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Zymoseptoria tritici - Wikipedia

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<div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Species of fungus</div> <p class="mw-empty-elt"> </p> <table class="infobox biota" style="text-align: left; width: 200px; font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(145,250,250)"><i>Zymoseptoria tritici</i> </th></tr> <tr> <td colspan="2" style="text-align: center"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Septoria-tritici.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Septoria-tritici.jpg/220px-Septoria-tritici.jpg" decoding="async" width="220" height="160" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Septoria-tritici.jpg/330px-Septoria-tritici.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Septoria-tritici.jpg/440px-Septoria-tritici.jpg 2x" data-file-width="1259" data-file-height="915" /></a></span> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%"><i>Zymoseptoria tritici</i> on leaves of wheat </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(145,250,250)"><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class="plainlinks taxobox-edit-taxonomy skin-invert" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><span typeof="mw:File"><a href="/wiki/Template:Taxonomy/Zymoseptoria" 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></span></span> </th></tr> <tr> <td>Domain: </td> <td><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> </td></tr> <tr> <td>Kingdom: </td> <td><a href="/wiki/Fungus" title="Fungus">Fungi</a> </td></tr> <tr> <td>Division: </td> <td><a href="/wiki/Ascomycota" title="Ascomycota">Ascomycota</a> </td></tr> <tr> <td>Class: </td> <td><a href="/wiki/Dothideomycetes" title="Dothideomycetes">Dothideomycetes</a> </td></tr> <tr> <td>Order: </td> <td><a href="/wiki/Capnodiales" title="Capnodiales">Capnodiales</a> </td></tr> <tr> <td>Family: </td> <td><a href="/wiki/Mycosphaerellaceae" title="Mycosphaerellaceae">Mycosphaerellaceae</a> </td></tr> <tr> <td>Genus: </td> <td><a href="/w/index.php?title=Zymoseptoria&amp;action=edit&amp;redlink=1" class="new" title="Zymoseptoria (page does not exist)"><i>Zymoseptoria</i></a> </td></tr> <tr> <td>Species: </td> <td><div style="display:inline" class="species"><i><b>Z.&#160;tritici</b></i></div> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(145,250,250)"><a href="/wiki/Binomial_nomenclature" title="Binomial nomenclature">Binomial name</a> </th></tr> <tr> <td colspan="2" style="text-align: center"><b><span class="binomial"><span style="font-weight:normal;"></span><i>Zymoseptoria tritici</i></span></b><br /><div style="font-size: 85%;">(Roberge ex Desm.) Quaedvl. &amp; Crous (2011)</div> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(145,250,250)"><a href="/wiki/Synonym_(taxonomy)" title="Synonym (taxonomy)">Synonyms</a> </th></tr> <tr> <td colspan="2" style="text-align: left"> <p><i>Septoria curtisiana</i> <small><a href="/wiki/Pier_Andrea_Saccardo" title="Pier Andrea Saccardo">Sacc.</a>, (1884)</small><sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><br /> <i>Septoria graminum</i> <small>Desm., (1843)</small><br /> <i>Septoria tritici</i> <small><a href="/wiki/John_Baptiste_Henri_Joseph_Desmazi%C3%A8res" title="John Baptiste Henri Joseph Desmazières">Desm.</a>, 1842</small><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> <i>Septoria tritici</i> <small>Berk. &amp; M.A. Curtis, (1874)</small><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup><br /> <i>Septoria tritici var. lolicola</i> <small>R. Sprague &amp; Aar. G. Johnson, (1944)</small><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><br /> <i>Sphaeria graminicola</i> <small>Fuckel, (1865)</small><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup><br /> <i>Sphaerella graminicola</i> <small>Fuckel, (1870)</small><sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup><br /> <i>Mycosphaerella graminicola</i> <small>(<a href="/wiki/Fuckel" class="mw-redirect" title="Fuckel">Fuckel</a>) <a href="/wiki/Joseph_Schr%C3%B6ter" title="Joseph Schröter">J. Schröt.</a>, (1894)</small><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> </p> </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> <p><i><b>Zymoseptoria tritici</b></i>, synonyms <i><b>Septoria tritici</b></i>, <i><b>Mycosphaerella graminicola</b></i>, is a species of <a href="/wiki/Filamentous_fungus" class="mw-redirect" title="Filamentous fungus">filamentous fungus</a>, an <a href="/wiki/Ascomycete" class="mw-redirect" title="Ascomycete">ascomycete</a> in the family <i><a href="/wiki/Mycosphaerellaceae" title="Mycosphaerellaceae">Mycosphaerellaceae</a></i>. It is a <a href="/wiki/Wheat" title="Wheat">wheat</a> plant pathogen causing <b>septoria leaf blotch</b> that is difficult to control due to <a href="/wiki/Drug_resistance" title="Drug resistance">resistance</a> to multiple <a href="/wiki/Fungicide" title="Fungicide">fungicides</a>. The pathogen today causes one of the most important <a href="/wiki/Wheat_diseases" title="Wheat diseases">diseases of wheat</a>.<sup id="cite_ref-Stukenbrock_2010_8-0" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> </p><p>In 2011, Quaedvlieg et al. introduced a <a href="/wiki/New_combination" class="mw-redirect" title="New combination">new combination</a> for this species: <i>Zymoseptoria tritici</i>,<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> as they found that the type strains of both the genus <i>Mycosphaerella</i> (linked to the anamorph genus <i>Ramularia</i>) and the genus <i>Septoria</i> (linked to the genus <i>Septoria</i>, an extensive clade of very distinct septoria-like species within the <i>Mycosphaerellaceae</i>) clustered separately from the clade containing both <i>Zymoseptoria tritici</i> and <i>Z. passerinii</i>. Since 2011, a total of eight <i>Zymoseptoria</i> species have been described within the genus <i>Zymoseptoria</i>; <i>Z. tritici</i> (the type of the genus <i>Zymoseptoria</i>), <i>Z. pseudotritici, Z. ardabiliae, Z. brevis, Z. passerinii, Z. halophila, Z. crescenta</i> and <i>Z. verkleyi</i> (Named after Gerard J.M. Verkleij, for the contribution that he has made to further the understanding of the genus <i>Septoria</i>).