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Self-fertile </div> </a> <ul id="toc-Self-fertile-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Physiology" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Physiology"> <div class="vector-toc-text"> 3 Physiology </div> </a> <ul id="toc-Physiology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genetics" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Genetics"> <div class="vector-toc-text"> 4 Genetics </div> </a> <ul id="toc-Genetics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Research" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Research"> <div class="vector-toc-text"> 5 Research </div> </a> <ul id="toc-Research-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Origin_and_diversity" class="vector-toc-list-item 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</li> <li id="toc-Reducing_inoculum" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Reducing_inoculum"> <div class="vector-toc-text"> 7.2 Reducing inoculum </div> </a> <ul id="toc-Reducing_inoculum-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Environmental_conditions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Environmental_conditions"> <div class="vector-toc-text"> 7.3 Environmental conditions </div> </a> <ul id="toc-Environmental_conditions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Use_of_fungicides" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Use_of_fungicides"> <div class="vector-toc-text"> 7.4 Use of fungicides </div> </a> <ul id="toc-Use_of_fungicides-sublist" class="vector-toc-list"> <li id="toc-In_organic_production" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#In_organic_production"> <div class="vector-toc-text"> 7.4.1 In organic production </div> </a> <ul id="toc-In_organic_production-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Control_of_tuber_blight" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Control_of_tuber_blight"> <div class="vector-toc-text"> 7.5 Control of tuber blight </div> </a> <ul id="toc-Control_of_tuber_blight-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Historical_impact" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Historical_impact"> <div class="vector-toc-text"> 8 Historical impact </div> </a> <ul id="toc-Historical_impact-sublist" class="vector-toc-list"> </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"> 9 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> 10 Further reading </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 11 External links </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" > Toggle the table of contents </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading">Phytophthora infestans</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 46 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-46" aria-hidden="true" > 46 languages </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%84%D9%81%D8%AD%D8%A9_%D9%85%D8%AA%D8%A3%D8%AE%D8%B1%D8%A9" title="لفحة متأخرة – Arabic" lang="ar" hreflang="ar" data-title="لفحة متأخرة" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Catalan" lang="ca" hreflang="ca" data-title="Phytophthora infestans" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-ceb mw-list-item"><a href="https://ceb.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Cebuano" lang="ceb" hreflang="ceb" data-title="Phytophthora infestans" data-language-autonym="Cebuano" data-language-local-name="Cebuano" class="interlanguage-link-target">Cebuano</a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Clwyf_tatws_hwyr" title="Clwyf tatws hwyr – Welsh" lang="cy" hreflang="cy" data-title="Clwyf tatws hwyr" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target">Cymraeg</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Kartoffelskimmel" title="Kartoffelskimmel – Danish" lang="da" hreflang="da" data-title="Kartoffelskimmel" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – German" lang="de" hreflang="de" data-title="Phytophthora infestans" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Kartuli-lehem%C3%A4danik" title="Kartuli-lehemädanik – Estonian" lang="et" hreflang="et" data-title="Kartuli-lehemädanik" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%A0%CE%B5%CF%81%CE%BF%CE%BD%CF%8C%CF%83%CF%80%CE%BF%CF%81%CE%BF%CF%82_%CF%84%CE%B7%CF%82_%CF%80%CE%B1%CF%84%CE%AC%CF%84%CE%B1%CF%82" title="Περονόσπορος της πατάτας – Greek" lang="el" hreflang="el" data-title="Περονόσπορος της πατάτας" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Spanish" lang="es" hreflang="es" data-title="Phytophthora infestans" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Basque" lang="eu" hreflang="eu" data-title="Phytophthora infestans" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D9%88%D8%AE%D8%AA%DA%AF%DB%8C_%D8%B3%DB%8C%D8%A8%E2%80%8C%D8%B2%D9%85%DB%8C%D9%86%DB%8C" title="سوختگی سیب‌زمینی – Persian" lang="fa" hreflang="fa" data-title="سوختگی سیب‌زمینی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Mildiou_de_la_pomme_de_terre" title="Mildiou de la pomme de terre – French" lang="fr" hreflang="fr" data-title="Mildiou de la pomme de terre" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/D%C3%BAchan_pr%C3%A1ta%C3%AD" title="Dúchan prátaí – Irish" lang="ga" hreflang="ga" data-title="Dúchan prátaí" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Galician" lang="gl" hreflang="gl" data-title="Phytophthora infestans" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EA%B0%90%EC%9E%90%EC%97%AD%EB%B3%91%EA%B7%A0" title="감자역병균 – Korean" lang="ko" hreflang="ko" data-title="감자역병균" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%96%D5%AB%D5%BF%D5%B8%D6%86%D5%BF%D5%B8%D6%80%D5%B8%D5%A6" title="Ֆիտոֆտորոզ – Armenian" lang="hy" hreflang="hy" data-title="Ֆիտոֆտորոզ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Indonesian" lang="id" hreflang="id" data-title="Phytophthora infestans" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Kart%C3%B6flumygla" title="Kartöflumygla – Icelandic" lang="is" hreflang="is" data-title="Kartöflumygla" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target">Íslenska</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Italian" lang="it" hreflang="it" data-title="Phytophthora infestans" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%9B%D7%99%D7%9E%D7%A9%D7%95%D7%9F_%D7%AA%D7%A4%D7%95%D7%97%D7%99_%D7%94%D7%90%D7%93%D7%9E%D7%94" title="כימשון תפוחי האדמה – Hebrew" lang="he" hreflang="he" data-title="כימשון תפוחי האדמה" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%A4%D0%B8%D1%82%D0%BE%D1%84%D1%82%D0%BE%D1%80%D0%BE%D0%B7" title="Фитофтороз – Kazakh" lang="kk" hreflang="kk" data-title="Фитофтороз" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%A4%D0%B8%D1%82%D0%BE%D1%84%D1%82%D0%BE%D1%80%D0%BE%D0%B7" title="Фитофтороз – Kyrgyz" lang="ky" hreflang="ky" data-title="Фитофтороз" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target">Кыргызча</a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Fitoftoroz%C4%97" title="Fitoftorozė – Lithuanian" lang="lt" hreflang="lt" data-title="Fitoftorozė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D9%84%D9%81%D8%AD%D9%87_%D9%85%D8%AA%D8%A7%D8%AE%D8%B1%D9%87" title="لفحه متاخره – Egyptian Arabic" lang="arz" hreflang="arz" data-title="لفحه متاخره" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target">مصرى</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Malay" lang="ms" hreflang="ms" data-title="Phytophthora infestans" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Dutch" lang="nl" hreflang="nl" data-title="Phytophthora infestans" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ne mw-list-item"><a href="https://ne.wikipedia.org/wiki/%E0%A4%AA%E0%A4%9B%E0%A5%8C%E0%A4%9F%E0%A5%87_%E0%A4%A1%E0%A4%A2%E0%A5%81%E0%A4%B5%E0%A4%BE" title="पछौटे डढुवा – Nepali" lang="ne" hreflang="ne" data-title="पछौटे डढुवा" data-language-autonym="नेपाली" data-language-local-name="Nepali" class="interlanguage-link-target">नेपाली</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Potett%C3%B8rr%C3%A5te" title="Potettørråte – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Potettørråte" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Mildi%C3%B3_de_la_trufa" title="Mildió de la trufa – Occitan" lang="oc" hreflang="oc" data-title="Mildió de la trufa" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target">Occitan</a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Fitoftoroz" title="Fitoftoroz – Uzbek" lang="uz" hreflang="uz" data-title="Fitoftoroz" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target">Oʻzbekcha / ўзбекча</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Polish" lang="pl" hreflang="pl" data-title="Phytophthora infestans" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Portuguese" lang="pt" hreflang="pt" data-title="Phytophthora infestans" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Phytophthora_infestans" title="Phytophthora infestans – Romanian" lang="ro" hreflang="ro" data-title="Phytophthora infestans" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-qu mw-list-item"><a href="https://qu.wikipedia.org/wiki/Papa_lancha" title="Papa lancha – Quechua" lang="qu" hreflang="qu" data-title="Papa lancha" data-language-autonym="Runa Simi" data-language-local-name="Quechua" class="interlanguage-link-target">Runa Simi</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A4%D0%B8%D1%82%D0%BE%D1%84%D1%82%D0%BE%D1%80%D0%BE%D0%B7" title="Фитофтороз – Russian" lang="ru" hreflang="ru" data-title="Фитофтороз" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-stq mw-list-item"><a href="https://stq.wikipedia.org/wiki/Tuffelke-Fuuler%C3%A4i" title="Tuffelke-Fuuleräi – Saterland Frisian" lang="stq" hreflang="stq" data-title="Tuffelke-Fuuleräi" data-language-autonym="Seeltersk" data-language-local-name="Saterland Frisian" class="interlanguage-link-target">Seeltersk</a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Vrugu_i_patates" title="Vrugu i patates – Albanian" lang="sq" hreflang="sq" data-title="Vrugu i patates" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target">Shqip</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Potato_blight" title="Potato blight – Simple English" lang="en-simple" hreflang="en-simple" data-title="Potato blight" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Krompirjeva_plesen" title="Krompirjeva