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class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#By_insects"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>By insects</span> </div> </a> <ul id="toc-By_insects-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-By_nematodes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#By_nematodes"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>By nematodes</span> </div> </a> <ul id="toc-By_nematodes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-By_plasmodiophorids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#By_plasmodiophorids"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>By plasmodiophorids</span> </div> </a> <ul id="toc-By_plasmodiophorids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-On_seed_and_pollen" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#On_seed_and_pollen"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>On seed and pollen</span> </div> </a> <ul id="toc-On_seed_and_pollen-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Directly_from_plant_to_humans" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Directly_from_plant_to_humans"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.6</span> <span>Directly from plant to humans</span> </div> </a> <ul id="toc-Directly_from_plant_to_humans-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Effects_on_hosts" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Effects_on_hosts"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Effects on hosts</span> </div> </a> <ul id="toc-Effects_on_hosts-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Translation_of_plant_viral_proteins" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Translation_of_plant_viral_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Translation of plant viral proteins</span> </div> </a> <button aria-controls="toc-Translation_of_plant_viral_proteins-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Translation of plant viral proteins subsection</span> </button> <ul id="toc-Translation_of_plant_viral_proteins-sublist" class="vector-toc-list"> <li id="toc-5'_Cap" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#5'_Cap"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>5' Cap</span> </div> </a> <ul id="toc-5'_Cap-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Readthrough" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Readthrough"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Readthrough</span> </div> </a> <ul id="toc-Readthrough-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Production_of_sub-genomic_RNAs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Production_of_sub-genomic_RNAs"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>Production of sub-genomic RNAs</span> </div> </a> <ul id="toc-Production_of_sub-genomic_RNAs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Segmented_genomes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Segmented_genomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.4</span> <span>Segmented genomes</span> </div> </a> <ul id="toc-Segmented_genomes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Polyprotein_processing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Polyprotein_processing"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.5</span> <span>Polyprotein processing</span> </div> </a> <ul id="toc-Polyprotein_processing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genome_packaging" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genome_packaging"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.6</span> <span>Genome packaging</span> </div> </a> <ul id="toc-Genome_packaging-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Applications_of_plant_viruses" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Applications_of_plant_viruses"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Applications of plant viruses</span> </div> </a> <ul id="toc-Applications_of_plant_viruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <button aria-controls="toc-References-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle References subsection</span> </button> <ul id="toc-References-sublist" class="vector-toc-list"> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > 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Available in 17 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-17" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">17 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%81%D9%8A%D8%B1%D9%88%D8%B3_%D9%86%D8%A8%D8%A7%D8%AA" title="فيروس نبات – Arabic" lang="ar" hreflang="ar" data-title="فيروس نبات" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%89%E0%A6%A6%E0%A7%8D%E0%A6%AD%E0%A6%BF%E0%A6%A6_%E0%A6%AD%E0%A6%BE%E0%A6%87%E0%A6%B0%E0%A6%BE%E0%A6%B8" title="উদ্ভিদ ভাইরাস – Bangla" lang="bn" hreflang="bn" data-title="উদ্ভিদ ভাইরাস" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Virus_de_les_plantes" title="Virus de les plantes – Catalan" lang="ca" hreflang="ca" data-title="Virus de les plantes" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Fytovirus" title="Fytovirus – Czech" lang="cs" hreflang="cs" data-title="Fytovirus" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Pflanzenvirus" title="Pflanzenvirus – German" lang="de" hreflang="de" data-title="Pflanzenvirus" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Fitovirus" title="Fitovirus – Spanish" lang="es" hreflang="es" data-title="Fitovirus" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%88%DB%8C%D8%B1%D9%88%D8%B3_%DA%AF%DB%8C%D8%A7%D9%87%DB%8C" title="ویروس گیاهی – Persian" lang="fa" hreflang="fa" data-title="ویروس گیاهی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Phytovirus" title="Phytovirus – French" lang="fr" hreflang="fr" data-title="Phytovirus" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%AA%E0%A4%BE%E0%A4%A6%E0%A4%AA_%E0%A4%B5%E0%A4%BE%E0%A4%87%E0%A4%B0%E0%A4%B8" title="पादप वाइरस – Hindi" lang="hi" hreflang="hi" data-title="पादप वाइरस" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Virus_tumbuhan" title="Virus tumbuhan – Indonesian" lang="id" hreflang="id" data-title="Virus tumbuhan" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B8%E0%B4%B8%E0%B5%8D%E0%B4%AF%E0%B4%B5%E0%B5%88%E0%B4%B1%E0%B4%B8%E0%B5%8D" title="സസ്യവൈറസ് – Malayalam" lang="ml" hreflang="ml" data-title="സസ്യവൈറസ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Plantenvirus" title="Plantenvirus – Dutch" lang="nl" hreflang="nl" data-title="Plantenvirus" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E6%A4%8D%E7%89%A9%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="植物ウイルス – Japanese" lang="ja" hreflang="ja" data-title="植物ウイルス" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D9%86%D8%A8%D8%A7%D8%AA%D9%8A_%D9%88%D9%8A%D8%B1%D9%88%D8%B3%D9%88%D9%86%D9%87" title="نباتي ويروسونه – Pashto" lang="ps" hreflang="ps" data-title="نباتي ويروسونه" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Wirusy_ro%C5%9Blinne" title="Wirusy roślinne – Polish" lang="pl" hreflang="pl" data-title="Wirusy roślinne" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Fitov%C3%ADrus" title="Fitovírus – Portuguese" lang="pt" hreflang="pt" data-title="Fitovírus" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E6%A4%8D%E7%89%A9%E7%97%85%E6%AF%92" title="植物病毒 – Chinese" lang="zh" hreflang="zh" data-title="植物病毒" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q1495434#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div 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//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Plant_virus" title="Special:EditPage/Plant virus">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Plant+virus%22">"Plant virus"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Plant+virus%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Plant+virus%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Plant+virus%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Plant+virus%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Plant+virus%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">December 2023</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Pepper_mild_mottle_virus.