<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Description">Description</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Zymoseptoria_tritici&amp;action=edit&amp;section=1" title="Edit section: Description"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Mycosphaerella_graminicola_8.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/6/6b/Mycosphaerella_graminicola_8.png" decoding="async" width="119" height="73" class="mw-file-element" data-file-width="119" data-file-height="73" /></a><figcaption>Ripe pycnidia of <i>Zymoseptoria tritici</i> in a primary leaf of a susceptible wheat seedling. High humidity stimulates the extrusion of cyrrhi, tendril-like mucilages containing asexual pycnidiospores that are rain-splash dispersed over short distances.</figcaption></figure> <p>This fungus causes septoria tritici blotch of wheat, a disease characterized by necrotic blotches on the foliage.<sup id="cite_ref-Wittenberg_2009_11-0" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> These blotches contain asexual (<a href="/wiki/Pycnidia" class="mw-redirect" title="Pycnidia">pycnidia</a>) and sexual (<a href="/wiki/Pseudothecia" class="mw-redirect" title="Pseudothecia">pseudothecia</a>) fructifications.<sup id="cite_ref-Wittenberg_2009_11-1" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p><p>Asexual state (<a href="/wiki/Anamorph" class="mw-redirect" title="Anamorph">anamorph</a>, asexual stage was previously named as <i>Septoria tritici</i>): <a href="/wiki/Pycnidiospore" title="Pycnidiospore">Pycnidiospores</a> are hyaline and threadlike and measure 1.7-3.4 x 39-86 μm, with 3 to 7 indistinct septations. Germiniation of pycnidiospores can be lateral or terminal. Cirrhi are milky white to buff. Sometimes in culture nonseptate, hyaline microspores, measuring 1-1.3 × 5-9 μm, occur outside pycnidia by yeastlike budding.<sup id="cite_ref-WheatCompendium_12-0" class="reference"><a href="#cite_note-WheatCompendium-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> </p> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_2.png" class="mw-file-description" title="Light stimulates yeast-like growth of Zymoseptoria tritici.[13] Close-up of yeast-like growth of Zymoseptoria tritici in vitro on V8 agar."><img alt="Light stimulates yeast-like growth of Zymoseptoria tritici.[13] Close-up of yeast-like growth of Zymoseptoria tritici in vitro on V8 agar." src="//upload.wikimedia.org/wikipedia/commons/e/e7/Mycosphaerella_graminicola_2.png" decoding="async" width="120" height="96" class="mw-file-element" data-file-width="113" data-file-height="90" /></a></span></div> <div class="gallerytext">Light stimulates yeast-like growth of <i>Zymoseptoria tritici</i>.<sup id="cite_ref-Goodwin_2011_13-0" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> Close-up of yeast-like growth of <i>Zymoseptoria tritici</i> <i>in vitro</i> on V8 <a href="/wiki/Agar" title="Agar">agar</a>.</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_3.png" class="mw-file-description" title="In vitro production of asexual fructifications (pycnidia; arrow) of Zymoseptoria tritici on wheat leaf extract agar."><img alt="In vitro production of asexual fructifications (pycnidia; arrow) of Zymoseptoria tritici on wheat leaf extract agar." src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Mycosphaerella_graminicola_3.png/120px-Mycosphaerella_graminicola_3.png" decoding="async" width="120" height="56" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Mycosphaerella_graminicola_3.png/180px-Mycosphaerella_graminicola_3.png 1.5x, //upload.wikimedia.org/wikipedia/commons/7/7f/Mycosphaerella_graminicola_3.png 2x" data-file-width="193" data-file-height="90" /></a></span></div> <div class="gallerytext"><i>In vitro</i> production of asexual fructifications (<a href="/wiki/Pycnidia" class="mw-redirect" title="Pycnidia">pycnidia</a>; arrow) of <i>Zymoseptoria tritici</i> on wheat leaf extract agar.</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_4.png" class="mw-file-description" title="Penetration of a wheat leaf stoma (arrow) by a pycnidiospore germ tube of Zymoseptoria tritici."><img alt="Penetration of a wheat leaf stoma (arrow) by a pycnidiospore germ tube of Zymoseptoria tritici." src="//upload.wikimedia.org/wikipedia/commons/c/ca/Mycosphaerella_graminicola_4.png" decoding="async" width="120" height="66" class="mw-file-element" data-file-width="119" data-file-height="65" /></a></span></div> <div class="gallerytext">Penetration of a wheat leaf stoma (arrow) by a pycnidiospore germ tube of <i>Zymoseptoria tritici</i>.</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_6.png" class="mw-file-description" title="Colonization of the mesophyll tissue by an intercellular hypha (arrows) of Zymoseptoria tritici during the symptomless biotrophic phase of pathogenesis."><img alt="Colonization of the mesophyll tissue by an intercellular hypha (arrows) of Zymoseptoria tritici during the symptomless biotrophic phase of pathogenesis." src="//upload.wikimedia.org/wikipedia/commons/e/ed/Mycosphaerella_graminicola_6.png" decoding="async" width="120" height="76" class="mw-file-element" data-file-width="103" data-file-height="65" /></a></span></div> <div class="gallerytext">Colonization of the mesophyll tissue by an intercellular hypha (arrows) of <i>Zymoseptoria tritici</i> during the symptomless biotrophic phase of pathogenesis.</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_7.png" class="mw-file-description" title="Initiation (arrow head) of a pycnidium of Zymoseptoria tritici in the substomatal cavity of a wheat leaf."><img alt="Initiation (arrow head) of a pycnidium of Zymoseptoria tritici in the substomatal cavity of a wheat leaf." src="//upload.wikimedia.org/wikipedia/commons/a/af/Mycosphaerella_graminicola_7.png" decoding="async" width="120" height="92" class="mw-file-element" data-file-width="85" data-file-height="65" /></a></span></div> <div class="gallerytext">Initiation (arrow head) of a pycnidium of <i>Zymoseptoria tritici</i> in the substomatal cavity of a wheat leaf.