plesen – Slovenian" lang="sl" hreflang="sl" data-title="Krompirjeva plesen" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Perunarutto" title="Perunarutto – Finnish" lang="fi" hreflang="fi" data-title="Perunarutto" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Potatisbladm%C3%B6gel" title="Potatisbladmögel – Swedish" lang="sv" hreflang="sv" data-title="Potatisbladmögel" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a 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width: 200px; font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(200,250,80)">Phytophthora infestans </th></tr> <tr> <td colspan="2" style="text-align: center"><a href="/wiki/File:Late_blight_on_potato_leaf_2.jpg" class="mw-file-description"><img alt="Symptom of late blight on the underside of a potato leaf" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Late_blight_on_potato_leaf_2.jpg/220px-Late_blight_on_potato_leaf_2.jpg" decoding="async" width="220" height="138" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Late_blight_on_potato_leaf_2.jpg/330px-Late_blight_on_potato_leaf_2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Late_blight_on_potato_leaf_2.jpg/440px-Late_blight_on_potato_leaf_2.jpg 2x" data-file-width="3072" data-file-height="1928" /></a> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%">Symptom of late blight on the underside of a <a href="/wiki/Potato" title="Potato">potato</a> leaf </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(200,250,80)"><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <a href="/wiki/Template:Taxonomy/Phytophthora" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a> </th></tr> <tr> <td>Domain: </td> <td><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> </td></tr> <tr> <td>Clade: </td> <td><a href="/wiki/Diaphoretickes" title="Diaphoretickes">Diaphoretickes</a> </td></tr> <tr> <td>Clade: </td> <td><a href="/wiki/SAR_supergroup" title="SAR supergroup">SAR</a> </td></tr> <tr> <td>Clade: </td> <td><a href="/wiki/Stramenopile" title="Stramenopile">Stramenopiles</a> </td></tr> <tr> <td>Phylum: </td> <td><a href="/wiki/Oomycete" title="Oomycete">Oomycota</a> </td></tr> <tr> <td>Order: </td> <td><a href="/wiki/Peronosporales" title="Peronosporales">Peronosporales</a> </td></tr> <tr> <td>Family: </td> <td><a href="/wiki/Peronosporaceae" title="Peronosporaceae">Peronosporaceae</a> </td></tr> <tr> <td>Genus: </td> <td><a href="/wiki/Phytophthora" title="Phytophthora">Phytophthora</a> </td></tr> <tr> <td>Species: </td> <td><div style="display:inline" class="species">P. infestans</div> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(200,250,80)"><a href="/wiki/Binomial_nomenclature" title="Binomial nomenclature">Binomial name</a> </th></tr> <tr> <td colspan="2" style="text-align: center">Phytophthora infestans <div style="font-size: 85%;">(<a href="/wiki/Jean_Pierre_Fran%C3%A7ois_Camille_Montagne" class="mw-redirect" title="Jean Pierre François Camille Montagne">Mont.</a>) <a href="/wiki/Heinrich_Anton_de_Bary" title="Heinrich Anton de Bary">de Bary</a></div> </td></tr> </tbody></table><style data-mw-deduplicate="TemplateStyles:r1238732961">@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .infobox.biota img{background:transparent}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .infobox.biota img{background:white}}.mw-parser-output .infobox.biota .taxobox-edit-taxonomy img{background:transparent!important}body.skin-vector .mw-parser-output table.biota.infobox{margin-top:0.5em}body.skin--responsive .mw-parser-output table.biota.infobox tr.taxonrow td{padding:2px 10px}</style> Phytophthora infestans is an <a href="/wiki/Oomycete" title="Oomycete">oomycete</a> or <a href="/wiki/Oomycete" title="Oomycete">water mold</a>, a fungus-like <a href="/wiki/Microorganism" title="Microorganism">microorganism</a> that causes the serious <a href="/wiki/Potato" title="Potato">potato</a> and <a href="/wiki/Tomato" title="Tomato">tomato</a> disease known as late blight or potato blight. <a href="/wiki/Early_blight" class="mw-redirect" title="Early blight">Early blight</a>, caused by <a href="/wiki/Alternaria_solani" title="Alternaria solani">Alternaria solani</a>, is also often called "potato blight". Late blight was a major culprit in the <a href="/wiki/European_Potato_Failure" class="mw-redirect" title="European Potato Failure">1840s European</a>, the <a href="/wiki/Great_Famine_(Ireland)" title="Great Famine (Ireland)">1845–1852 Irish</a>, and the <a href="/wiki/Highland_Potato_Famine" title="Highland Potato Famine">1846 Highland</a> potato <a href="/wiki/Famine" title="Famine">famines</a>. The organism can also infect some other members of the <a href="/wiki/Solanaceae" title="Solanaceae">Solanaceae</a>.<a href="#cite_note-bbc_news-1">[1]</a><a href="#cite_note-PlDis2011-2">[2]</a><a href="#cite_note-Wiley2013-3">[3]</a> The pathogen is favored by moist, cool environments: sporulation is optimal at 12–18 °C (54–64 °F) in water-saturated or nearly saturated environments, and <a href="/wiki/Zoospore" title="Zoospore">zoospore</a> production is favored at temperatures below 15 °C (59 °F). Lesion growth rates are typically optimal at a slightly warmer temperature range of 20 to 24 °C (68 to 75 °F).<a href="#cite_note-4">[4]</a> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Etymology">Etymology</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=1" title="Edit section: Etymology">edit</a>]</div> The genus name Phytophthora comes from the Greek φυτό (phyto), meaning "plant" – plus the Greek φθορά (phthora), meaning "decay, ruin, perish". The species name infestans is the present participle of the Latin verb infestare, meaning "attacking, destroying", from which the word "to infest" is derived. The name Phytophthora infestans was coined in 1876 by the German mycologist <a href="/wiki/Heinrich_Anton_de_Bary" title="Heinrich Anton de Bary">Heinrich Anton de Bary</a> (1831–1888).<a href="#cite_note-5">[5]</a><a href="#cite_note-6">[6]</a> <div class="mw-heading mw-heading2"><h2 id="Life_cycle,_signs_and_symptoms">Life cycle, signs and symptoms</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=2" title="Edit section: Life cycle, signs and symptoms">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Frameless"><a href="/wiki/File:Tomato_with_Phytophthora_infestans_(late_blight).jpg" class="mw-file-description"><img alt="Infected ripe tomato" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Tomato_with_Phytophthora_infestans_%28late_blight%29.jpg/220px-Tomato_with_Phytophthora_infestans_%28late_blight%29.jpg" decoding="async" width="220" height="111" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Tomato_with_Phytophthora_infestans_%28late_blight%29.jpg/330px-Tomato_with_Phytophthora_infestans_%28late_blight%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Tomato_with_Phytophthora_infestans_%28late_blight%29.jpg/440px-Tomato_with_Phytophthora_infestans_%28late_blight%29.jpg 2x" data-file-width="1440" data-file-height="726" /></a><figcaption></figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Frameless"><a href="/wiki/File:Tomato_late_blight_stem_lesion_3_(5816739322).jpg" class="mw-file-description"><img alt="Infected tomato plant" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Tomato_late_blight_stem_lesion_3_%285816739322%29.jpg/220px-Tomato_late_blight_stem_lesion_3_%285816739322%29.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Tomato_late_blight_stem_lesion_3_%285816739322%29.jpg/330px-Tomato_late_blight_stem_lesion_3_%285816739322%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Tomato_late_blight_stem_lesion_3_%285816739322%29.jpg/440px-Tomato_late_blight_stem_lesion_3_%285816739322%29.jpg 2x" data-file-width="2050" data-file-height="1367" /></a><figcaption></figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Frameless"><a href="/wiki/File:Tomato_late_blight_fruit_cluster_(5816739612).jpg" class="mw-file-description"><img alt="Infected unripe tomatoes" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/Tomato_late_blight_fruit_cluster_%285816739612%29.jpg/220px-Tomato_late_blight_fruit_cluster_%285816739612%29.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e3/Tomato_late_blight_fruit_cluster_%285816739612%29.jpg/330px-Tomato_late_blight_fruit_cluster_%285816739612%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e3/Tomato_late_blight_fruit_cluster_%285816739612%29.jpg/440px-Tomato_late_blight_fruit_cluster_%285816739612%29.jpg 2x" data-file-width="2784" data-file-height="1856" /></a><figcaption></figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Phytophtora_infestans-effects.jpg" class="mw-file-description"><img alt="Infected potatoes are shrunken on the outside, and corky as well as rotted on the inside." src="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Phytophtora_infestans-effects.jpg/220px-Phytophtora_infestans-effects.jpg" decoding="async" width="220" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Phytophtora_infestans-effects.jpg/330px-Phytophtora_infestans-effects.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/92/Phytophtora_infestans-effects.jpg/440px-Phytophtora_infestans-effects.jpg 2x" data-file-width="2790" data-file-height="2003" /></a><figcaption>Infected <a href="/wiki/Potato" title="Potato">potato</a></figcaption></figure> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:07_08_life_cycle,_Phytophthora_infestans_on_potato,_Peronosporales,_Oomycota_(M._Piepenbring).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/07_08_life_cycle%2C_Phytophthora_infestans_on_potato%2C_Peronosporales%2C_Oomycota_%28M._Piepenbring%29.png/293px-07_08_life_cycle%2C_Phytophthora_infestans_on_potato%2C_Peronosporales%2C_Oomycota_%28M._Piepenbring%29.png" decoding="async" width="293" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/07_08_life_cycle%2C_Phytophthora_infestans_on_potato%2C_Peronosporales%2C_Oomycota_%28M._Piepenbring%29.png/440px-07_08_life_cycle%2C_Phytophthora_infestans_on_potato%2C_Peronosporales%2C_Oomycota_%28M._Piepenbring%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c0/07_08_life_cycle%2C_Phytophthora_infestans_on_potato%2C_Peronosporales%2C_Oomycota_%28M._Piepenbring%29.png/586px-07_08_life_cycle%2C_Phytophthora_infestans_on_potato%2C_Peronosporales%2C_Oomycota_%28M._Piepenbring%29.png 2x" data-file-width="3070" data-file-height="2302" /></a><figcaption><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycle</a> on potato</figcaption></figure> The asexual life cycle of Phytophthora infestans is characterized by alternating phases of <a href="/wiki/Hypha" title="Hypha">hyphal</a> growth, sporulation, <a href="/wiki/Sporangia" class="mw-redirect" title="Sporangia">sporangia</a> germination (either through <a href="/wiki/Zoospore" title="Zoospore">zoospore</a> release or direct germination, i.e. germ tube emergence from the <a href="/wiki/Sporangium" title="Sporangium">sporangium</a>), and the re-establishment of hyphal growth.