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Pepper_mild_mottle_virus.png/220px-Pepper_mild_mottle_virus.png" decoding="async" width="220" height="191" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Pepper_mild_mottle_virus.png/330px-Pepper_mild_mottle_virus.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Pepper_mild_mottle_virus.png/440px-Pepper_mild_mottle_virus.png 2x" data-file-width="650" data-file-height="565" /></a><figcaption><i><a href="/wiki/Pepper_mild_mottle_virus" title="Pepper mild mottle virus">Pepper mild mottle virus</a></i></figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:SequoiaBio_Leaf_Curl.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/SequoiaBio_Leaf_Curl.jpg/220px-SequoiaBio_Leaf_Curl.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/SequoiaBio_Leaf_Curl.jpg/330px-SequoiaBio_Leaf_Curl.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fc/SequoiaBio_Leaf_Curl.jpg/440px-SequoiaBio_Leaf_Curl.jpg 2x" data-file-width="640" data-file-height="480" /></a><figcaption>Leaf curl virus</figcaption></figure> <p><b>Plant viruses</b> are <a href="/wiki/Virus" title="Virus">viruses</a> that have the potential to affect <a href="/wiki/Plant" title="Plant">plants</a>. Like all other viruses, plant viruses are obligate <a href="/wiki/Intracellular_parasite" title="Intracellular parasite">intracellular parasites</a> that do not have the molecular machinery to <a href="/wiki/Self-replication" title="Self-replication">replicate</a> without a <a href="/wiki/Host_(biology)" title="Host (biology)">host</a>. Plant viruses can be <a href="/wiki/Pathogen" title="Pathogen">pathogenic</a> to <a href="/wiki/Vascular_plant" title="Vascular plant">vascular plants ("higher plants")</a>. </p><p>Many plant viruses are <a href="/wiki/Rod-shaped" class="mw-redirect" title="Rod-shaped">rod-shaped</a>, with protein discs forming a tube surrounding the viral <a href="/wiki/Genome" title="Genome">genome</a>; isometric particles are another common structure. They rarely have an <a href="/wiki/Viral_envelope" title="Viral envelope">envelope</a>. The great majority have an RNA genome, which is usually small and single stranded (ss), but some viruses have double-stranded (ds) RNA, ssDNA or dsDNA genomes. Although plant viruses are not as well understood as their animal counterparts, one plant virus has become very recognizable: <i><a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">tobacco mosaic virus</a></i> (TMV), the first virus to be discovered. This and other viruses cause an estimated US$60 billion loss in crop yields worldwide each year. Plant viruses are grouped into 73 <a href="/wiki/Genus_(biology)" class="mw-redirect" title="Genus (biology)">genera</a> and 49 <a href="/wiki/Family_(biology)" title="Family (biology)">families</a>. However, these figures relate only to cultivated plants, which represent only a tiny fraction of the total number of plant species. Viruses in wild plants have not been well-studied, but the interactions between wild plants and their viruses often do not appear to cause disease in the host plants.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>To transmit from one plant to another and from one plant cell to another, plant viruses must use strategies that are usually different from <a href="/wiki/Animal_virus" title="Animal virus">animal viruses</a>. Most plants do not move, and so plant-to-plant transmission usually involves vectors (such as insects). Plant cells are surrounded by solid <a href="/wiki/Cell_wall" title="Cell wall">cell walls</a>, therefore transport through <a href="/wiki/Plasmodesmata" class="mw-redirect" title="Plasmodesmata">plasmodesmata</a> is the preferred path for virions to move between plant cells. Plants have specialized mechanisms for transporting <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNAs</a> through plasmodesmata, and these mechanisms are thought to be used by <a href="/wiki/RNA_virus" title="RNA virus">RNA viruses</a> to spread from one cell to another.<sup id="cite_ref-Plasmodesmata_2-0" class="reference"><a href="#cite_note-Plasmodesmata-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Innate_immune_system#Plants" title="Innate immune system">Plant defenses against viral infection</a> include, among other measures, the use of <a href="/wiki/SiRNA" class="mw-redirect" title="SiRNA">siRNA</a> in response to <a href="/wiki/DsRNA" class="mw-redirect" title="DsRNA">dsRNA</a>.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Most plant viruses encode a protein to suppress this response.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Plants also reduce transport through <a href="/wiki/Plasmodesmata" class="mw-redirect" title="Plasmodesmata">plasmodesmata</a> in response to injury.<sup id="cite_ref-Plasmodesmata_2-1" class="reference"><a href="#cite_note-Plasmodesmata-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:TobaccoMosaicVirus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/TobaccoMosaicVirus.jpg/170px-TobaccoMosaicVirus.jpg" decoding="async" width="170" height="219" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/TobaccoMosaicVirus.jpg/255px-TobaccoMosaicVirus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/TobaccoMosaicVirus.jpg/340px-TobaccoMosaicVirus.jpg 2x" data-file-width="900" data-file-height="1158" /></a><figcaption><a href="/wiki/Electron_microscope" title="Electron microscope">Electron micrograph</a> of the rod-shaped particles of <a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">tobacco mosaic virus</a></figcaption></figure> <p>The discovery of plant viruses causing <a href="/wiki/Disease" title="Disease">disease</a> is often accredited to A. Mayer (1886) working in the Netherlands demonstrated that the sap of mosaic obtained from tobacco leaves developed mosaic symptom when injected in healthy plants. However the infection of the sap was destroyed when it was boiled. He thought that the causal agent was bacteria. However, after larger inoculation with a large number of bacteria, he failed to develop a mosaic symptom. </p><p>In 1898, Martinus Beijerinck, who was a professor of microbiology at the Technical University the Netherlands, put forth his concepts that viruses were small and determined that the "mosaic disease" remained infectious when passed through a <a href="/wiki/Chamberland_filter" title="Chamberland filter">Chamberland filter-candle</a>. This was in contrast to bacteria <a href="/wiki/Microorganisms" class="mw-redirect" title="Microorganisms">microorganisms</a>, which were retained by the filter. <a href="/wiki/Beijerinck" class="mw-redirect" title="Beijerinck">Beijerinck</a> referred to the infectious filtrate as a "<a href="/wiki/Contagium_vivum_fluidum" title="Contagium vivum fluidum">contagium vivum fluidum</a>", thus the coinage of the modern term "virus". </p><p>After