</div> </li> </ul> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Mycosphaerella_graminicola_5.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/b/b2/Mycosphaerella_graminicola_5.png" decoding="async" width="113" height="65" class="mw-file-element" data-file-width="113" data-file-height="65" /></a><figcaption>Simultaneous penetration of a wheat leaf stoma by three germ tubes of sexual airborne <a href="/wiki/Ascospores" class="mw-redirect" title="Ascospores">ascospores</a> (arrows) of <i>Zymoseptoria tritici</i>.</figcaption></figure> <p>Sexual state (<a href="/wiki/Teleomorph" class="mw-redirect" title="Teleomorph">teleomorph</a>): <a href="/wiki/Pseudothecia" class="mw-redirect" title="Pseudothecia">Pseudothecia</a> are subepidermal, globose, dark brown, and 68-114 μm in diameter. Asci measure 11-14 × 30-40 μm. <a href="/wiki/Ascospore" class="mw-redirect" title="Ascospore">Ascospores</a> are hyaline, elliptical, and 2.5-4 × 9-16 μm, with two cells of unequal length.<sup id="cite_ref-WheatCompendium_12-1" class="reference"><a href="#cite_note-WheatCompendium-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Genetics">Genetics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Zymoseptoria_tritici&amp;action=edit&amp;section=2" title="Edit section: Genetics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Mycosphaerella_graminicola_chromosomes.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Mycosphaerella_graminicola_chromosomes.png/220px-Mycosphaerella_graminicola_chromosomes.png" decoding="async" width="220" height="98" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Mycosphaerella_graminicola_chromosomes.png/330px-Mycosphaerella_graminicola_chromosomes.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/35/Mycosphaerella_graminicola_chromosomes.png/440px-Mycosphaerella_graminicola_chromosomes.png 2x" data-file-width="663" data-file-height="296" /></a><figcaption>Chromosomes 1-13 are the largest and essential. Chromosomes 14-21 are smaller and dispensable.</figcaption></figure> <p><i>Zymoseptoria tritici</i> represents an intriguing model for fundamental genetic studies of plant-pathogenic fungi.<sup id="cite_ref-Wittenberg_2009_11-2" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> It is <a href="/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a> plant-pathogenic fungus.<sup id="cite_ref-Wittenberg_2009_11-3" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> Many fungi are haploid, which greatly simplifies genetic studies.<sup id="cite_ref-Wittenberg_2009_11-4" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p><p><i>Zymoseptoria tritici</i> was the first species, in 2002, of the family Mycosphaerellaceae to have a <a href="/wiki/Linkage_map" class="mw-redirect" title="Linkage map">linkage map</a> created.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup> </p><p>The first report of fully sequenced <a href="/wiki/Genome" title="Genome">genome</a> of <i>Zymoseptoria tritici</i> from 2011 was the first genome of a <a href="/wiki/Filamentous_fungus" class="mw-redirect" title="Filamentous fungus">filamentous fungus</a> to be finished according to current standards.<sup id="cite_ref-Goodwin_2011_13-1" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> The length of the genome is 39.7 Mb,<sup id="cite_ref-Goodwin_2011_13-2" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> that is similar to other filamentous ascomycetes.<sup id="cite_ref-Wittenberg_2009_11-5" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> The genome contains 21 <a href="/wiki/Chromosomes" class="mw-redirect" title="Chromosomes">chromosomes</a>,<sup id="cite_ref-Goodwin_2011_13-3" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> that is the highest number reported among ascomycetes.<sup id="cite_ref-Wittenberg_2009_11-6" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> Furthermore, these chromosomes have an extraordinary size range, varying from 0.39 to 6.09 Mb.<sup id="cite_ref-Wittenberg_2009_11-7" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p><p>A striking aspect of <i>Zymoseptoria tritici</i> genetics is the presence of many <a href="/w/index.php?title=Dispensable_chromosome&amp;action=edit&amp;redlink=1" class="new" title="Dispensable chromosome (page does not exist)">dispensable chromosomes</a>.<sup id="cite_ref-Goodwin_2011_13-4" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> Eight of chromosomes could be lost with no visible effect on the fungus and thus are dispensable.<sup id="cite_ref-Goodwin_2011_13-5" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> Dispensable chromosomes have been found in other fungi but they usually occur at a low frequency and typically represent single or a few chromosomes.<sup id="cite_ref-Wittenberg_2009_11-8" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> Dispensable chromosomes have originated by ancient <a href="/wiki/Horizontal_transfer" class="mw-redirect" title="Horizontal transfer">horizontal transfer</a> from an unknown donor, that was followed by extensive <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a>, a possible mechanism of stealth <a href="/wiki/Pathogenicity" class="mw-redirect" title="Pathogenicity">pathogenicity</a> and exciting new aspects of genome structure.<sup id="cite_ref-Goodwin_2011_13-6" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>A surprising<sup class="noprint Inline-Template noprint Template-Opinion" style="white-space:nowrap;">&#91;<i><a href="/wiki/Wikipedia:Neutral_point_of_view/FAQ#Assert_facts,_not_opinions" title="Wikipedia:Neutral point of view/FAQ"><span title="This statement may be opinion presented as fact. (July 2024)">opinion</span></a></i>&#93;</sup> feature of the <i>Zymoseptoria tritici</i> genome compared to other sequenced plant pathogens was that it contained very few genes for <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> that break down plant <a href="/wiki/Cell_wall" title="Cell wall">cell walls</a>, which was more similar to <a href="/wiki/Endophyte" title="Endophyte">endophytes</a> than to <a href="/wiki/Pathogen" title="Pathogen">pathogens</a>.