<a href="#cite_note-VCRB2012b-7">[7]</a> There is also a sexual cycle, which occurs when isolates of opposite mating type (A1 and A2, see <a href="#Mating_types">§ Mating types</a> below) meet. Hormonal communication triggers the formation of the sexual <a href="/wiki/Spore" title="Spore">spores</a>, called <a href="/wiki/Oospore" title="Oospore">oospores</a>.<a href="#cite_note-8">[8]</a> The different types of spores play major roles in the dissemination and survival of P. infestans. Sporangia are spread by wind or water and enable the movement of P. infestans between different <a href="/wiki/Host_(biology)" title="Host (biology)">host</a> plants. The zoospores released from sporangia are <a href="/wiki/Biflagellate" class="mw-redirect" title="Biflagellate">biflagellated</a> and <a href="/wiki/Chemotactic" class="mw-redirect" title="Chemotactic">chemotactic</a>, allowing further movement of P. infestans on water films found on leaves or soils. Both sporangia and zoospores are short-lived, in contrast to oospores which can persist in a viable form for many years. People can observe P. infestans produce dark green, then brown then black spots on the surface of potato leaves and stems, often near the tips or edges, where water or dew collects.<a href="#cite_note-ndsu-9">[9]</a> The sporangia and <a href="/wiki/Sporangiophore" title="Sporangiophore">sporangiophores</a> appear white on the lower surface of the foliage. As for tuber blight, the white <a href="/wiki/Mycelium" title="Mycelium">mycelium</a> often shows on the tubers' surface.<a href="#cite_note-10">[10]</a> Under ideal conditions, P. infestans completes its life cycle on potato or tomato foliage in about five days.<a href="#cite_note-VCRB2012b-7">[7]</a> Sporangia develop on the leaves, spreading through the crop when temperatures are above 10 °C (50 °F) and humidity is over 75–80% for 2 days or more. <a href="/wiki/Rain" title="Rain">Rain</a> can wash spores into the <a href="/wiki/Soil" title="Soil">soil</a> where they infect young tubers, and the spores can also travel long distances on the wind. The early stages of blight are easily missed. Symptoms include the appearance of dark blotches on leaf tips and plant stems. White mold will appear under the leaves in humid conditions and the whole plant may quickly collapse. Infected tubers develop grey or dark patches that are reddish brown beneath the skin, and quickly decay to a foul-smelling mush caused by the infestation of secondary <a href="/wiki/Bacterial_soft_rot" title="Bacterial soft rot">soft bacterial rots</a>. Seemingly healthy tubers may rot later when in store. P. infestans survives poorly in nature apart from on its plant hosts. Under most conditions, the hyphae and asexual sporangia can survive for only brief periods in plant debris or soil, and are generally killed off during frosts or very warm weather. The exceptions involve oospores, and hyphae present within tubers. The persistence of viable pathogen within tubers, such as those that are left in the ground after the previous year's harvest or left in cull piles is a major problem in disease management. In particular, <a href="/wiki/Volunteer_plant" class="mw-redirect" title="Volunteer plant">volunteer plants</a> sprouting from infected tubers are thought to be a major source of inoculum (or <a href="/wiki/Propagule" title="Propagule">propagules</a>) at the start of a growing season.<a href="#cite_note-pnw-11">[11]</a> This can have devastating effects by destroying entire crops. <div class="mw-heading mw-heading3"><h3 id="Mating_types">Mating types</h3>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=3" title="Edit section: Mating types">edit</a>]</div> The <a href="/wiki/Mating_type" title="Mating type">mating types</a> are broadly divided into A1 and A2.<a href="#cite_note-Judelson-2017-12">[12]</a><a href="#cite_note-Fry-2008-13">[13]</a> Until the 1980s populations could only be distinguished by <a href="/w/index.php?title=Virulence_assay&action=edit&redlink=1" class="new" title="Virulence assay (page does not exist)">virulence assays</a> and mating types, but since then more detailed analysis has shown that mating type and <a href="/wiki/Genotype" title="Genotype">genotype</a> are substantially decoupled.<a href="#cite_note-Fry-2016-14">[14]</a> These types each produce a <a href="/w/index.php?title=Mating_hormone&action=edit&redlink=1" class="new" title="Mating hormone (page does not exist)">mating hormone</a> of their own.<a href="#cite_note-Fry-2008-13">[13]</a><a href="#cite_note-Judelson-2017-12">[12]</a> Pathogen populations are grouped into clonal lineages of these mating types and includes: <div class="mw-heading mw-heading4"><h4 id="A1">A1</h4>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=4" title="Edit section: A1">edit</a>]</div> A1 produces a mating hormone, a <a href="/wiki/Diterpene" title="Diterpene">diterpene</a><a href="#cite_note-Fry-2008-13">[13]</a> α1.<a href="#cite_note-Judelson-2017-12">[12]</a> Clonal lineages of A1 include: <ul><li><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>CN-1CN-1, -2, -4, -5, -6, -7, -8 – mtDNA haplotype Ia, China in 1996–97<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">CN-3 – Ia, China, 1996–97<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">CN-10 – Ia, China, 2004<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">CN-11 – IIb, China, 2000 & 2002<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">CN01 – IIa, China, 2004–09<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">CN03 – Ia/IIb, China, 2004–09<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">FAM-1 – (only presumed to be A1), mtDNA haplo Ia subtype <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">HERB-1, Japan, Philippines, India, China, Malaysia, Nepal, present some time before 1950<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">IN-1 – Ia, India, Nepal, 1993<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">IN-2 – Ia, India, 1993<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">JP-2JP-2/SIB-1/RF006 – mtDNA haplo IIa, distinguishable by <a href="/w/index.php?title=RG57&action=edit&redlink=1" class="new" title="RG57 (page does not exist)">RG57</a>, intermediate level of <a href="/wiki/Metalaxyl_resistance" class="mw-redirect" title="Metalaxyl resistance">metalaxyl resistance</a>, Japan, China, Korea, Thailand, 1996–present<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">JP-3 – IIa, distinguishable by <a href="/w/index.php?title=RG57&action=edit&redlink=1" class="new" title="RG57 (page does not exist)">RG57</a>, intermediate level of <a href="/wiki/Metalaxyl_resistance" class="mw-redirect" title="Metalaxyl resistance">metalaxyl resistance</a>, Japan, 1996–present<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">JP-4 – IIa, distinguishable by <a href="/w/index.php?title=RG57&action=edit&redlink=1" class="new" title="RG57 (page does not exist)">RG57</a>, intermediate level of <a href="/wiki/Metalaxyl_resistance" class="mw-redirect" title="Metalaxyl resistance">metalaxyl resistance</a>, Japan, 1996–present<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">KR-1 Zhang sensu Zhang (not to be confused with <a href="#KR-1_Gotoh">#KR-1 sensu Gotoh</a> below) – IIa, Korea, 2002–04<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">KR_1_A1KR_1_A1 – mtDNA haplo unknown, Korea, 2009–16<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">MO-6 – Ia, China, 2004<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP-1 – Ia, India, Nepal, 1993, 1996–97<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP-2 – Ia, Nepal, 1997<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP1 – Ia, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP2 A1 – (Also A2, see <a href="#NP2_A2">#the A2 type of NP2</a> below) Ia, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP3 (not to be confused with <a href="#US-1">#US-1</a> below) – Ib, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-1 (not to be confused with <a href="#NP3/US-1">#NP3/US-1</a> above) – Ib,<a href="#cite_note-Fry-2008-13">[13]</a><a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a> China, India, Nepal, Japan, Taiwan, Thailand, Vietnam, 1940–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP4, 5, 7, and 9 – Ia, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP6 – mtDNA haplo unknown, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-11 – IIb, Taiwan, Korea, Vietnam, 1998–2016<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-16 – IIb, China, 2002 & 2004<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-17<a href="#cite_note-Fry-2008-13">[13]</a> – IIa, Korea, 2003–04<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-23<a href="#cite_note-Fry-2016-14">[14]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-24<a href="#cite_note-Fry-2016-14">[14]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">2_A1 – Ia, Indonesia, 2016–19<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">T30-4<a href="#cite_note-Fry-2016-14">[14]</a></li></ul> <div class="mw-heading mw-heading4"><h4 id="A2">A2</h4>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=5" title="Edit section: A2">edit</a>]</div> Discovered by <a href="/wiki/John_Niederhauser" title="John Niederhauser">John Niederhauser</a> in the 1950s, in the <a href="/wiki/Toluca_Valley" title="Toluca Valley">Toluca Valley</a> in Central Mexico, while working for the <a href="/wiki/Rockefeller_Foundation" title="Rockefeller Foundation">Rockefeller Foundation</a>'s Mexican Agriculture Program. Published in Niederhauser 1956.<a href="#cite_note-Fry-2008-13">[13]</a><a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a> A2 produces a mating hormone α2.