the initial discovery of the 'viral concept' there was need to classify any other known <a href="/wiki/Virus" title="Virus">viral</a> diseases based on the mode of transmission even though <a href="/wiki/Microscopic" class="mw-redirect" title="Microscopic">microscopic</a> observation proved fruitless. In 1939 Holmes published a classification list of 129 plant viruses. This was expanded and in 1999 there were 977 officially recognized, and some provisional, plant virus species. </p><p>The purification (crystallization) of TMV was first performed by <a href="/wiki/Wendell_Stanley" class="mw-redirect" title="Wendell Stanley">Wendell Stanley</a>, who published his findings in 1935, although he did not determine that the RNA was the infectious material. However, he received the <a href="/wiki/Nobel_Prize" title="Nobel Prize">Nobel Prize</a> in Chemistry in 1946. In the 1950s a discovery by two labs simultaneously proved that the purified <a href="/wiki/RNA" title="RNA">RNA</a> of the TMV was infectious which reinforced the argument. The RNA carries <a href="/wiki/Genetics" title="Genetics">genetic</a> information to code for the production of new infectious particles. </p><p>More recently virus research has been focused on understanding the genetics and molecular biology of plant virus <a href="/wiki/Genomes" class="mw-redirect" title="Genomes">genomes</a>, with a particular interest in determining how the virus can replicate, move and infect plants. Understanding the virus genetics and protein functions has been used to explore the potential for commercial use by <a href="/wiki/Biotechnology" title="Biotechnology">biotechnology</a> companies. In particular, viral-derived sequences have been used to provide an understanding of novel forms of <a href="/wiki/Plant_disease_resistance" title="Plant disease resistance">resistance</a>. The recent boom in technology allowing humans to manipulate plant viruses may provide new strategies for production of value-added proteins in plants. </p> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=2" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Molecules-23-02311-g001.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Molecules-23-02311-g001.png/220px-Molecules-23-02311-g001.png" decoding="async" width="220" height="145" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Molecules-23-02311-g001.png/330px-Molecules-23-02311-g001.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Molecules-23-02311-g001.png/440px-Molecules-23-02311-g001.png 2x" data-file-width="2954" data-file-height="1952" /></a><figcaption>Structural comparison of some plant viruses</figcaption></figure> <p>Viruses are so small that they can only be observed under an <a href="/wiki/Electron_microscope" title="Electron microscope">electron microscope</a>. The structure of a virus is given by its coat of <a href="/wiki/Proteins" class="mw-redirect" title="Proteins">proteins</a>, which surround the viral <a href="/wiki/Genome" title="Genome">genome</a>. Assembly of viral particles takes place <a href="/wiki/Self-organization#Self-organization_in_biology" title="Self-organization">spontaneously</a>. </p><p>Over 50% of known plant viruses are <a href="/wiki/Rod-shaped" class="mw-redirect" title="Rod-shaped">rod-shaped</a> (<a href="/wiki/Flexuous" title="Flexuous">flexuous</a> or rigid). The length of the particle is normally dependent on the genome but it is usually between 300 and 500 <a href="/wiki/Nanometers" class="mw-redirect" title="Nanometers">nm</a> with a <a href="/wiki/Diameter" title="Diameter">diameter</a> of 15–20 nm. Protein subunits can be placed around the <a href="/wiki/Circumference" title="Circumference">circumference</a> of a circle to form a disc. In the presence of the viral genome, the discs are stacked, then a tube is created with room for the <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acid</a> genome in the middle.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>The second most common structure amongst plant viruses are <a href="/wiki/Isometric_(disambiguation)" class="mw-redirect mw-disambig" title="Isometric (disambiguation)">isometric</a> particles. They are 25–50 nm in diameter. In cases when there is only a single coat protein, the basic structure consists of 60 T subunits, where T is an <a href="/wiki/Integer" title="Integer">integer</a>. Some viruses may have 2 coat proteins that associate to form an <a href="/wiki/Icosahedron" title="Icosahedron">icosahedral</a> shaped particle. </p><p>There are three genera of <i><a href="/wiki/Geminiviridae" title="Geminiviridae">Geminiviridae</a></i> that consist of particles that are like two isometric particles stuck together. </p><p>A few number of plant viruses have, in addition to their coat proteins, a <a href="/wiki/Cell_membrane" title="Cell membrane">lipid envelope</a>. This is derived from the plant cell membrane as the virus particle <a href="/wiki/Viral_shedding" title="Viral shedding">buds</a> off from the <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Transmission">Transmission</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=3" title="Edit section: Transmission"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Through_sap">Through sap</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=4" title="Edit section: Through sap"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Viruses can be spread by direct transfer of sap by contact of a wounded plant with a healthy one. Such contact may occur during agricultural practices, as by damage caused by tools or hands, or naturally, as by an animal feeding on the plant. Generally TMV, potato viruses and cucumber mosaic viruses are transmitted via sap. </p> <div class="mw-heading mw-heading3"><h3 id="By_insects">By insects</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=5" title="Edit section: By insects"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Viruses-08-00303-g001.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Viruses-08-00303-g001.png/440px-Viruses-08-00303-g001.png" decoding="async" width="440" height="415" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Viruses-08-00303-g001.png/660px-Viruses-08-00303-g001.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/24/Viruses-08-00303-g001.png/880px-Viruses-08-00303-g001.png 2x" data-file-width="3190" data-file-height="3010" /></a><figcaption>Plant virus transmission strategies in insect vectors</figcaption></figure> <p>Plant viruses need to be transmitted by a <a href="/wiki/Vector_(epidemiology)" class="mw-redirect" title="Vector (epidemiology)">vector</a>, most often <a href="/wiki/Insects" class="mw-redirect" title="Insects">insects</a> such as <a href="/wiki/Leafhoppers" class="mw-redirect" title="Leafhoppers">leafhoppers</a>. One class of viruses, the <a href="/wiki/Rhabdoviridae" title="Rhabdoviridae">Rhabdoviridae</a>, has been proposed to actually be insect viruses that have evolved to replicate in plants. The chosen insect vector of a plant virus will often be the determining factor in that virus's host range: it can only infect plants that the insect vector feeds upon. This was shown in part when the <a href="/wiki/Old_world" class="mw-redirect" title="Old world">old world</a> <a href="/wiki/White_fly" class="mw-redirect" title="White fly">white fly</a> made it to the United States, where it transferred many plant viruses into new hosts. Depending on the way they are transmitted, plant viruses are classified as non-persistent, semi-persistent and persistent. In non-persistent transmission, viruses become attached to the distal tip of the <a href="/wiki/Insect_mouthparts" title="Insect mouthparts">stylet</a> of the insect and on the next plant it feeds on, it inoculates it with the virus.