<sup id="cite_ref-Goodwin_2011_13-7" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> Goodwin et al. (2011)<sup id="cite_ref-Goodwin_2011_13-8" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> suggested, that the stealth pathogenesis of <i>Zymoseptoria tritici</i> probably involves <a href="/wiki/Proteolysis" title="Proteolysis">degradation of proteins</a> rather than <a href="/wiki/Carbohydrates" class="mw-redirect" title="Carbohydrates">carbohydrates</a> to evade host defenses during the biotrophic stage of infection and may have evolved from endophytic ancestors.<sup id="cite_ref-Goodwin_2011_13-9" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Zymoseptoria_tritici&amp;action=edit&amp;section=3" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The fungus <i>Zymoseptoria tritici</i> has been a pathogen of wheat since host domestication 10,000–12,000 years ago in the <a href="/wiki/Fertile_Crescent" title="Fertile Crescent">Fertile Crescent</a>.<sup id="cite_ref-Stukenbrock_2010_8-1" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> The wheat-infecting lineage emerged from closely related <i>Mycosphaerella</i> pathogens infecting wild <a href="/wiki/Grass" class="mw-redirect" title="Grass">grasses</a>.<sup id="cite_ref-Stukenbrock_2010_8-2" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> It has <a href="/wiki/Coevolved" class="mw-redirect" title="Coevolved">coevolved</a> and spread with its host globally.<sup id="cite_ref-Stukenbrock_2010_8-3" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> <i>Zymoseptoria tritici</i> shows a significantly higher degree of <a href="/wiki/Host_specificity" class="mw-redirect" title="Host specificity">host specificity</a> and virulence in a detached leaf assay.<sup id="cite_ref-Stukenbrock_2010_8-4" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> </p><p>The emergence and "co-domestication" of <i>Zymoseptoria tritici</i> was associated with an <a href="/wiki/Adaptation" title="Adaptation">adaptation</a> to wheat and an agricultural environment.<sup id="cite_ref-Stukenbrock_2010_8-5" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> Endemic descendants of the <a href="/wiki/Progenitor" title="Progenitor">progenitor</a> of <i>Zymoseptoria tritici</i> are still found on wild grasses in the <a href="/wiki/Middle_East" title="Middle East">Middle East</a>; however these "wild" pathogens show a broader host range than the "domesticated" wheat pathogen.<sup id="cite_ref-Stukenbrock_2010_8-6" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> The closest known relative of <i>Zymoseptoria tritici</i> is named <i>Z. pseudotritici</i> B.<sup id="cite_ref-Stukenbrock_2010_8-7" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> <i>Zymoseptoria pseudotritici</i> was isolated in <a href="/wiki/Iran" title="Iran">Iran</a> from the two grass species <i><a href="/wiki/Agropyron_repens" class="mw-redirect" title="Agropyron repens">Agropyron repens</a></i> and <i><a href="/wiki/Dactylis_glomerata" title="Dactylis glomerata">Dactylis glomerata</a></i> growing in close proximity to fields planted to bread wheat (<i><a href="/wiki/Triticum_aestivum" class="mw-redirect" title="Triticum aestivum">Triticum aestivum</a></i>).<sup id="cite_ref-Stukenbrock_2010_8-8" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> Although <i>Z. tritici</i> is a frequent pathogen of wheat in Iran, no evidence of <a href="/wiki/Gene_flow" title="Gene flow">gene flow</a> between <i>Z. pseudotritici</i> and <i>Z. tritici</i> was detected based on sequence analysis of six nuclear loci.<sup id="cite_ref-Stukenbrock_2010_8-9" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Life_cycle">Life cycle</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Zymoseptoria_tritici&amp;action=edit&amp;section=4" title="Edit section: Life cycle"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i>Zymoseptoria tritici</i> overwinters as fruiting bodies on crop debris, mostly as pseudothecia (sexual fruiting bodies) but sometimes also some pycnidia (asexual fruiting bodies).<sup id="cite_ref-:0_15-0" class="reference"><a href="#cite_note-:0-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> The sexual spores are quantitatively the more significant spores involved in primary inoculum of the disease, while the asexual spores are more significant in the secondary cycle.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup> In early spring, <a href="/wiki/Ascospore" class="mw-redirect" title="Ascospore">ascospores</a>, the sexual spores of the fungus, are released from the pseudothecia. Ascospores are wind-dispersed and eventually land on the leaves of a host plant (bread wheat or durum wheat). Unlike most other plant pathogens, <i>Zymoseptoria tritici</i> uses a germ tube to enter the host leaf through <a href="/wiki/Stoma" title="Stoma">stomata</a> rather than by direct penetration.<sup id="cite_ref-:1_17-0" class="reference"><a href="#cite_note-:1-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> There is a long <a href="/wiki/Incubation_period" title="Incubation period">latent period</a> of up to two weeks following infection before symptoms develop.<sup id="cite_ref-Goodwin_2011_13-10" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> The fungus evades host defenses during the latent phase, followed by a rapid switch to necrotrophy immediately prior to symptom expression 12–20 days after penetration.<sup id="cite_ref-Goodwin_2011_13-11" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> The period between infection and formation of sporulating structures (latent period) was estimated to be 20.35 ± 4.15 days for <i>Zymoseptoria tritici</i> in Northern Germany and decreased with increasing temperature.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> Such a switch from <a href="/wiki/Biotrophic" class="mw-redirect" title="Biotrophic">biotrophic</a> to <a href="/wiki/Necrotrophic" class="mw-redirect" title="Necrotrophic">necrotrophic</a> growth at the end of a long latent period is an unusual characteristic shared by most fungi in the genus <i>Mycosphaerella</i>.