<a href="#cite_note-Judelson-2017-12">[12]</a> Clonal lineages of A2 include: <ul><li>CN02 – See <a href="#CN02">#13_A2/CN02</a> below</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-22 – with <a href="/wiki/MtDNA" class="mw-redirect" title="MtDNA">mtDNA</a> <a href="/wiki/Haplotype" title="Haplotype">haplotype</a> H-20<a href="#cite_note-Fry-2016-14">[14]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">JP-1 – IIa, Japan, Korea, Indonesia, late 1980s–present<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">KR-1 Gotoh sensu Gotoh (not to be confused with <a href="#KR-1_Zhang">#KR-1 sensu Zhang</a> above) – IIa, differs from JP-1 by one RG57 band, Korea, 1992<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">KR_2_A2 – mtDNA haplo unknown, Korea, 2009–16<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">CN-9 – Ia, China, 2001<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP2 A2 – (Also A1, see <a href="#NP2_A1">#the A1 type of NP2</a> above) Ia, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP8 – Ib, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">NP10 & 11 – Ia, Nepal, 1999–2000<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">TH-1 – Ia, Thailand, China, Nepal, 1994 & 1997<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li>Unknown – Ib, India, 1996–2003<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">BR-1 – Brazil<a href="#cite_note-Fry-2008-13">[13]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-7<a href="#cite_note-Fry-2008-13">[13]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-8<a href="#cite_note-Fry-2008-13">[13]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">US-14 – IIa, Korea, 2002–03<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">13_A2<a href="#cite_note-16">[16]</a><a href="#cite_note-Fry-2016-14">[14]</a><a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a>/CN02 – Ia, China, India, Bangladesh, Nepal, Pakistan, Myanmar, 2005–19<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a></li></ul> <div class="mw-heading mw-heading4"><h4 id="Self-fertile">Self-fertile</h4>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=6" title="Edit section: Self-fertile">edit</a>]</div> A self-fertile type was present in China between 2009 and 2013.<a href="#cite_note-Guha-Roy-et-al-2021-15">[15]</a> <div class="mw-heading mw-heading2"><h2 id="Physiology">Physiology</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=7" title="Edit section: Physiology">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509"><a href="/wiki/PiINF1" class="mw-redirect" title="PiINF1">PiINF1</a> is the <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509"><a href="/wiki/INF1" title="INF1">INF1</a> in P. infestans. Hosts respond with <a href="/wiki/Autophagy" title="Autophagy">autophagy</a> upon detection of this <a href="/wiki/Elicitor" title="Elicitor">elicitor</a>, Liu et al. 2005 finding this to be the only alternative to mass <a href="/wiki/Hypersensitive_response" title="Hypersensitive response">hypersensitivity</a> leading to mass <a href="/wiki/Programmed_cell_death" title="Programmed cell death">programmed cell death</a>.<a href="#cite_note-Yang-et-al-2020-17">[17]</a> <div class="mw-heading mw-heading2"><h2 id="Genetics">Genetics</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=8" title="Edit section: Genetics">edit</a>]</div> P. infestans is <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a>, with about 8–10 <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a>, and in 2009 scientists completed the sequencing of its <a href="/wiki/Genome" title="Genome">genome</a>. The genome was found to be considerably larger (240 <a href="/wiki/Megabase" class="mw-redirect" title="Megabase">Mbp</a>) than that of most other Phytophthora species whose genomes have been sequenced; <a href="/wiki/Phytophthora_sojae" title="Phytophthora sojae">P. sojae</a> has a 95 Mbp genome and <a href="/wiki/Phytophthora_ramorum" title="Phytophthora ramorum">P. ramorum</a> had a 65 Mbp genome. About 18,000 <a href="/wiki/Gene" title="Gene">genes</a> were detected within the P. infestans genome. It also contained a diverse variety of <a href="/wiki/Transposon" class="mw-redirect" title="Transposon">transposons</a> and many <a href="/wiki/Gene_families" class="mw-redirect" title="Gene families">gene families</a> encoding for <a href="/wiki/Effector_protein" class="mw-redirect" title="Effector protein">effector proteins</a> that are involved in causing <a href="/wiki/Pathogen" title="Pathogen">pathogenicity</a>. These proteins are split into two main groups depending on whether they are produced by the water mold in the <a href="/wiki/Symplast" title="Symplast">symplast</a> (inside plant cells) or in the <a href="/wiki/Apoplast" title="Apoplast">apoplast</a> (between plant cells). Proteins produced in the symplast included <a href="/wiki/RXLR" class="mw-redirect" title="RXLR">RXLR</a> proteins, which contain an <a href="/wiki/Arginine" title="Arginine">arginine</a>-X-<a href="/wiki/Leucine" title="Leucine">leucine</a>-arginine (where X can be any <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a>) sequence at the <a href="/wiki/Amino_terminus" class="mw-redirect" title="Amino terminus">amino terminus</a> of the protein. Some RXLR proteins are <a href="/wiki/Avirulence" class="mw-redirect" title="Avirulence">avirulence</a> proteins, meaning that they can be detected by the plant and lead to a <a href="/wiki/Hypersensitive_response" title="Hypersensitive response">hypersensitive response</a> which restricts the growth of the pathogen. P. infestans was found to encode around 60% more of these proteins than most other Phytophthora species. Those found in the apoplast include <a href="/wiki/Hydrolytic_enzyme" class="mw-redirect" title="Hydrolytic enzyme">hydrolytic enzymes</a> such as <a href="/wiki/Protease" title="Protease">proteases</a>, <a href="/wiki/Lipase" title="Lipase">lipases</a> and <a href="/wiki/Glycosylase" title="Glycosylase">glycosylases</a> that act to degrade plant tissue, <a href="/wiki/Enzyme_inhibitor" title="Enzyme inhibitor">enzyme inhibitors</a> to protect against host defence enzymes and <a href="/wiki/Necrosis" title="Necrosis">necrotizing</a> toxins. Overall the genome was found to have an extremely high repeat content (around 74%) and to have an unusual gene distribution in that some areas contain many genes whereas others contain very few.<a href="#cite_note-bbc_news-1">[1]</a><a href="#cite_note-genome-18">[18]</a> The pathogen shows high <a href="/wiki/Allelic" class="mw-redirect" title="Allelic">allelic</a> diversity in many <a href="/wiki/Microbial_isolate" class="mw-redirect" title="Microbial isolate">isolates</a> collected in <a href="/wiki/Europe" title="Europe">Europe</a>.<a href="#cite_note-Global-Perspective-19">[19]</a> This may be due to widespread <a href="/wiki/Trisomy" title="Trisomy">trisomy</a> or <a href="/wiki/Polyploidy" title="Polyploidy">polyploidy</a> in those populations.<a href="#cite_note-Global-Perspective-19">[19]</a>: 61  <div class="mw-heading mw-heading2"><h2 id="Research">Research</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=9" title="Edit section: Research">edit</a>]</div> <a href="/wiki/Mycology" title="Mycology">Study</a> of P. infestans presents <a href="/wiki/Microbial_isolate" class="mw-redirect" title="Microbial isolate">sampling</a> difficulties in the United States.<a href="#cite_note-Genetics-Ecology-Evolution-20">[20]</a>: 43  It occurs only sporadically and usually has significant <a href="/wiki/Founder_effect" title="Founder effect">founder effects</a> due to each epidemic starting from introduction of a single <a href="/wiki/Genotype" title="Genotype">genotype</a>.<a href="#cite_note-Genetics-Ecology-Evolution-20">[20]</a>: 43  <div class="mw-heading mw-heading2"><h2 id="Origin_and_diversity">Origin and diversity</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=10" title="Edit section: Origin and diversity">edit</a>]</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">See also: <a href="/wiki/Plant_disease_epidemiology" title="Plant disease epidemiology">Plant disease epidemiology</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Modell_eines_Querschnitts_durch_ein_Kartoffelblatt_mit_Phytophthora_infestans_(Kartoffelpilz)_-Osterloh_Nr._55-_-Brendel_10_k,_1-_(2).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/80/Modell_eines_Querschnitts_durch_ein_Kartoffelblatt_mit_Phytophthora_infestans_%28Kartoffelpilz%29_-Osterloh_Nr._55-_-Brendel_10_k%2C_1-_%282%29.jpg/220px-Modell_eines_Querschnitts_durch_ein_Kartoffelblatt_mit_Phytophthora_infestans_%28Kartoffelpilz%29_-Osterloh_Nr._55-_-Brendel_10_k%2C_1-_%282%29.jpg" decoding="async" width="220" height="336" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/80/Modell_eines_Querschnitts_durch_ein_Kartoffelblatt_mit_Phytophthora_infestans_%28Kartoffelpilz%29_-Osterloh_Nr._55-_-Brendel_10_k%2C_1-_%282%29.jpg/330px-Modell_eines_Querschnitts_durch_ein_Kartoffelblatt_mit_Phytophthora_infestans_%28Kartoffelpilz%29_-Osterloh_Nr._55-_-Brendel_10_k%2C_1-_%282%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/80/Modell_eines_Querschnitts_durch_ein_Kartoffelblatt_mit_Phytophthora_infestans_%28Kartoffelpilz%29_-Osterloh_Nr._55-_-Brendel_10_k%2C_1-_%282%29.jpg/440px-Modell_eines_Querschnitts_durch_ein_Kartoffelblatt_mit_Phytophthora_infestans_%28Kartoffelpilz%29_-Osterloh_Nr._55-_-Brendel_10_k%2C_1-_%282%29.jpg 2x" data-file-width="2298" data-file-height="3510" /></a><figcaption>Histological model of a potato leaf cross-section, <a href="/wiki/Botanical_Museum_Greifswald" title="Botanical Museum Greifswald">Botanical Museum Greifswald</a></figcaption></figure> The highlands of central <a href="/wiki/Mexico" title="Mexico">Mexico</a> are considered by many to be the center of origin of P. infestans, although others have proposed its origin to be in the <a href="/wiki/Andes" title="Andes">Andes</a>, which is also the origin of potatoes.<a href="#cite_note-21">[21]</a><a href="#cite_note-grunwald-2005-22">[22]</a> A recent study evaluated these two alternate hypotheses and found conclusive support for central Mexico being the center of origin.