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Semi-persistent viral transmission involves the virus entering the <a href="/wiki/Foregut" title="Foregut">foregut</a> of the insect. Those viruses that manage to pass through the gut into the <a href="/wiki/Haemolymph" class="mw-redirect" title="Haemolymph">haemolymph</a> and then to the <a href="/wiki/Salivary_glands" class="mw-redirect" title="Salivary glands">salivary glands</a> are known as persistent. There are two sub-classes of persistent viruses: propagative and circulative. Propagative viruses are able to replicate in both the plant and the insect (and may have originally been insect viruses), whereas circulative can not. Circulative viruses are protected inside aphids by the chaperone protein <a href="/w/index.php?title=Symbionin&action=edit&redlink=1" class="new" title="Symbionin (page does not exist)">symbionin</a>, produced by bacterial <a href="/wiki/Symbionts" class="mw-redirect" title="Symbionts">symbionts</a>. Many plant viruses encode within their genome <a href="/wiki/Polypeptide" class="mw-redirect" title="Polypeptide">polypeptides</a> with domains essential for transmission by insects. In non-persistent and semi-persistent viruses, these domains are in the coat protein and another protein known as the helper component. A bridging <a href="/wiki/Hypothesis" title="Hypothesis">hypothesis</a> has been proposed to explain how these proteins aid in insect-mediated viral transmission. The helper component will bind to the specific domain of the coat protein, and then the insect mouthparts – creating a bridge. In persistent propagative viruses, such as <a href="/wiki/Tomato_spotted_wilt_virus" class="mw-redirect" title="Tomato spotted wilt virus">tomato spotted wilt virus</a> (TSWV), there is often a lipid coat surrounding the proteins that is not seen in other classes of plant viruses. In the case of TSWV, 2 viral proteins are expressed in this lipid envelope. It has been proposed that the viruses bind via these proteins and are then taken into the insect <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> by receptor-mediated <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a>. </p> <div class="mw-heading mw-heading3"><h3 id="By_nematodes">By nematodes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=6" title="Edit section: By nematodes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Soil-borne <a href="/wiki/Nematode" title="Nematode">nematodes</a> have been shown to transmit viruses. They acquire and transmit them by feeding on infected <a href="/wiki/Root" title="Root">roots</a>. Viruses can be transmitted both non-persistently and persistently, but there is no evidence of viruses being able to replicate in nematodes. The <a href="/wiki/Virions" class="mw-redirect" title="Virions">virions</a> attach to the stylet (feeding organ) or to the gut when they feed on an infected plant and can then detach during later feeding to infect other plants. Nematodes transmit viruses such as <a href="/wiki/Tobacco_ringspot_virus" title="Tobacco ringspot virus">tobacco ringspot virus</a> and <a href="/wiki/Tobacco_rattle_virus" title="Tobacco rattle virus">tobacco rattle virus</a>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="By_plasmodiophorids">By plasmodiophorids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=7" title="Edit section: By plasmodiophorids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A number of virus genera are transmitted, both persistently and non-persistently, by soil borne <a href="/wiki/Zoospore" title="Zoospore">zoosporic</a> <a href="/wiki/Protozoa" title="Protozoa">protozoa</a>. These protozoa are not phytopathogenic themselves, but <a href="/wiki/Parasitic" class="mw-redirect" title="Parasitic">parasitic</a>. Transmission of the virus takes place when they become associated with the plant roots. Examples include <i><a href="/wiki/Polymyxa_graminis" title="Polymyxa graminis">Polymyxa graminis</a></i>, which has been shown to transmit plant viral diseases in cereal crops<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> and <i><a href="/w/index.php?title=Polymyxa_betae&action=edit&redlink=1" class="new" title="Polymyxa betae (page does not exist)">Polymyxa betae</a></i> which transmits <a href="/wiki/Beet_necrotic_yellow_vein_virus" title="Beet necrotic yellow vein virus">Beet necrotic yellow vein virus</a>. Plasmodiophorids also create wounds in the plant's root through which other viruses can enter. </p> <div class="mw-heading mw-heading3"><h3 id="On_seed_and_pollen">On seed and pollen</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=8" title="Edit section: On seed and pollen"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-Unreferenced_section plainlinks metadata ambox ambox-content ambox-Unreferenced" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>does not <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">cite</a> any <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">sources</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Plant_virus" title="Special:EditPage/Plant virus">improve this section</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and <a href="/wiki/Wikipedia:Verifiability#Burden_of_evidence" title="Wikipedia:Verifiability">removed</a>.</span> <span class="date-container"><i>(<span class="date">May 2024</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>Plant virus transmission from generation to generation occurs in about 20% of plant viruses. When viruses are transmitted by seeds, the seed is infected in the generative cells and the virus is maintained in the germ cells and sometimes, but less often, in the seed coat. When the growth and development of plants is delayed because of situations like unfavorable weather, there is an increase in the amount of virus infections in seeds. There does not seem to be a correlation between the location of the seed on the plant and its chances of being infected. Little is known about the mechanisms involved in the transmission of plant viruses via seeds, although it is known that it is environmentally influenced and that seed transmission occurs because of a direct invasion of the embryo via the ovule or by an indirect route with an attack on the embryo mediated by infected gametes. These processes can occur concurrently or separately depending on the host plant. It is unknown how the virus is able to directly invade and cross the embryo and boundary between the parental and progeny generations in the ovule. Many plants species can be infected through seeds including but not limited to the families <a href="/wiki/Leguminosae" class="mw-redirect" title="Leguminosae">Leguminosae</a>, <a href="/wiki/Solanaceae" title="Solanaceae">Solanaceae</a>, <a href="/wiki/Compositae" class="mw-redirect" title="Compositae">Compositae</a>, <a href="/wiki/Rosaceae" title="Rosaceae">Rosaceae</a>, <a href="/wiki/Cucurbitaceae" title="Cucurbitaceae">Cucurbitaceae</a>, <a href="/wiki/Gramineae" class="mw-redirect" title="Gramineae">Gramineae</a>. Bean common mosaic virus is transmitted through seeds. </p> <div class="mw-heading mw-heading3"><h3 id="Directly_from_plant_to_humans">Directly from plant to humans</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=9" title="Edit section: Directly from plant to humans"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There is tenuous evidence that a virus common to peppers, the <a href="/wiki/Pepper_mild_mottle_virus" title="Pepper mild mottle virus">Pepper Mild Mottle Virus</a> (PMMoV) may have moved on to infect humans.