<sup id="cite_ref-Goodwin_2011_13-12" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> Very little is known about the cause or mechanism of this lifestyle switch even though <i>Mycosphaerella</i> is one of the largest and most economically important genera of plant-pathogenic fungi.<sup id="cite_ref-Goodwin_2011_13-13" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>Primary inoculum requires wet conditions and cool temperatures of 50-68&#160;°F.<sup id="cite_ref-:2_19-0" class="reference"><a href="#cite_note-:2-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> Under appropriate environmental conditions, lesions are able to develop on infected leaves, and soon pycnidia begin to develop on the lesions.<sup id="cite_ref-:2_19-1" class="reference"><a href="#cite_note-:2-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> The pycnidia appear as small dark dots on the lesions. From the pycnidia, conidiospores, the asexual spores of the fungus, are released. These asexual spores are dispersed via rain splash and are response for the secondary inoculum of this polycyclic disease cycle.<sup id="cite_ref-:1_17-1" class="reference"><a href="#cite_note-:1-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> When the <a href="/wiki/Conidium" title="Conidium">conidiospores</a> are splashed onto leaves, they act similarly to ascospores and cause the development of foliar lesions. In addition to pycnidia, pseudothecia also develop within these lesions. Pycnidia and pseudothecia are the structures in which the fungus overwinters, and the cycle begins again.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">&#91;<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (June 2024)">citation needed</span></a></i>&#93;</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Disease_Management">Disease Management</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Zymoseptoria_tritici&amp;action=edit&amp;section=5" title="Edit section: Disease Management"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i>Zymoseptoria tritici</i> is a difficult fungus to <a href="/wiki/Pest_control" title="Pest control">control</a> because populations contain extremely high levels of <a href="/wiki/Genetic_variability" title="Genetic variability">genetic variability</a> and it has very unusual biology for a pathogen.<sup id="cite_ref-Goodwin_2011_13-14" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> <i>Z. tritici</i> has an active sexual cycle under natural conditions, which is an important driver of septoria tritici blotch <a href="/wiki/Epidemics" class="mw-redirect" title="Epidemics">epidemics</a> and results in high genetic diversity of populations in the field.<sup id="cite_ref-Wittenberg_2009_11-9" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p><p>The most effective, economical, and simple method of <i>Z. tritici</i> management is planting <a href="/wiki/Resistant_cultivar" class="mw-redirect" title="Resistant cultivar">resistant cultivars</a>. Twenty-one resistant genes have been named, mapped, and published.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> Mikaberidze and McDonald 2020 found a <a href="/wiki/Biological_fitness" class="mw-redirect" title="Biological fitness">fitness</a> tradeoff between genes for <i>Septoria</i> <a href="/wiki/Disease_tolerance" class="mw-redirect" title="Disease tolerance">tolerance</a> and <i>Septoria</i> resistance in wheat.<sup id="cite_ref-Pagan-Garcia-Arenal-2020_21-0" class="reference"><a href="#cite_note-Pagan-Garcia-Arenal-2020-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> Some cultivars are resistant in one region but susceptible in another; it depends on the local pathogen population. All varieties of bread wheat and durum wheat are susceptible to the disease to some extent, but planting varieties that have at least partial resistance to the local population of <i>Zymoseptoria tritici</i> can greatly improve yield. </p><p>There are also cultural management strategies that may be effective, including regular rotation of crops, deep plowing, and late planting.<sup id="cite_ref-:0_15-1" class="reference"><a href="#cite_note-:0-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> More specifically, rotating a recently infected field to any non-host crop can be useful in minimizing the amount of fungus present in the field. Planting winter wheat after the first ascospore flights in September is a way to reduce primary inoculum of winter wheat.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> </p><p>Fungicide use often simply is not economical for Septoria Leaf Blotch. The rapid evolution of pathogen resistance to fungicides is a major barrier. <i>Zymoseptoria tritici</i> has <a href="/wiki/Drug_resistance" title="Drug resistance">resistance</a> to multiple fungicides, because it has number of <a href="/wiki/Substitution_reaction" title="Substitution reaction">substitutions</a> of CYP51. CYP51 substitutions include <a href="/w/index.php?title=Y137F&amp;action=edit&amp;redlink=1" class="new" title="Y137F (page does not exist)">Y137F</a> which confers resistance to <a href="/w/index.php?title=Triadimenol&amp;action=edit&amp;redlink=1" class="new" title="Triadimenol (page does not exist)">triadimenol</a>, <a href="/w/index.php?title=I381V&amp;action=edit&amp;redlink=1" class="new" title="I381V (page does not exist)">I381V</a> which confers resistance to <a href="/wiki/Tebuconazole" title="Tebuconazole">tebuconazole</a> and <a href="/w/index.php?title=V136A&amp;action=edit&amp;redlink=1" class="new" title="V136A (page does not exist)">V136A</a> that confers resistance to <a href="/wiki/Prochloraz" title="Prochloraz">prochloraz</a>.<sup id="cite_ref-Mullins_2011_23-0" class="reference"><a href="#cite_note-Mullins_2011-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> Chemical control of the pathogen (using <a href="/wiki/Fungicide" title="Fungicide">fungicides</a>) now relies on the application of SDHIs,<sup id="cite_ref-Downie-et-al-2021_24-0" class="reference"><a href="#cite_note-Downie-et-al-2021-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> <a href="/wiki/Azole" title="Azole">azole</a> fungicides which are <a href="/wiki/Demethylase" title="Demethylase">demethylase</a> inhibitors that inhibit <a href="/wiki/Lanosterol_14_alpha-demethylase" title="Lanosterol 14 alpha-demethylase">lanosterol 14 alpha-demethylase</a> (<a href="/wiki/CYP51" class="mw-redirect" title="CYP51">CYP51</a>) activity.