<a href="#cite_note-goss-2014-23">[23]</a> Support for Mexico – specifically the <a href="/wiki/Toluca_Valley" title="Toluca Valley">Toluca Valley</a><a href="#cite_note-Genetics-Ecology-Evolution-20">[20]</a> – comes from multiple observations including the fact that populations are genetically most diverse in Mexico, late blight is observed in native tuber-bearing <a href="/wiki/Solanum" title="Solanum">Solanum</a> species, populations of the pathogen are in <a href="/wiki/Hardy%E2%80%93Weinberg_equilibrium" class="mw-redirect" title="Hardy–Weinberg equilibrium">Hardy–Weinberg equilibrium</a>, the two mating (see <a href="#Mating_types">§ Mating types</a> above) types occur in a 1:1 ratio, and detailed <a href="/wiki/Phylogeographic" class="mw-redirect" title="Phylogeographic">phylogeographic</a> and evolutionary studies.<a href="#cite_note-grunwald-2005-22">[22]</a><a href="#cite_note-goss-2014-23">[23]</a><a href="#cite_note-24">[24]</a><a href="#cite_note-25">[25]</a><a href="#cite_note-26">[26]</a><a href="#cite_note-27">[27]</a><a href="#cite_note-28">[28]</a> Furthermore, the closest relatives of P. infestans, namely <a href="/w/index.php?title=Phytophthora_mirabilis&action=edit&redlink=1" class="new" title="Phytophthora mirabilis (page does not exist)">P. mirabilis</a> and <a href="/w/index.php?title=Phytophthora_ipomoeae&action=edit&redlink=1" class="new" title="Phytophthora ipomoeae (page does not exist)">P. ipomoeae</a> are endemic to central Mexico.<a href="#cite_note-29">[29]</a> On the other hand, the only close relative found in <a href="/wiki/South_America" title="South America">South America</a>, namely <a href="/w/index.php?title=Phytophthora_andina&action=edit&redlink=1" class="new" title="Phytophthora andina (page does not exist)">P. andina</a>, is a hybrid that does not share a single common ancestor with P. infestans. Finally, populations of P. infestans in South America lack genetic diversity and are <a href="/wiki/Vegetative_reproduction" title="Vegetative reproduction">clonal</a>.<a href="#cite_note-goss-2014-23">[23]</a><a href="#cite_note-30">[30]</a><a href="#cite_note-31">[31]</a> Migrations from Mexico to <a href="/wiki/North_America" title="North America">North America</a> or <a href="/wiki/Europe" title="Europe">Europe</a> have occurred several times throughout history, probably linked to the movement of tubers.<a href="#cite_note-32">[32]</a><a href="#cite_note-33">[33]</a> Until the 1970s, the A2 mating type was restricted to Mexico, but now in many regions of the world both A1 and A2 isolates can be found in the same region.<a href="#cite_note-Fry-2008-13">[13]</a> The co-occurrence of the two mating types is significant due to the possibility of sexual recombination and formation of oospores, which can survive the winter. Only in Mexico and <a href="/wiki/Scandinavia" title="Scandinavia">Scandinavia</a>, however, is oospore formation thought to play a role in overwintering.<a href="#cite_note-grunwald-2005-22">[22]</a><a href="#cite_note-34">[34]</a> In other parts of Europe, increasing genetic diversity has been observed as a consequence of sexual reproduction.<a href="#cite_note-PLOS2014-35">[35]</a> This is notable since different forms of P. infestans vary in their aggressiveness on potato or tomato, in sporulation rate, and sensitivity to <a href="/wiki/Fungicide" title="Fungicide">fungicides</a>.<a href="#cite_note-36">[36]</a> Variation in such traits also occurs in North America, however importation of new <a href="/wiki/Genotype" title="Genotype">genotypes</a> from Mexico appears to be the predominant cause of genetic diversity, as opposed to sexual recombination within potato or tomato fields.<a href="#cite_note-Fry-2008-13">[13]</a> In 1976 – due to a summer drought in Europe – there was a potato production shortfall and so eating potatoes were imported to fill the shortfall. It is thought that this was the vehicle for mating type A2 to reach the rest of the world. In any case, there had been little diversity, consisting of the US-1 strain, and of that only one type of: mating type, mtDNA, <a href="/wiki/Restriction_fragment_length_polymorphism" title="Restriction fragment length polymorphism">restriction fragment length polymorphism</a>, and di-locus[<a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify">clarification needed</a>] <a href="/wiki/Isozyme" title="Isozyme">isozyme</a>. Then in 1980 suddenly greater diversity and A2 appeared in Europe. In 1981 it was found in the Netherlands, United Kingdom, 1985 in Sweden, the early 1990s in Norway and Finland, 1996 in Denmark, and 1999 in Iceland. In the UK new A1 lineages only replaced the old lineage by end of the '80s, and A2 spread even more slowly, with Britain having low levels and Ireland (north and Republic) having none-to-trace detections through the '90s.<a href="#cite_note-Cooke-et-al-2011-37">[37]</a> Many of the strains that appeared outside of Mexico since the 1980s have been more aggressive, leading to increased crop losses.<a href="#cite_note-Fry-2008-13">[13]</a> In Europe since 2013 the populations have been tracked by the EuroBlight network (see links below). Some of the differences between strains may be related to variation in the RXLR effectors that are present. <div class="mw-heading mw-heading2"><h2 id="Disease_management">Disease management</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=11" title="Edit section: Disease management">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Plant_pathology" title="Plant pathology">Plant pathology</a></div> P. infestans is still a difficult disease to control.<a href="#cite_note-Wiley2013-3">[3]</a><a href="#cite_note-Desiree-Victoria-no-late-blight-Africa-38">[38]</a><a href="#cite_note-Ghislain-et-al-2018-39">[39]</a><a href="#cite_note-Modern-Farmer-Victoria-immunity-40">[40]</a> There are many chemical options in <a href="/wiki/Agriculture" title="Agriculture">agriculture</a> for the control of damage to the <a href="/wiki/Foliage" class="mw-redirect" title="Foliage">foliage</a> as well as the fruit (for tomatoes) and the <a href="/wiki/Tuber" title="Tuber">tuber</a><a href="#cite_note-41">[41]</a> (for potatoes). A few of the most common foliar-applied fungicides are <a href="/w/index.php?title=Ridomil&action=edit&redlink=1" class="new" title="Ridomil (page does not exist)">Ridomil</a>, a Gavel/<a href="/w/index.php?title=SuperTin&action=edit&redlink=1" class="new" title="SuperTin (page does not exist)">SuperTin</a> tank mix, and <a href="/w/index.php?title=Previcur_Flex&action=edit&redlink=1" class="new" title="Previcur Flex (page does not exist)">Previcur Flex</a>. All of the aforementioned fungicides need to be tank mixed with a broad-spectrum fungicide such as <a href="/wiki/Mancozeb" title="Mancozeb">mancozeb</a> or <a href="/wiki/Chlorothalonil" title="Chlorothalonil">chlorothalonil</a> not just for resistance management but also because the potato plants will be attacked by other pathogens at the same time. If adequate field scouting occurs and late blight is found soon after disease development, localized patches of potato plants can be killed with a <a href="/wiki/Desiccant" title="Desiccant">desiccant</a> (e.g. <a href="/wiki/Paraquat" title="Paraquat">paraquat</a>) through the use of a backpack sprayer. This management technique can be thought of as a field-scale <a href="/wiki/Hypersensitive_response" title="Hypersensitive response">hypersensitive response</a> similar to what occurs in some plant-viral interactions whereby cells surrounding the initial point of infection are killed in order to prevent proliferation of the pathogen. If infected tubers make it into a storage bin, there is a very high risk to the storage life of the entire bin. Once in storage, there isn't much that can be done besides emptying the parts of the bin that contain tubers infected with Phytophthora infestans. To increase the probability of successfully storing potatoes from a field where late blight was known to occur during the growing season, some products can be applied just prior to entering storage (e.g. <a href="/w/index.php?title=Phostrol&action=edit&redlink=1" class="new" title="Phostrol (page does not exist)">Phostrol</a>).<a href="#cite_note-42">[42]</a> Around the world the disease causes around $6 billion of damage to crops each year.<a href="#cite_note-bbc_news-1">[1]</a><a href="#cite_note-PlDis2011-2">[2]</a> <div class="mw-heading mw-heading3"><h3 id="Resistant_plants">Resistant plants</h3>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=12" title="Edit section: Resistant plants">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Plant_disease_resistance" title="Plant disease resistance">Plant disease resistance</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Cisgenicpotatoes.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/2/21/Cisgenicpotatoes.JPG/220px-Cisgenicpotatoes.JPG" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/2/21/Cisgenicpotatoes.JPG/330px-Cisgenicpotatoes.JPG 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/2/21/Cisgenicpotatoes.JPG/440px-Cisgenicpotatoes.JPG 2x" data-file-width="1600" data-file-height="1200" /></a><figcaption>Potatoes after exposure. The normal potatoes have blight but the <a href="/wiki/Cisgenic" class="mw-redirect" title="Cisgenic">cisgenic</a> potatoes are healthy.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:%C3%84lskade_kn%C3%B6l_%E2%80%93_utflykt_med_SLU_till_deras_potatisf%C3%B6rs%C3%B6ksodlingar_%E2%80%93_20190825_09_(cropped).