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> This is a rare and unlikely event as, to enter a cell and replicate, a virus must "bind to a receptor on its surface, and a plant virus would be highly unlikely to recognize a receptor on a human cell. One possibility is that the virus does not infect human cells directly. Instead, the naked viral RNA may alter the function of the cells through a mechanism similar to <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a>, in which the presence of certain RNA sequences can turn genes on and off," according to Virologist Robert Garry.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Effects_on_hosts">Effects on hosts</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=10" title="Edit section: Effects on hosts"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The intracellular life of plant viruses in hosts is still understudied especially the earliest <a href="/wiki/Viral_life_cycle" title="Viral life cycle">stages of infection</a>.<sup id="cite_ref-Laliberte-Zheng-2014_11-0" class="reference"><a href="#cite_note-Laliberte-Zheng-2014-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Many membranous structures which viruses induce plant cells to produce are motile, often being used to traffic new virions within the producing cell and into their neighbors.<sup id="cite_ref-Laliberte-Zheng-2014_11-1" class="reference"><a href="#cite_note-Laliberte-Zheng-2014-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Viruses also induce various changes to plants' own <a href="/wiki/Intracellular_membrane" class="mw-redirect" title="Intracellular membrane">intracellular membranes</a>.<sup id="cite_ref-Laliberte-Zheng-2014_11-2" class="reference"><a href="#cite_note-Laliberte-Zheng-2014-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> The work of Perera et al. 2012 in <a href="/wiki/Mosquito" title="Mosquito">mosquito</a> virus infection and various others studying <a href="/wiki/Yeast" title="Yeast">yeast</a> models of plant viruses find this to be due to changes in <a href="/wiki/Homeostasis" title="Homeostasis">homeostasis</a> of the lipids that compose <i>their</i> intracellular membranes, including increasing <a href="/wiki/Membrane_lipid_synthesis" class="mw-redirect" title="Membrane lipid synthesis">synthesis</a>.<sup id="cite_ref-Laliberte-Zheng-2014_11-3" class="reference"><a href="#cite_note-Laliberte-Zheng-2014-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> These comparable lipid alterations inform our expectations and research directions for the lesser understood area of plant viruses.<sup id="cite_ref-Laliberte-Zheng-2014_11-4" class="reference"><a href="#cite_note-Laliberte-Zheng-2014-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Translation_of_plant_viral_proteins">Translation of plant viral proteins</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=11" title="Edit section: Translation of plant viral proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Fpls-09-00666-g002.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Fpls-09-00666-g002.jpg/220px-Fpls-09-00666-g002.jpg" decoding="async" width="220" height="309" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Fpls-09-00666-g002.jpg/330px-Fpls-09-00666-g002.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Fpls-09-00666-g002.jpg/440px-Fpls-09-00666-g002.jpg 2x" data-file-width="1818" data-file-height="2550" /></a><figcaption>Polyprotein processing is used by 45% of plant viruses. Plant virus families that produce <a href="/wiki/Protein" title="Protein">polyproteins</a>, their genomes, and colored triangles indicating <a href="/wiki/Proteolysis" title="Proteolysis">self-cleavage</a> sites.<sup id="cite_ref-:0_12-0" class="reference"><a href="#cite_note-:0-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>75% of plant viruses have genomes that consist of single stranded RNA (ssRNA). 65% of plant viruses have +ssRNA, meaning that they are in the same sense orientation as <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> but 10% have -ssRNA, meaning they must be converted to +ssRNA before they can be translated. 5% are double stranded RNA and so can be immediately translated as +ssRNA viruses. 3% require a <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a> enzyme to convert between RNA and DNA. 17% of plant viruses are ssDNA and very few are dsDNA, in contrast a quarter of animal viruses are dsDNA and three-quarters of <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophage</a> are dsDNA.<sup id="cite_ref-Hull_2001_13-0" class="reference"><a href="#cite_note-Hull_2001-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Viruses use the plant <a href="/wiki/Ribosomes" class="mw-redirect" title="Ribosomes">ribosomes</a> to produce the 4-10 proteins encoded by their genome. However, since many of the proteins are encoded on a single strand (that is, they are <a href="/wiki/Polycistronic" class="mw-redirect" title="Polycistronic">polycistronic</a>) this will mean that the ribosome will either only produce one protein, as it will terminate translation at the first <a href="/wiki/Stop_codon" title="Stop codon">stop codon</a>, or that a <a href="/wiki/Polyprotein" class="mw-redirect" title="Polyprotein">polyprotein</a> will be produced. Plant viruses have had to evolve special techniques to allow the production of viral proteins by <a href="/wiki/Plant_cell" title="Plant cell">plant cells</a>. </p> <div class="mw-heading mw-heading3"><h3 id="5'_Cap"><span id="5.27_Cap"></span>5' Cap</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=12" title="Edit section: 5' Cap"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>For <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> to occur, <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> mRNAs require a <a href="/wiki/5%27_Cap" class="mw-redirect" title="5' Cap">5' Cap</a> structure. This means that viruses must also have one. This normally consists of 7MeGpppN where N is normally <a href="/wiki/Adenine" title="Adenine">adenine</a> or <a href="/wiki/Guanine" title="Guanine">guanine</a>. The viruses encode a protein, normally a <a href="/wiki/Replicase" class="mw-redirect" title="Replicase">replicase</a>, with a <a href="/wiki/Methyltransferase" title="Methyltransferase">methyltransferase</a> activity to allow this. </p><p>Some viruses are cap-snatchers. During this process, a <sup>7m</sup>G-capped host mRNA is recruited by the viral transcriptase complex and subsequently cleaved by a virally encoded endonuclease. The resulting capped leader RNA is used to prime transcription on the viral genome.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>However some plant viruses do not use cap, yet translate efficiently due to cap-independent translation enhancers present in 5' and 3' untranslated regions of viral mRNA.