<sup id="cite_ref-Mullins_2011_23-1" class="reference"><a href="#cite_note-Mullins_2011-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> </p><p>The last method of control for <i>Zymoseptoria tritici</i> is biological control using bacteria. <i><a href="/wiki/Bacillus_megaterium" class="mw-redirect" title="Bacillus megaterium">Bacillus megaterium</a></i> has been shown to cause about an 80% decrease in disease development in the trials done so far.<sup id="cite_ref-:1_17-2" class="reference"><a href="#cite_note-:1-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> <a href="/wiki/Pseudomonadaceae" title="Pseudomonadaceae">Pseudomonads</a> are also a promising bacterial control option. A benefit to using pseudomonads or bacillus is that they are not harmed by most fungicides, so they can be used in combination with chemical controls.<sup id="cite_ref-:1_17-3" class="reference"><a href="#cite_note-:1-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> However, a lack of commercial availability limits the use of biological controls.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">&#91;<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (June 2024)">citation needed</span></a></i>&#93;</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Disease_Importance">Disease Importance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Zymoseptoria_tritici&amp;action=edit&amp;section=6" title="Edit section: Disease Importance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ascomycete fungus <i>Zymoseptoria tritici</i> causes <b>septoria tritici blotch</b>, a foliar disease of <a href="/wiki/Wheat" title="Wheat">wheat</a> that poses a significant threat to global <a href="/wiki/Food_production" class="mw-redirect" title="Food production">food production</a>.<sup id="cite_ref-Goodwin_2011_13-15" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> It is the primary foliar disease of <a href="/wiki/Winter_wheat" title="Winter wheat">winter wheat</a> in most western European countries.<sup id="cite_ref-Mullins_2011_23-2" class="reference"><a href="#cite_note-Mullins_2011-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> <i>Zymoseptoria tritici</i> infects wheat crops throughout the world and is also currently a big problem in Iran, Tunisia, and Morocco.<sup id="cite_ref-:1_17-4" class="reference"><a href="#cite_note-:1-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> Severe epidemics of the disease have decreased wheat yields by 35-50%.<sup id="cite_ref-:1_17-5" class="reference"><a href="#cite_note-:1-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> In the United States, Septoria leaf blotch is a very important disease in wheat, second only to <a href="/wiki/Wheat_leaf_rust" title="Wheat leaf rust">wheat rust</a>. An estimated $275 million is lost per year in the US due to this disease. In Europe the annual losses are equivalent to over 400 million USD.<sup id="cite_ref-:1_17-6" class="reference"><a href="#cite_note-:1-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> </p><p>Different areas of the world are currently trying different management strategies. For example, in the Nordic-Baltic region, one of the largest wheat-producing regions of the world, the use of fungicides has substantially increased wheat yields.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> The fungicides that have been shown to be effective include <a href="/wiki/QoI" title="QoI">quinone outside inhibitors</a> (QoIs), which, like most fungicides, are expensive to apply in large quantities. As <a href="/wiki/Climate_change" title="Climate change">climate change</a> begins to increase temperatures around the globe, <i>Zymoseptoria tritici</i>, along with many other fungal pathogens, is likely to show increased overwintering survival and therefore more substantial primary inocula.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> The need for effective management techniques will become even more important as the prevalence of Septoria leaf blotch increases with climate change.<sup id="cite_ref-Fones_27-0" class="reference"><a href="#cite_note-Fones-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup> </p> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_9.png" class="mw-file-description" title="Typical infection caused by Zymoseptoria tritici of the primary leaf of a resistant cultivar. Note the low fungal density in the apoplast (arrow) and the response of the mesophyll cells (arrow head), particularly the chloroplasts, to the presence of intercellular hyphae."><img alt="Typical infection caused by Zymoseptoria tritici of the primary leaf of a resistant cultivar. Note the low fungal density in the apoplast (arrow) and the response of the mesophyll cells (arrow head), particularly the chloroplasts, to the presence of intercellular hyphae." src="//upload.wikimedia.org/wikipedia/commons/b/b6/Mycosphaerella_graminicola_9.png" decoding="async" width="120" height="78" class="mw-file-element" data-file-width="113" data-file-height="73" /></a></span></div> <div class="gallerytext">Typical infection caused by <i>Zymoseptoria tritici</i> of the primary leaf of a resistant <a href="/wiki/Cultivar" title="Cultivar">cultivar</a>. Note the low fungal density in the <a href="/wiki/Apoplast" title="Apoplast">apoplast</a> (arrow) and the response of the <a href="/wiki/Mesophyll_tissue" class="mw-redirect" title="Mesophyll tissue">mesophyll</a> cells (arrow head), particularly the chloroplasts, to the presence of intercellular <a href="/wiki/Hyphae" class="mw-redirect" title="Hyphae">hyphae</a>.</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_10.png" class="mw-file-description" title="(upper image) Typical symptoms of Zymoseptoria tritici on a primary seedling leaf of a highly susceptible wheat cultivar. (lower image) Typical response to Zymoseptoria tritici on a primary leaf of a highly resistant wheat cultivar."