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bb/%C3%84lskade_kn%C3%B6l_%E2%80%93_utflykt_med_SLU_till_deras_potatisf%C3%B6rs%C3%B6ksodlingar_%E2%80%93_20190825_09_%28cropped%29.jpg/220px-%C3%84lskade_kn%C3%B6l_%E2%80%93_utflykt_med_SLU_till_deras_potatisf%C3%B6rs%C3%B6ksodlingar_%E2%80%93_20190825_09_%28cropped%29.jpg" decoding="async" width="220" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bb/%C3%84lskade_kn%C3%B6l_%E2%80%93_utflykt_med_SLU_till_deras_potatisf%C3%B6rs%C3%B6ksodlingar_%E2%80%93_20190825_09_%28cropped%29.jpg/330px-%C3%84lskade_kn%C3%B6l_%E2%80%93_utflykt_med_SLU_till_deras_potatisf%C3%B6rs%C3%B6ksodlingar_%E2%80%93_20190825_09_%28cropped%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bb/%C3%84lskade_kn%C3%B6l_%E2%80%93_utflykt_med_SLU_till_deras_potatisf%C3%B6rs%C3%B6ksodlingar_%E2%80%93_20190825_09_%28cropped%29.jpg/440px-%C3%84lskade_kn%C3%B6l_%E2%80%93_utflykt_med_SLU_till_deras_potatisf%C3%B6rs%C3%B6ksodlingar_%E2%80%93_20190825_09_%28cropped%29.jpg 2x" data-file-width="3990" data-file-height="2464" /></a><figcaption>Genetically modified <a href="/wiki/King_Edward_potato" title="King Edward potato">King Edward</a> (right) next to King Edward which has not been genetically modified (left). Research field, <a href="/wiki/Swedish_University_of_Agricultural_Sciences" title="Swedish University of Agricultural Sciences">Swedish University of Agricultural Sciences</a>, 2019</figcaption></figure> Breeding for resistance, particularly in potato plants, has had limited success in part due to difficulties in crossing cultivated potato to its wild relatives,<a href="#cite_note-Desiree-Victoria-no-late-blight-Africa-38">[38]</a><a href="#cite_note-Ghislain-et-al-2018-39">[39]</a><a href="#cite_note-Modern-Farmer-Victoria-immunity-40">[40]</a> which are the source of potential resistance genes.<a href="#cite_note-Desiree-Victoria-no-late-blight-Africa-38">[38]</a><a href="#cite_note-Ghislain-et-al-2018-39">[39]</a><a href="#cite_note-Modern-Farmer-Victoria-immunity-40">[40]</a> In addition, most resistance genes only work against a subset of P. infestans isolates, since effective <a href="/wiki/Plant_disease_resistance" title="Plant disease resistance">plant disease resistance</a> only results when the pathogen expresses a RXLR effector gene that matches the corresponding plant resistance (R) gene; <a href="/wiki/Gene-for-gene_relationship" title="Gene-for-gene relationship">effector-R</a> gene interactions trigger a range of plant defenses, such as the production of compounds toxic to the pathogen. Potato and tomato varieties vary in their susceptibility to blight.<a href="#cite_note-PLOS2014-35">[35]</a><a href="#cite_note-Desiree-Victoria-no-late-blight-Africa-38">[38]</a><a href="#cite_note-Ghislain-et-al-2018-39">[39]</a><a href="#cite_note-Modern-Farmer-Victoria-immunity-40">[40]</a> Most early varieties are very vulnerable; they should be planted early so that the crop matures before blight starts (usually in July in the Northern Hemisphere). Many old crop varieties, such as <a href="/wiki/King_Edward_potato" title="King Edward potato">King Edward potato</a> are also very susceptible but are grown because they are wanted commercially. Maincrop varieties which are very slow to develop blight include <a href="/wiki/List_of_potato_cultivars" title="List of potato cultivars">Cara</a>, Stirling, Teena, Torridon, <a href="/wiki/List_of_potato_cultivars" title="List of potato cultivars">Remarka</a>, and Romano. Some so-called resistant varieties can resist some strains of blight and not others, so their performance may vary depending on which are around.<a href="#cite_note-PLOS2014-35">[35]</a><a href="#cite_note-Desiree-Victoria-no-late-blight-Africa-38">[38]</a><a href="#cite_note-Ghislain-et-al-2018-39">[39]</a><a href="#cite_note-Modern-Farmer-Victoria-immunity-40">[40]</a> These crops have had polygenic resistance bred into them, and are known as "field resistant". New varieties<a href="#cite_note-Desiree-Victoria-no-late-blight-Africa-38">[38]</a><a href="#cite_note-Ghislain-et-al-2018-39">[39]</a><a href="#cite_note-Modern-Farmer-Victoria-immunity-40">[40]</a> such as Sarpo Mira and Sarpo Axona show great resistance to blight even in areas of heavy infestation. Defender is an American cultivar whose parentage includes <a href="/wiki/Ranger_Russet" title="Ranger Russet">Ranger Russet</a> and <a href="/wiki/Poland" title="Poland">Polish</a> potatoes resistant to late blight. It is a long white-skinned cultivar with both foliar and tuber resistance to late blight. Defender was released in 2004.<a href="#cite_note-43">[43]</a> Genetic engineering may also provide options for generating resistance cultivars. A resistance gene effective against most known strains of blight has been identified from a wild relative of the potato, <a href="/wiki/Solanum_bulbocastanum" title="Solanum bulbocastanum">Solanum bulbocastanum</a>, and introduced by genetic engineering into cultivated varieties of potato.<a href="#cite_note-44">[44]</a> This is an example of <a href="/wiki/Cisgenesis" title="Cisgenesis">cisgenic</a> genetic engineering.<a href="#cite_note-45">[45]</a> <a href="/wiki/Melatonin" title="Melatonin">Melatonin</a> in the plant/P. infestans co-environment reduces the stress tolerance of the parasite.<a href="#cite_note-Socaciu-et-al-2020-46">[46]</a> <div class="mw-heading mw-heading3"><h3 id="Reducing_inoculum">Reducing inoculum</h3>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=13" title="Edit section: Reducing inoculum">edit</a>]</div> Blight can be controlled by limiting the source of <a href="https://en.wiktionary.org/wiki/inoculum" class="extiw" title="wikt:inoculum">inoculum</a>.<a href="#cite_note-PLOS2014-35">[35]</a> Only good-quality <a href="/wiki/Seed_potato" class="mw-redirect" title="Seed potato">seed potatoes</a> and tomatoes obtained from <a href="/wiki/Certified" class="mw-redirect" title="Certified">certified</a> suppliers should be planted. Often discarded potatoes from the previous <a href="/wiki/Season" title="Season">season</a> and self-sown <a href="/wiki/Tuber" title="Tuber">tubers</a> can act as sources of inoculum.<a href="#cite_note-47">[47]</a> <a href="/wiki/Compost" title="Compost">Compost</a>, soil or potting medium can be heat-treated to kill oomycetes such as Phytophthora infestans. The recommended sterilisation temperature for oomycetes is 120 °F (49 °C) for 30 minutes.<a href="#cite_note-48">[48]</a><a href="#cite_note-49">[49]</a> <div class="mw-heading mw-heading3"><h3 id="Environmental_conditions">Environmental conditions</h3>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=14" title="Edit section: Environmental conditions">edit</a>]</div> There are several environmental conditions that are conducive to P. infestans. An example of such took place in the United States during the 2009 growing season. As colder than average for the season and with greater than average rainfall, there was a major infestation of tomato plants, specifically in the eastern states.<a href="#cite_note-50">[50]</a> By using <a href="/wiki/Weather_forecasting" title="Weather forecasting">weather forecasting</a> systems, such as <a href="/w/index.php?title=BLITECAST&action=edit&redlink=1" class="new" title="BLITECAST (page does not exist)">BLITECAST</a>, if the following conditions occur as the <a href="/wiki/Canopy_(biology)" title="Canopy (biology)">canopy</a> of the crop closes, then the use of <a href="/wiki/Fungicide" title="Fungicide">fungicides</a> is recommended to prevent an <a href="/wiki/Epidemic" title="Epidemic">epidemic</a>.<a href="#cite_note-51">[51]</a> <ul><li>A <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">Beaumont Period is a period of 48 consecutive hours, in at least 46 of which the hourly readings of <a href="/wiki/Temperature" title="Temperature">temperature</a> and <a href="/wiki/Relative_humidity" class="mw-redirect" title="Relative humidity">relative humidity</a> at a given place have not been less than 10 °C (50 °F) and 75%, respectively.<a href="#cite_note-52">[52]</a><a href="#cite_note-Microbial-53">[53]</a></li> <li>A <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238216509">Smith Period is at least two <a href="/wiki/Consecutive" title="Consecutive">consecutive</a> days where min temperature is 10 °C (50 °F) or above and on each day at least 11 hours when the relative humidity is greater than 90%.</li></ul> The Beaumont and Smith periods have traditionally been used by growers in the <a href="/wiki/United_Kingdom" title="United Kingdom">United Kingdom</a>, with different criteria developed by growers in other regions.<a href="#cite_note-Microbial-53">[53]</a> The Smith period has been the preferred system used in the UK since its introduction in the 1970s.<a href="#cite_note-54">[54]</a> Based on these conditions and other factors, several tools have been developed to help growers manage the disease and plan fungicide applications. Often these are deployed as part of <a href="/wiki/Decision_support_system" title="Decision support system">Decision support systems</a> accessible through web sites or smart phones. Several studies have attempted to develop systems for real-time detection via <a href="/wiki/Flow_cytometry" title="Flow cytometry">flow cytometry</a> or <a href="/wiki/Microscopy" title="Microscopy">microscopy</a> of airborne sporangia collected in air samplers.<a href="#cite_note-55">[55]</a><a href="#cite_note-56">[56]</a><a href="#cite_note-57">[57]</a> Whilst these methods show potential to allow detection of sporangia in advance of occurrence of detectable disease symptoms on plants, and would thus be useful in enhancing existing <a href="/wiki/Decision_support_system" title="Decision support system">Decision support systems</a>, none have been commercially deployed to date. <div class="mw-heading mw-heading3"><h3 id="Use_of_fungicides">Use of fungicides</h3>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=15" title="Edit section: Use of fungicides">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Potato_blight_spraying_system.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/80/Potato_blight_spraying_system.jpg/220px-Potato_blight_spraying_system.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/80/Potato_blight_spraying_system.jpg/330px-Potato_blight_spraying_system.