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Readthrough">Readthrough</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=13" title="Edit section: Readthrough"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some viruses (e.g. <a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">tobacco mosaic virus</a> (TMV)) have RNA sequences that contain a "leaky" stop codon. In TMV 95% of the time the host ribosome will terminate the synthesis of the polypeptide at this codon but the rest of the time it continues past it. This means that 5% of the proteins produced are larger than and different from the others normally produced, which is a form of <a href="/wiki/Translational_regulation" title="Translational regulation">translational regulation</a>. In TMV, this extra sequence of polypeptide is an <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a> that replicates its genome. </p> <div class="mw-heading mw-heading3"><h3 id="Production_of_sub-genomic_RNAs">Production of sub-genomic RNAs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=14" title="Edit section: Production of sub-genomic RNAs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some viruses use the production of <a href="/wiki/Subgenomic" class="mw-redirect" title="Subgenomic">subgenomic</a> RNAs to ensure the translation of all proteins within their genomes. In this process the first protein encoded on the genome, and is the first to be translated, is a <a href="/wiki/RNA_replicase" class="mw-redirect" title="RNA replicase">replicase</a>. This protein will act on the rest of the genome producing negative strand sub-genomic RNAs then act upon these to form positive strand sub-genomic RNAs that are essentially mRNAs ready for translation. </p> <div class="mw-heading mw-heading3"><h3 id="Segmented_genomes">Segmented genomes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=15" title="Edit section: Segmented genomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some viral families, such as the <i><a href="/wiki/Bromoviridae" title="Bromoviridae">Bromoviridae</a></i> instead opt to have <a href="/wiki/Multipartite" title="Multipartite">multipartite</a> genomes, genomes split between multiple viral particles. For infection to occur, the plant must be infected with all particles across the genome. For instance <i><a href="/wiki/Brome_mosaic_virus" title="Brome mosaic virus">Brome mosaic virus</a></i> has a genome split between 3 viral particles, and all 3 particles with the different RNAs are required for <a href="/wiki/Infection" title="Infection">infection</a> to take place. </p> <div class="mw-heading mw-heading3"><h3 id="Polyprotein_processing">Polyprotein processing</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=16" title="Edit section: Polyprotein processing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Polyprotein processing is adopted by 45% of plant viruses, such as the <a href="/wiki/Potyviridae" title="Potyviridae">Potyviridae</a> and <a href="/wiki/Tymoviridae" title="Tymoviridae">Tymoviridae</a>.<sup id="cite_ref-:0_12-1" class="reference"><a href="#cite_note-:0-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The ribosome translates a single protein from the viral genome. Within the polyprotein is an enzyme (or enzymes) with <a href="/wiki/Proteinase" class="mw-redirect" title="Proteinase">proteinase</a> function that is able to cleave the polyprotein into the various single proteins or just cleave away the protease, which can then cleave other polypeptides producing the mature proteins. </p> <div class="mw-heading mw-heading3"><h3 id="Genome_packaging">Genome packaging</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=17" title="Edit section: Genome packaging"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Besides involvement in the infection process, <a href="/wiki/Viral_replicase" class="mw-redirect" title="Viral replicase">viral replicase</a> is a directly necessary part of the <a href="/wiki/Genome_packaging" class="mw-redirect" title="Genome packaging">packaging of RNA viruses' genetic material</a>. This was expected due to replicase involvement already being confirmed in various other viruses.<sup id="cite_ref-Rao-2006_16-0" class="reference"><a href="#cite_note-Rao-2006-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>The genome of Beet necrotic yellow vein virus (BNYVV) consists of five RNAs, each encapsidated into rod-shaped virus particles. RNA 1, which is 6746 nucleotides long, encodes a single open reading frame (ORF) that produces the 237 kDa protein P237. This protein is cleaved into P150 and P66 by a papain-like proteinase. RNA 2, 4612 nucleotides long, encodes six proteins, including movement proteins (P42, P13, P15), a coat protein (P21), and a regulatory protein (P14). RNA 3, 1775 nucleotides long, encodes P25, which is involved in symptom expression. RNA 4, 1431 nucleotides long, encodes P31, crucial for vector transmission. RNA 5, found in certain isolates, encodes P26 and is associated with more severe symptoms.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Applications_of_plant_viruses">Applications of plant viruses</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=18" title="Edit section: Applications of plant viruses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Plant viruses can be used to engineer <a href="/wiki/Viral_vector" title="Viral vector">viral vectors</a>, tools commonly used by molecular <a href="/wiki/Biologist" title="Biologist">biologists</a> to deliver <a href="/wiki/Genetic_material" class="mw-redirect" title="Genetic material">genetic material</a> into plant <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a>; they are also sources of biomaterials and nanotechnology devices.<sup id="cite_ref-:1_18-0" class="reference"><a href="#cite_note-:1-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_19-0" class="reference"><a href="#cite_note-:2-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Knowledge of plant viruses and their components has been instrumental for the development of modern plant biotechnology. The use of plant viruses to enhance the beauty of ornamental plants can be considered the first recorded application of plant viruses. <a href="/wiki/Tulip_breaking_virus" title="Tulip breaking virus">Tulip breaking virus</a> is famous for its dramatic effects on the color of the tulip <a href="/wiki/Perianth" title="Perianth">perianth</a>, an effect highly sought after during the 17th-century Dutch "<a href="/wiki/Tulip_mania" title="Tulip mania">tulip mania</a>." <a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">Tobacco mosaic virus</a> (TMV) and <a href="/wiki/Cauliflower_mosaic_virus" title="Cauliflower mosaic virus">cauliflower mosaic virus</a> (CaMV) are frequently used in plant molecular biology. Of special interest is the CaMV 35S <a href="/wiki/Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoter</a>, which is a very strong promoter most frequently used in plant <a href="/wiki/Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">transformations</a>. Viral vectors based on <a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">tobacco mosaic virus</a> include those of the <a rel="nofollow" class="external text" href="https://www.icongenetics.com/technology/">magnICON®</a> and TRBO plant expression technologies.<sup id="cite_ref-:2_19-1" class="reference"><a href="#cite_note-:2-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Semper_Augustus_Tulip_17th_century.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Semper_Augustus_Tulip_17th_century.jpg/220px-Semper_Augustus_Tulip_17th_century.jpg" decoding="async" width="220" height="340" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Semper_Augustus_Tulip_17th_century.jpg/330px-Semper_Augustus_Tulip_17th_century.