><img alt="(upper image) Typical symptoms of Zymoseptoria tritici on a primary seedling leaf of a highly susceptible wheat cultivar. (lower image) Typical response to Zymoseptoria tritici on a primary leaf of a highly resistant wheat cultivar." src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3d/Mycosphaerella_graminicola_10.png/120px-Mycosphaerella_graminicola_10.png" decoding="async" width="120" height="52" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/3/3d/Mycosphaerella_graminicola_10.png 1.5x" data-file-width="168" data-file-height="73" /></a></span></div> <div class="gallerytext">(upper image) Typical symptoms of <i>Zymoseptoria tritici</i> on a primary seedling leaf of a highly susceptible wheat cultivar. (lower image) Typical response to <i>Zymoseptoria tritici</i> on a primary leaf of a highly resistant wheat cultivar.</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mycosphaerella_graminicola_14.png" class="mw-file-description" title="Symptoms of Zymoseptoria tritici on a naturally infected adult plant flag leaf of wheat."><img alt="Symptoms of Zymoseptoria tritici on a naturally infected adult plant flag leaf of wheat." src="//upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Mycosphaerella_graminicola_14.png/120px-Mycosphaerella_graminicola_14.png" decoding="async" width="120" height="52" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/e/eb/Mycosphaerella_graminicola_14.png 1.5x" data-file-width="168" data-file-height="73" /></a></span></div> <div class="gallerytext">Symptoms of <i>Zymoseptoria tritici</i> on a naturally infected adult plant flag leaf of wheat.</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Zymoseptoria_tritici&amp;action=edit&amp;section=7" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>This article incorporates CC-BY-2.5 text from references<sup id="cite_ref-Stukenbrock_2010_8-10" class="reference"><a href="#cite_note-Stukenbrock_2010-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Wittenberg_2009_11-10" class="reference"><a href="#cite_note-Wittenberg_2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Goodwin_2011_13-16" class="reference"><a href="#cite_note-Goodwin_2011-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Mullins_2011_23-3" class="reference"><a href="#cite_note-Mullins_2011-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><a href="/wiki/Pier_Andrea_Saccardo" title="Pier Andrea Saccardo">Saccardo P. A.</a> (1884). <i>Syll. fung.</i> (Abellini) <b>3</b>: 561.</span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><a href="/wiki/John_Baptiste_Henri_Joseph_Desmazi%C3%A8res" title="John Baptiste Henri Joseph Desmazières">Desmazières J. B. H. J.</a> (1842). "Neuvième notice sur quelques plantes cryptogames, la plupart inédites, récemment découvertes en France, et que vont paraître en nature dans la collection publiée par l’auteur". <i><a href="/w/index.php?title=Annales_Des_Sciences_Naturelles&amp;action=edit&amp;redlink=1" class="new" title="Annales Des Sciences Naturelles (page does not exist)">Annales Des Sciences Naturelles</a>, Bot.</i>, sér. 2, <b>17</b>: <a rel="nofollow" class="external text" href="https://archive.org/stream/annalesdesscienc17cram#page/91/mode/1up">91</a>-118. <a rel="nofollow" class="external text" href="https://archive.org/stream/annalesdesscienc17cram#page/106/mode/2up">page 107</a>.</span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">Berk. &amp; Curtis M. A. (1874). <i>N. Amer. Fung.</i>: no. 441 bis.</span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text">Sprague R. &amp; Johnson A. G. (1944). In: Sprague, <i>Ore. St. Monog., Bot.</i> <b>6</b>: 32.</span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text">Fuckel (1865). <i>Fungi rhenani exsic.</i>: no. 1578.</span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text">Fuckel (1870). <i>Jb. nassau. Ver. Naturk.</i> <b>23-24</b>: 101.</span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><a href="/wiki/Joseph_Schr%C3%B6ter" title="Joseph Schröter">Schröter J.</a> (1894). 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font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Mycosphaerella_graminicola" class="extiw" title="commons:Category:Mycosphaerella graminicola">Mycosphaerella graminicola</a></span>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070820101227/http://nt.ars-grin.gov/fungaldatabases/">USDA ARS Fungal Database</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFvan_GinkelA._McNabJ._Krupinsky1999" class="citation book cs1">van Ginkel, M.; A. McNab; J. Krupinsky (1999). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20031029075604/http://www.cimmyt.org/Research/Wheat/pdf/Septoria/septoria.pdf"><i>Septoria and stagonospora diseases of cereals: A compilation of global research</i></a> <span class="cs1-format">(PDF)</span>. CIMMYT. p.&#160;186pp. Archived from <a rel="nofollow" class="external text" href="http://www.cimmyt.org/Research/Wheat/pdf/Septoria/septoria.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 2003-10-29.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Septoria+and+stagonospora+diseases+of+cereals%3A+A+compilation+of+global+research&amp;rft.pages=186pp&amp;rft.pub=CIMMYT&amp;rft.date=1999&amp;rft.aulast=van+Ginkel&amp;rft.aufirst=M.&amp;rft.au=A.+McNab&amp;rft.au=J.+Krupinsky&amp;rft_id=http%3A%2F%2Fwww.cimmyt.org%2FResearch%2FWheat%2Fpdf%2FSeptoria%2Fseptoria.pdf&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AZymoseptoria+tritici" class="Z3988"></span></li></ul> <ul><li>Orton E. S., Sian Deller S. &amp; Brown J. K. M. (2011). "<i>Mycosphaerella graminicola</i>: from genomics to disease control". <i><a href="/wiki/Molecular_Plant_Pathology" title="Molecular Plant Pathology">Molecular Plant Pathology</a></i> <b>12</b>(5): 413-424. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1364-3703.2010.00688.x">10.1111/j.1364-3703.2010.00688.x</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 ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl 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.