" title="History of botany">History</a></li> <li><a href="/wiki/Outline_of_botany" title="Outline of botany">Outline</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background: #C3EEC3;;width:1%"><a href="/wiki/Branches_of_botany" title="Branches of botany">Subdisciplines</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/Archaeobotany" class="mw-redirect" title="Archaeobotany">Archaeobotany</a></li> <li><a href="/wiki/Astrobotany" title="Astrobotany">Astrobotany</a></li> <li><a href="/wiki/Bryology" 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_intelligence" title="Plant intelligence">Plant intelligence</a></li> <li><a href="/wiki/Plant_pathology" title="Plant pathology">Plant pathology</a></li> <li><a href="/wiki/Plant_physiology" title="Plant physiology">Plant physiology</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><br /><span class="nobold">(<a href="/wiki/Glossary_of_plant_morphology" title="Glossary of plant morphology">glossary</a>)</span></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><br />(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 class="mw-selflink selflink">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><br />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<br />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 href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">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" 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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><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" 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control">Authority control databases</a>: National <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q216635#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" 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