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cc/Semper_Augustus_Tulip_17th_century.jpg/440px-Semper_Augustus_Tulip_17th_century.jpg 2x" data-file-width="1182" data-file-height="1829" /></a><figcaption><b>Application of plant viruses to enhance the plant beauty</b>. The <i>Semper Augustus</i>, famous for being the most expensive tulip sold during <a href="/wiki/Tulip_mania" title="Tulip mania">tulip mania</a>. The effects of <a href="/wiki/Tulip_breaking_virus" title="Tulip breaking virus">tulip breaking virus</a> are seen in the striking streaks of white in its red petals.</figcaption></figure> <p>Building on the market approvals and sales of recombinant virus-based biopharmaceuticals for veterinary and human medicine, the use of engineered plant viruses has been proposed to enhance crop performance and promote sustainable production.<sup id="cite_ref-:3_20-0" class="reference"><a href="#cite_note-:3-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>Representative applications of plant viruses are listed below. </p> <table class="wikitable"> <caption>Applications of plant viruses<sup id="cite_ref-:1_18-1" class="reference"><a href="#cite_note-:1-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <td><b>Use</b> </td> <td><b>Description</b> </td> <th><b>References</b> </th></tr> <tr> <td>Enhanced plant aesthetics </td> <td>Increase beauty and commercial value of ornamental plants </td> <td><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Cross‐protection </td> <td>Delivery of mild virus strains to prevent infections by their severe relatives </td> <td><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Weed biocontrol </td> <td>Viruses triggering lethal systemic necrosis as bioherbicides </td> <td><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Pest biocontrol </td> <td>Enhanced toxin and pesticide delivery for insect and nematode control </td> <td><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Nanoparticle scaffolds </td> <td>Virion surfaces are functionalized and used to assemble nanoparticles </td> <td><sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Nanocarriers </td> <td>Virions are used to transport cargo compounds </td> <td><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><a href="/wiki/Nanoreactor" title="Nanoreactor">Nanoreactors</a> </td> <td>Enzymes are encapsulated into virions to engineer cascade reactions </td> <td><sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Recombinant protein/peptide expression </td> <td>Fast, transient overproduction of recombinant peptide, polypeptide libraries and protein complexes </td> <td><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Functional genomic studies </td> <td>Targeted gene silencing using <a href="/wiki/VIGS" class="mw-redirect" title="VIGS">VIGS</a> and miRNA viral vectors </td> <td><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><a href="/wiki/Genome_editing" title="Genome editing">Genome editing</a> </td> <td>Targeted genome editing <i>via</i> transient delivery of sequence‐specific nucleases </td> <td><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Metabolic pathway engineering </td> <td>Biosynthetic pathway rewiring to improve production of native and foreign metabolites </td> <td><sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Flowering induction </td> <td>Viral expression of <i>FLOWERING LOCUS T</i> to accelerate flowering induction and crop breeding </td> <td><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>Crop <a href="/wiki/Gene_therapy" title="Gene therapy">gene therapy</a> </td> <td>Open‐field use of viral vectors for transient reprogramming of crop traits within a single growing season </td> <td><sup id="cite_ref-:3_20-1" class="reference"><a href="#cite_note-:3-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=19" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Plant_disease" title="Plant disease">plant disease</a> – Diseases of plants</li> <li><a href="/wiki/Plant_pathology" title="Plant pathology">plant pathology</a> – Scientific study of plant diseases</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=Plant_virus&action=edit&section=20" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <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"><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="CITEREFRoossinck2011" class="citation journal cs1">Roossinck, Marilyn J. 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Llorente, Briardo; Rodríguez-Concepción, Manuel; Daròs, José-Antonio (31 January 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5282570">"Rewiring carotenoid biosynthesis in plants using a viral vector"</a>. <i>Scientific Reports</i>. <b>7</b>: 41645. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2017NatSR...741645M">2017NatSR...741645M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fsrep41645">10.1038/srep41645</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2045-2322">2045-2322</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5282570">5282570</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/28139696">28139696</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Scientific+Reports&rft.atitle=Rewiring+carotenoid+biosynthesis+in+plants+using+a+viral+vector&rft.volume=7&rft.pages=41645&rft.date=2017-01-31&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5282570%23id-name%3DPMC&rft_id=info%3Abibcode%2F2017NatSR...741645M&rft_id=info%3Apmid%2F28139696&rft_id=info%3Adoi%2F10.1038%2Fsrep41645&rft.issn=2045-2322&rft.aulast=Majer&rft.aufirst=Eszter&rft.au=Llorente%2C+Briardo&rft.au=Rodr%C3%ADguez-Concepci%C3%B3n%2C+Manuel&rft.au=Dar%C3%B2s%2C+Jos%C3%A9-Antonio&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5282570&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+virus" class="Z3988"></span></span> </li> <li id="cite_note-34"><span class="mw-cite-backlink"><b><a href="#cite_ref-34">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMcGarryKlockoPangStrauss2017" class="citation journal cs1">McGarry, Roisin C.; Klocko, Amy L.; Pang, Mingxiong; Strauss, Steven H.; Ayre, Brian G. (January 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5210732">"Virus-Induced Flowering: An Application of Reproductive Biology to Benefit Plant Research and Breeding"</a>. <i>Plant Physiology</i>. <b>173</b> (1): <span class="nowrap">47–</span>55. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1104%2Fpp.16.01336">10.1104/pp.16.01336</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1532-2548">1532-2548</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5210732">5210732</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/27856915">27856915</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Plant+Physiology&rft.atitle=Virus-Induced+Flowering%3A+An+Application+of+Reproductive+Biology+to+Benefit+Plant+Research+and+Breeding&rft.volume=173&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E47-%3C%2Fspan%3E55&rft.date=2017-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5210732%23id-name%3DPMC&rft.issn=1532-2548&rft_id=info%3Apmid%2F27856915&rft_id=info%3Adoi%2F10.1104%2Fpp.16.01336&rft.aulast=McGarry&rft.aufirst=Roisin+C.&rft.au=Klocko%2C+Amy+L.&rft.au=Pang%2C+Mingxiong&rft.au=Strauss%2C+Steven+H.&rft.au=Ayre%2C+Brian+G.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5210732&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+virus" class="Z3988"></span></span> </li> <li id="cite_note-35"><span class="mw-cite-backlink"><b><a href="#cite_ref-35">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTortiSchlesierThümmlerBartels2021" class="citation journal cs1">Torti, Stefano; Schlesier, René; Thümmler, Anka; Bartels, Doreen; Römer, Patrick; Koch, Birgit; Werner, Stefan; Panwar, Vinay; Kanyuka, Kostya; Wirén, Nicolaus von; Jones, Jonathan D. G.; Hause, Gerd; Giritch, Anatoli; Gleba, Yuri (February 2021). <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/33594264">"Transient reprogramming of crop plants for agronomic performance"</a>. <i>Nature Plants</i>. <b>7</b> (2): <span class="nowrap">159–</span>171. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2021NatPl...7..159T">2021NatPl...7..159T</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fs41477-021-00851-y">10.1038/s41477-021-00851-y</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2055-0278">2055-0278</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/33594264">33594264</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:231945168">231945168</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Plants&rft.atitle=Transient+reprogramming+of+crop+plants+for+agronomic+performance&rft.volume=7&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E159-%3C%2Fspan%3E171&rft.date=2021-02&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A231945168%23id-name%3DS2CID&rft_id=info%3Abibcode%2F2021NatPl...7..159T&rft.issn=2055-0278&rft_id=info%3Adoi%2F10.1038%2Fs41477-021-00851-y&rft_id=info%3Apmid%2F33594264&rft.aulast=Torti&rft.aufirst=Stefano&rft.au=Schlesier%2C+Ren%C3%A9&rft.au=Th%C3%BCmmler%2C+Anka&rft.au=Bartels%2C+Doreen&rft.au=R%C3%B6mer%2C+Patrick&rft.au=Koch%2C+Birgit&rft.au=Werner%2C+Stefan&rft.au=Panwar%2C+Vinay&rft.au=Kanyuka%2C+Kostya&rft.au=Wir%C3%A9n%2C+Nicolaus+von&rft.au=Jones%2C+Jonathan+D.+G.&rft.au=Hause%2C+Gerd&rft.au=Giritch%2C+Anatoli&rft.au=Gleba%2C+Yuri&rft_id=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F33594264&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+virus" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading3"><h3 id="Further_reading">Further reading</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=21" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZaitlinPalukaitis2000" class="citation journal cs1"><a href="/wiki/Milton_Zaitlin" title="Milton Zaitlin">Zaitlin, Milton</a>; Palukaitis, Peter (2000). "Advances in Understanding Plant Viruses and Virus Diseases". <i>Annual Review of Phytopathology</i>. <b>38</b>: <span class="nowrap">117–</span>143. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.phyto.38.1.117">10.1146/annurev.phyto.38.1.117</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11701839">11701839</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Phytopathology&rft.atitle=Advances+in+Understanding+Plant+Viruses+and+Virus+Diseases&rft.volume=38&rft.pages=%3Cspan+class%3D%22nowrap%22%3E117-%3C%2Fspan%3E143&rft.date=2000&rft_id=info%3Adoi%2F10.1146%2Fannurev.phyto.38.1.117&rft_id=info%3Apmid%2F11701839&rft.aulast=Zaitlin&rft.aufirst=Milton&rft.au=Palukaitis%2C+Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+virus" class="Z3988"></span></li> <li>Zaitlin, Milton (1998), <i>Discoveries in Plant Biology</i>, New York 14853, USA. pp. 105–110. S. D. Kung and S. F. Yang (eds).</li> <li>Dickinson, M. (2003), <i>Molecular Plant Pathology</i>. <a href="/wiki/BIOS_Scientific_Publishers" title="BIOS Scientific Publishers">BIOS Scientific Publishers</a>.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWang_DaowenMaule_Andrew_J1994" class="citation journal cs1">Wang Daowen; Maule Andrew J (1994). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC160477">"A Model for Seed Transmission of a Plant Virus: Genetic and Structural Analyses of Pea Embryo Invasion by Pea Seed-Borne Mosaic Virus"</a>. <i>The Plant Cell</i>. <b>6</b> (6): <span class="nowrap">777–</span>787. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1994PlanC...6..777W">1994PlanC...6..777W</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2307%2F3869957">10.2307/3869957</a>. <a href="/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/3869957">3869957</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC160477">160477</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12244258">12244258</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Plant+Cell&rft.atitle=A+Model+for+Seed+Transmission+of+a+Plant+Virus%3A+Genetic+and+Structural+Analyses+of+Pea+Embryo+Invasion+by+Pea+Seed-Borne+Mosaic+Virus&rft.volume=6&rft.issue=6&rft.pages=%3Cspan+class%3D%22nowrap%22%3E777-%3C%2Fspan%3E787&rft.date=1994&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC160477%23id-name%3DPMC&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F3869957%23id-name%3DJSTOR&rft_id=info%3Abibcode%2F1994PlanC...6..777W&rft_id=info%3Apmid%2F12244258&rft_id=info%3Adoi%2F10.2307%2F3869957&rft.au=Wang+Daowen&rft.au=Maule+Andrew+J&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC160477&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlant+virus" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plant_virus&action=edit&section=22" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output 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style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini 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this template">e</abbr></a></li></ul></div><div id="Microbiology:_Virus661" style="font-size:114%;margin:0 4em"><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a>: <a href="/wiki/Virus" title="Virus">Virus</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li><a href="/wiki/History_of_virology" title="History of virology">History</a></li> <li><a href="/wiki/Introduction_to_viruses" title="Introduction to viruses">Introduction</a></li> <li><a href="/wiki/Social_history_of_viruses" title="Social history of viruses">Social history of viruses</a></li> <li><a href="/wiki/Virology" title="Virology">Virology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Components</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/Capsid" title="Capsid">Capsid</a></li> <li><a href="/wiki/Viral_envelope" title="Viral envelope">Viral envelope</a></li> <li><a href="/wiki/Viral_protein" title="Viral protein">Viral protein</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Rotavirus_Reconstruction.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/100px-Rotavirus_Reconstruction.jpg" decoding="async" width="100" height="88" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/150px-Rotavirus_Reconstruction.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/200px-Rotavirus_Reconstruction.jpg 2x" data-file-width="1986" data-file-height="1739" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Viral_life_cycle" title="Viral life cycle">Viral life cycle</a></th><td class="navbox-list-with-group 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style="width:1%">Other</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/Antiviral_drug" title="Antiviral drug">Antiviral drug</a></li> <li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li> <li><a href="/wiki/Helper_virus" title="Helper virus">Helper virus</a></li> <li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a> <ul><li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li><a href="/wiki/Laboratory_diagnosis_of_viral_infections" title="Laboratory diagnosis of viral infections">Laboratory diagnosis of viral infections</a></li> <li><a href="/wiki/Marine_viruses" title="Marine viruses">Marine viruses</a></li> <li><a href="/wiki/Neurotropic_virus" title="Neurotropic virus">Neurotropic virus</a></li> <li><a href="/wiki/Oncovirus" title="Oncovirus">Oncovirus</a></li> <li><a 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