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_identifiers5166" 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_identifiers5166" 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;"><i>Zymoseptoria tritici</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><span style="white-space:nowrap;"><a href="/wiki/Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q104837794" class="extiw" title="wikidata:Q104837794">Q104837794</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Catalogue_of_Life" title="Catalogue of Life">CoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.catalogueoflife.org/data/taxon/5DCF9">5DCF9</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/EPPO_Code" title="EPPO Code">EPPO</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://gd.eppo.int/taxon/SEPTTR">SEPTTR</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.gbif.org/species/7897674">7897674</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/1096267">1096267</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Index_Fungorum" title="Index Fungorum">IndexFungorum</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.indexfungorum.org/names/NamesRecord.asp?RecordID=517926">517926</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/MycoBank" title="MycoBank">MycoBank</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.mycobank.org/MB/517926">517926</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&amp;id=1047171">1047171</a></span></span></li> <li><span style="white-space:nowrap;">NZOR: <span class="uid"><a rel="nofollow" class="external text" href="https://www.nzor.org.nz/names/2b1ddbe0-c4cd-4d46-9d24-ea13bc9e954d">2b1ddbe0-c4cd-4d46-9d24-ea13bc9e954d</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Open_Tree_of_Life" title="Open Tree of Life">Open Tree of Life</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://tree.opentreeoflife.org/taxonomy/browse?id=906425">906425</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Index_Fungorum" title="Index Fungorum">SpeciesFungorum</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.speciesfungorum.org/Names/NamesRecord.asp?RecordID=517926">517926</a></span></span></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;"><i>Mycosphaerella graminicola</i></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><span style="white-space:nowrap;"><a href="/wiki/Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q138622" class="extiw" title="wikidata:Q138622">Q138622</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://eol.org/pages/192235">192235</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.gbif.org/species/2622594">2622594</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/383238">383238</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Index_Fungorum" title="Index Fungorum">IndexFungorum</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.indexfungorum.org/names/NamesRecord.asp?RecordID=318177">318177</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&amp;id=10646219">10646219</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/MycoBank" title="MycoBank">MycoBank</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.mycobank.org/MB/318177">318177</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/NatureServe" title="NatureServe">NatureServe</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://explorer.natureserve.org/Taxon/ELEMENT_GLOBAL.2.1157071/">2.1157071</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/National_Biodiversity_Network" title="National Biodiversity Network">NBN</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://species.nbnatlas.org/species/BMSSYS0000011814">BMSSYS0000011814</a></span></span></li> <li><span style="white-space:nowrap;">NZOR: <span class="uid"><a rel="nofollow" class="external text" href="https://www.nzor.org.nz/names/d907e3b0-2cf8-4d99-b0d5-169d82826933">d907e3b0-2cf8-4d99-b0d5-169d82826933</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Open_Tree_of_Life" title="Open Tree of Life">Open Tree of Life</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://tree.opentreeoflife.org/taxonomy/browse?id=5319573">5319573</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Index_Fungorum" title="Index Fungorum">SpeciesFungorum</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.speciesfungorum.org/Names/NamesRecord.asp?RecordID=318177">318177</a></span></span></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;"><i>Septoria tritici</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><span style="white-space:nowrap;"><a href="/wiki/Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q103817219" class="extiw" title="wikidata:Q103817219">Q103817219</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.gbif.org/species/9624429">9624429</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/384314">384314</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Index_Fungorum" title="Index Fungorum">IndexFungorum</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.indexfungorum.org/names/NamesRecord.asp?RecordID=205691">205691</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&amp;id=10502706">10502706</a></span></span></li> <li><span style="white-space:nowrap;">NZOR: <span class="uid"><a rel="nofollow" class="external text" href="https://www.nzor.org.nz/names/ab280441-b0b6-4145-ba06-570623bb3478">ab280441-b0b6-4145-ba06-570623bb3478</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Open_Tree_of_Life" title="Open Tree of Life">Open Tree of Life</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://tree.opentreeoflife.org/taxonomy/browse?id=5319978">5319978</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Index_Fungorum" title="Index Fungorum">SpeciesFungorum</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.speciesfungorum.org/Names/NamesRecord.asp?RecordID=205691">205691</a></span></span></li></ul> </div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐api‐ext.eqiad.main‐84b6579fc7‐fvr9p Cached time: 20250313093042 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 0.735 seconds Real time usage: 1.053 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