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/80/Potato_blight_spraying_system.jpg/440px-Potato_blight_spraying_system.jpg 2x" data-file-width="2400" data-file-height="1600" /></a><figcaption><a href="/wiki/Fungicide_application" class="mw-redirect" title="Fungicide application">Spraying</a> potato, <a href="/wiki/Nottinghamshire" title="Nottinghamshire">Nottinghamshire</a></figcaption></figure> <a href="/wiki/Fungicide" title="Fungicide">Fungicides</a> for the control of potato blight are normally only used in a preventative manner, optionally in <a href="https://en.wiktionary.org/wiki/conjunction" class="extiw" title="wikt:conjunction">conjunction</a> with disease <a href="/wiki/Forecasting" title="Forecasting">forecasting</a>. In susceptible varieties, sometimes fungicide applications may be needed weekly. An early spray is most effective. The choice of fungicide can depend on the nature of local strains of P. infestans. <a href="/wiki/Metalaxyl" title="Metalaxyl">Metalaxyl</a> is a fungicide that was marketed for use against P. infestans, but suffered serious <a href="/wiki/Pesticide_resistance" title="Pesticide resistance">resistance</a> issues when used on its own. In some regions of the world during the 1980s and 1990s, most strains of P. infestans became resistant to metalaxyl, but in subsequent years many populations shifted back to sensitivity. To reduce the occurrence of resistance, it is strongly advised to use single-target fungicides such as metalaxyl along with <a href="/wiki/Carbamate" title="Carbamate">carbamate</a> compounds. A combination of other compounds are recommended for managing metalaxyl-resistant strains. These include <a href="/w/index.php?title=Mandipropamid&action=edit&redlink=1" class="new" title="Mandipropamid (page does not exist)">mandipropamid</a>, <a href="/wiki/Chlorothalonil" title="Chlorothalonil">chlorothalonil</a>, <a href="/wiki/Fluazinam" title="Fluazinam">fluazinam</a>, <a href="/wiki/Triphenyltin_hydroxide" title="Triphenyltin hydroxide">triphenyltin</a>, <a href="/wiki/Mancozeb" title="Mancozeb">mancozeb</a>, and others. In the United States, the <a href="/wiki/Environmental_Protection_Agency" class="mw-redirect" title="Environmental Protection Agency">Environmental Protection Agency</a> has approved <a href="/wiki/Oxathiapiprolin" title="Oxathiapiprolin">oxathiapiprolin</a> for use against late blight.<a href="#cite_note-58">[58]</a> In African <a href="/wiki/Smallholder" class="mw-redirect" title="Smallholder">smallholder</a> production <a href="/wiki/Fungicide_application" class="mw-redirect" title="Fungicide application">fungicide application</a> can be necessary up to once every three days.<a href="#cite_note-Ally4Sci-59">[59]</a> <div class="mw-heading mw-heading4"><h4 id="In_organic_production">In organic production</h4>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=16" title="Edit section: In organic production">edit</a>]</div> In the past, <a href="/wiki/Copper(II)_sulfate" title="Copper(II) sulfate">copper(II) sulfate</a> solution (called '<a href="/wiki/Copper(II)_sulfate" title="Copper(II) sulfate">bluestone</a>') was used to combat potato blight. <a href="/wiki/Copper_pesticide" title="Copper pesticide">Copper pesticides</a> remain in use on organic crops, both in the form of <a href="/wiki/Copper_hydroxide" class="mw-redirect" title="Copper hydroxide">copper hydroxide</a> and copper sulfate. Given the dangers of <a href="/wiki/Copper_toxicity" title="Copper toxicity">copper toxicity</a>, other organic control options that have been shown to be effective include <a href="/wiki/Horticultural_oil" title="Horticultural oil">horticultural oils</a>, <a href="/wiki/Phosphorous_acid" title="Phosphorous acid">phosphorous acids</a>, and <a href="/wiki/Rhamnolipid" title="Rhamnolipid">rhamnolipid</a> <a href="/wiki/Biosurfactant" title="Biosurfactant">biosurfactants</a>, while sprays containing "beneficial" microbes such as <a href="/wiki/Bacillus_subtilis" title="Bacillus subtilis">Bacillus subtilis</a> or compounds that encourage the plant to produce defensive chemicals (such as <a href="/wiki/Fallopia_sachalinensis" class="mw-redirect" title="Fallopia sachalinensis">knotweed</a> extract) have not performed as well.<a href="#cite_note-60">[60]</a> During the crop year 2008, many of the <a href="/wiki/Certified_organic_food" class="mw-redirect" title="Certified organic food">certified organic</a> potatoes produced in the United Kingdom and certified by the <a href="/wiki/Soil_Association" title="Soil Association">Soil Association</a> as organic were sprayed with a <a href="/wiki/Copper_pesticide" title="Copper pesticide">copper pesticide</a><a href="#cite_note-61">[61]</a> to control potato blight. According to the Soil Association, the total copper that can be applied to organic land is 6 kilograms per hectare (5.4 lb/acre)/year.<a href="#cite_note-62">[62]</a> <div class="mw-heading mw-heading3"><h3 id="Control_of_tuber_blight">Control of tuber blight</h3>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=17" title="Edit section: Control of tuber blight">edit</a>]</div> Ridging is often used to reduce tuber contamination by blight. This normally involves piling <a href="/wiki/Soil" title="Soil">soil</a> or <a href="/wiki/Mulch" title="Mulch">mulch</a> around the <a href="/wiki/Plant_stem" title="Plant stem">stems</a> of the potato blight meaning the <a href="/wiki/Pathogen" title="Pathogen">pathogen</a> has farther to travel to get to the tuber.<a href="#cite_note-63">[63]</a> Another approach is to destroy the canopy around five weeks before <a href="/wiki/Harvest" title="Harvest">harvest</a>, using a contact <a href="/wiki/Herbicide" title="Herbicide">herbicide</a> or <a href="/wiki/Sulfuric_acid" title="Sulfuric acid">sulfuric acid</a> to burn off the foliage. Eliminating infected foliage reduces the likelihood of tuber infection. <div class="mw-heading mw-heading2"><h2 id="Historical_impact">Historical impact</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=18" title="Edit section: Historical impact">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Timeline_of_plant_pathology" title="Timeline of plant pathology">Timeline of plant pathology</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Suggested_paths_of_migration_and_diversification_of_P._infestans_lineages_HERB-1_and_US-1.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Suggested_paths_of_migration_and_diversification_of_P._infestans_lineages_HERB-1_and_US-1.jpg/220px-Suggested_paths_of_migration_and_diversification_of_P._infestans_lineages_HERB-1_and_US-1.jpg" decoding="async" width="220" height="125" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Suggested_paths_of_migration_and_diversification_of_P._infestans_lineages_HERB-1_and_US-1.jpg/330px-Suggested_paths_of_migration_and_diversification_of_P._infestans_lineages_HERB-1_and_US-1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d3/Suggested_paths_of_migration_and_diversification_of_P._infestans_lineages_HERB-1_and_US-1.jpg/440px-Suggested_paths_of_migration_and_diversification_of_P._infestans_lineages_HERB-1_and_US-1.jpg 2x" data-file-width="2790" data-file-height="1582" /></a><figcaption>Suggested paths of migration and diversification of lineages <a href="#HERB-1">HERB-1</a> and <a href="#US-1">US-1</a></figcaption></figure> The first recorded instances of the disease were in the United States, in <a href="/wiki/Philadelphia" title="Philadelphia">Philadelphia</a> and <a href="/wiki/New_York_City" title="New York City">New York City</a> in early 1843.<a href="#cite_note-64">[64]</a> Winds then spread the spores, and in 1845 it was found from <a href="/wiki/Illinois" title="Illinois">Illinois</a> to <a href="/wiki/Nova_Scotia" title="Nova Scotia">Nova Scotia</a>, and from <a href="/wiki/Virginia" title="Virginia">Virginia</a> to <a href="/wiki/Ontario" title="Ontario">Ontario</a>. It crossed the <a href="/wiki/Atlantic_Ocean" title="Atlantic Ocean">Atlantic Ocean</a> with a shipment of seed potatoes for <a href="/wiki/Belgium" title="Belgium">Belgian</a> farmers in 1845.<a href="#cite_note-65">[65]</a><a href="#cite_note-66">[66]</a> The disease being first identified in Europe around <a href="/wiki/Kortrijk" title="Kortrijk">Kortrijk</a>, Flanders, in the June 1845, and resulted in the Flemish potato harvest failing that summer, yields declining 75-80%, leading to an estimated forty thousand deaths in the locale.<a href="#cite_note-67">[67]</a> All of the potato-growing countries in Europe would be affected, within a year. The effect of Phytophthora infestans in Ireland in 1845–52 was one of the factors which caused over one million to starve to death<a href="#cite_note-Bourouiba-2021-68">[68]</a> and forced another two million to emigrate. Most commonly referenced is the <a href="/wiki/Great_Famine_(Ireland)" title="Great Famine (Ireland)">Great Irish Famine</a>, during the late 1840s. Implicated in Ireland's fate was the island's disproportionate dependency on a single variety of potato, the <a href="/wiki/Irish_Lumper" title="Irish Lumper">Irish Lumper</a>. The lack of <a href="/wiki/Genetic_variability" title="Genetic variability">genetic variability</a> created a susceptible host population for the organism<a href="#cite_note-IrishCentral-69">[69]</a> after the blight strains <a href="/wiki/Potatoes_of_Chilo%C3%A9" title="Potatoes of Chiloé">originating in Chiloé Archipelago</a> replaced earlier potatoes of Peruvian origin in Europe.<a href="#cite_note-70">[70]</a> During the First World War, all of the copper in Germany was used for <a href="/wiki/Shell_casing" class="mw-redirect" title="Shell casing">shell casings</a> and <a href="/wiki/Electric_wire" class="mw-redirect" title="Electric wire">electric wire</a> and therefore none was available for making copper sulfate to spray potatoes. A major late blight outbreak on potato in Germany therefore went untreated, and the <a href="/wiki/Turnip_winter" class="mw-redirect" title="Turnip winter">resulting scarcity of potatoes</a> contributed to the <a href="/wiki/Blockade_of_Germany_(1914%E2%80%931919)#Death_toll" title="Blockade of Germany (1914–1919)">deaths from the blockade</a>.<a href="#cite_note-71">[71]</a><a href="#cite_note-72">[72]</a> Since 1941, Eastern Africa has been suffering potato production losses because of strains of P. infestans from Europe.<a href="#cite_note-73">[73]</a> <a href="/wiki/France" title="France">France</a>, <a href="/wiki/Canada" title="Canada">Canada</a>, the <a href="/wiki/United_States" title="United States">United States</a>, and the <a href="/wiki/Soviet_Union" title="Soviet Union">Soviet Union</a> researched P. infestans as a <a href="/wiki/Biological_weapon" class="mw-redirect" title="Biological weapon">biological weapon</a> in the 1940s and 1950s.<a href="#cite_note-weapon-74">[74]</a> Potato blight was one of more than 17 agents that the United States researched as potential biological weapons before the nation suspended <a href="/wiki/United_States_biological_weapons_program" title="United States biological weapons program">its biological weapons program</a>.<a href="#cite_note-jmcenter-75">[75]</a> Whether a weapon based on the pathogen would be effective is questionable, due to the difficulties in delivering viable pathogen to an enemy's fields, and the role of uncontrollable environmental factors in spreading the disease.<a href="#cite_note-76">[76]</a> Late blight (A2 type) has not yet been detected in Australia and strict biosecurity measures are in place. The disease has been seen in China, India and south-east Asian countries. A large outbreak of P. infestans occurred on tomato plants in the Northeast United States in 2009.<a href="#cite_note-77">[77]</a> In light of the periodic epidemics of P. infestans ever since its first emergence, it may be regarded as a periodically <a href="/wiki/Emerging_agricultural_pathogen" class="mw-redirect" title="Emerging agricultural pathogen">emerging pathogen</a> – or a periodically "re-emerging pathogen".<a href="#cite_note-Fones-et-al-2020-78">[78]</a><a href="#cite_note-Fry-et-al-2015-79">[79]</a> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=19" title="Edit section: References">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-bbc_news-1">^ <a href="#cite_ref-bbc_news_1-0">a</a> <a href="#cite_ref-bbc_news_1-1">b</a> <a href="#cite_ref-bbc_news_1-2">c</a> <style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFChand2009" class="citation cs2">Chand, Sudeep (9 September 2009), <a rel="nofollow" class="external text" href="http://news.bbc.co.uk/1/hi/sci/tech/8246944.stm">Killer genes cause potato famine</a>, <a href="/wiki/BBC_News" title="BBC News">BBC News</a>, retrieved 26 September 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Killer+genes+cause+potato+famine&rft.pub=BBC+News&rft.date=2009-09-09&rft.aulast=Chand&rft.aufirst=Sudeep&rft_id=http%3A%2F%2Fnews.bbc.co.uk%2F1%2Fhi%2Fsci%2Ftech%2F8246944.stm&rfr_id=info%3Asid%2Fen.wikipedia.org%3APhytophthora+infestans" class="Z3988"> </li> <li id="cite_note-PlDis2011-2">^ <a href="#cite_ref-PlDis2011_2-0">a</a> <a href="#cite_ref-PlDis2011_2-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNowicki2011" class="citation cs2">Nowicki, Marcin; et al. 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F; Jacobsen, E (2009), <a rel="nofollow" class="external text" href="http://library.wur.nl/WebQuery/wurpubs/381896">"Applied Biotechnology to Combat Late Blight in Potato Caused by Phytophthora Infestans"</a>, <a href="/wiki/Potato_Research" title="Potato Research">Potato Research</a> (Submitted manuscript), 52 (3): 249, <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%2Fs11540-009-9136-3">10.1007/s11540-009-9136-3</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:2850128">2850128</a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Potato+Research&rft.atitle=Applied+Biotechnology+to+Combat+Late+Blight+in+Potato+Caused+by+Phytophthora+Infestans&rft.volume=52&rft.issue=3&rft.pages=249&rft.date=2009&rft_id=info%3Adoi%2F10.1007%2Fs11540-009-9136-3&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A2850128%23id-name%3DS2CID&rft.aulast=Haverkort&rft.aufirst=A.+J&rft.au=Struik%2C+P.+C&rft.au=Visser%2C+R.+G.+F&rft.au=Jacobsen%2C+E&rft_id=http%3A%2F%2Flibrary.wur.nl%2FWebQuery%2Fwurpubs%2F381896&rfr_id=info%3Asid%2Fen.wikipedia.org%3APhytophthora+infestans" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFErwinRibeiro1996" class="citation cs2">Erwin, Donald C.; Ribeiro, Olaf K. (1996), Phytophthora Diseases Worldwide, <a href="/wiki/St._Paul,_MN" class="mw-redirect" title="St. Paul, MN">St. Paul, MN</a>: <a href="/w/index.php?title=American_Phytopathological_Society_Press&action=edit&redlink=1" class="new" title="American Phytopathological Society Press (page does not exist)">American Phytopathological Society Press</a>, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-89054-212-5" title="Special:BookSources/978-0-89054-212-5"><bdi>978-0-89054-212-5</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Phytophthora+Diseases+Worldwide&rft.place=St.+Paul%2C+MN&rft.pub=American+Phytopathological+Society+Press&rft.date=1996&rft.isbn=978-0-89054-212-5&rft.aulast=Erwin&rft.aufirst=Donald+C.&rft.au=Ribeiro%2C+Olaf+K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APhytophthora+infestans" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLucasShattockShawCooke1991" class="citation cs2">Lucas, J. 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Factsheets for Farmers. CABI (Centre for Agriculture and Bioscience International). <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1079%2Fpwkb.20147801535">10.1079/pwkb.20147801535</a>. Retrieved 2021-11-15.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Plantwise+Knowledge+Bank&rft.atitle=Potato+Late+Blight&rft.volume=Factsheets+for+Farmers&rft.date=2016&rft_id=info%3Adoi%2F10.1079%2Fpwkb.20147801535&rft.aulast=Omid&rft.aufirst=Shir+Ahmad&rft.au=%28Aga+Khan+Foundation%29&rft_id=http%3A%2F%2Fwww.plantwise.org%2Fknowledgebank%2Ffactsheetforfarmers%2F20147801535&rfr_id=info%3Asid%2Fen.wikipedia.org%3APhytophthora+infestans" class="Z3988"></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=Phytophthora_infestans&action=edit&section=21" title="Edit section: External links">edit</a>]</div> <ul><li><a rel="nofollow" class="external text" href="http://www.USAblight.org">USAblight A National Web Portal on Late Blight</a></li> <li><a rel="nofollow" class="external text" href="http://www.cipotato.org">International Potato Center</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20061014125542/http://www.btny.purdue.edu/USDA-ARS/Goodwin_lab/results/Phytoph_biblio.html">Online Phytophtora bibliography</a></li> <li><a rel="nofollow" class="external text" href="http://www.euroblight.net">EuroBlight a potato blight network in Europe</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20050519074314/http://www.barc.usda.gov/psi/vl/lateblight.htm">USDA-BARC Phytophthora infestans page</a></li> <li><a rel="nofollow" class="external text" href="https://attra.ncat.org/attra-pub-summaries/?pub=123">Organic Alternatives for Late Blight Control in Potatoes, from ATTRA</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200925005014/https://attra.ncat.org/attra-pub-summaries/?pub=123">Archived</a> 2020-09-25 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="http://uspest.org/risk/tom_pot_map">Google Map of Tomato Potato Blight Daily Risk across NE USA</a></li> <li><a rel="nofollow" class="external text" href="https://www.invasivespeciesinfo.gov/profile/late-blight">Species Profile – Late Blight (Phytophthora infestans)</a>, National Invasive Species Information Center, <a href="/wiki/United_States_National_Agricultural_Library" title="United States National Agricultural Library">United States National Agricultural Library</a>. Lists general information and resources for Late Blight.</li> <li><a rel="nofollow" class="external text" href="http://www.apsnet.org/edcenter/intropp/lessons/fungi/oomycetes/pages/lateblight.aspx">Continuing education lesson</a> created by The <a href="/wiki/American_Phytopathological_Society" title="American Phytopathological Society">American Phytopathological Society</a></li> <li><a rel="nofollow" class="external text" href="https://plantvillage.org/topics/potato/infos">entry on Late Blight</a> by <a href="/wiki/PlantVillage" class="mw-redirect" title="PlantVillage">PlantVillage</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline 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.navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Taxon_identifiers" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Taxon_identifiers" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Taxon_identifiers" title="Help:Taxon identifiers">Taxon identifiers</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;">Phytophthora infestans</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/Wikidata" title="Wikidata">Wikidata</a>: <a href="https://www.wikidata.org/wiki/Q149072" class="extiw" title="wikidata:Q149072">Q149072</a></li> <li><a href="/wiki/Wikispecies" title="Wikispecies">Wikispecies</a>: <a href="https://species.wikimedia.org/wiki/Phytophthora_infestans" class="extiw" title="wikispecies:Phytophthora infestans">Phytophthora infestans</a></li> <li>BioLib: <a rel="nofollow" class="external text" href="https://www.biolib.cz/en/taxon/id61264">61264</a></li> <li><a href="/wiki/Catalogue_of_Life" title="Catalogue of Life">CoL</a>: <a rel="nofollow" class="external text" href="https://www.catalogueoflife.org/data/taxon/4HNSM">4HNSM</a></li> <li><a href="/wiki/Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <a rel="nofollow" class="external text" href="https://eol.org/pages/1004648">1004648</a></li> <li><a href="/wiki/EPPO_Code" title="EPPO Code">EPPO</a>: <a rel="nofollow" class="external text" href="https://gd.eppo.int/taxon/PHYTIN">PHYTIN</a></li> <li><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <a rel="nofollow" class="external text" href="https://www.gbif.org/species/3203716">3203716</a></li> <li><a href="/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/53860">53860</a></li> <li><a href="/wiki/Index_Fungorum" title="Index Fungorum">IndexFungorum</a>: <a rel="nofollow" class="external text" href="http://www.indexfungorum.org/names/NamesRecord.asp?RecordID=232148">232148</a></li> <li><a href="/wiki/Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&id=10873176">10873176</a></li> <li><a href="/wiki/Invasive_Species_Compendium" title="Invasive Species Compendium">ISC</a>: <a rel="nofollow" class="external text" href="https://www.cabi.org/isc/datasheet/40970">40970</a></li> <li><a href="/wiki/Integrated_Taxonomic_Information_System" title="Integrated Taxonomic Information System">ITIS</a>: <a rel="nofollow" class="external text" href="https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=181472">181472</a></li> <li><a href="/wiki/MycoBank" title="MycoBank">MycoBank</a>: <a rel="nofollow" class="external text" href="https://www.mycobank.org/MB/232148">232148</a></li> <li><a href="/wiki/National_Biodiversity_Network" title="National Biodiversity Network">NBN</a>: <a rel="nofollow" class="external text" href="https://data.nbn.org.uk/Taxa/NHMSYS0001493755">NHMSYS0001493755</a></li> <li><a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=4787">4787</a></li> <li>NZOR: <a rel="nofollow" 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