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vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#RNA_silencing"> <div class="vector-toc-text"> 3 RNA silencing </div> </a> <ul id="toc-RNA_silencing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Viroid-like_elements" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Viroid-like_elements"> <div class="vector-toc-text"> 4 Viroid-like elements </div> </a> <button aria-controls="toc-Viroid-like_elements-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Viroid-like elements subsection </button> <ul id="toc-Viroid-like_elements-sublist" class="vector-toc-list"> <li id="toc-Viroid-like_satellite_RNAs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Viroid-like_satellite_RNAs"> <div class="vector-toc-text"> 4.1 Viroid-like satellite RNAs </div> </a> <ul id="toc-Viroid-like_satellite_RNAs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ambiviruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ambiviruses"> <div class="vector-toc-text"> 4.2 Ambiviruses </div> </a> <ul id="toc-Ambiviruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Retroviroids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Retroviroids"> <div class="vector-toc-text"> 4.3 Retroviroids </div> </a> <ul id="toc-Retroviroids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Obelisks" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Obelisks"> <div class="vector-toc-text"> 4.4 Obelisks </div> </a> <ul id="toc-Obelisks-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-RNA_world_hypothesis" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#RNA_world_hypothesis"> <div class="vector-toc-text"> 5 RNA world hypothesis </div> </a> <ul id="toc-RNA_world_hypothesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Control" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Control"> <div class="vector-toc-text"> 6 Control </div> </a> <ul id="toc-Control-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> 7 History </div> </a> <ul id="toc-History-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"> 8 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 9 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-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"> 10 External links </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" 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Available in 47 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-47" aria-hidden="true" > 47 languages </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%81%D9%8A%D8%B1%D9%88%D8%B3%D9%8A_%D8%A7%D9%84%D8%B4%D9%83%D9%84" title="فيروسي الشكل – Arabic" lang="ar" hreflang="ar" data-title="فيروسي الشكل" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Viroide" title="Viroide – Asturian" lang="ast" hreflang="ast" data-title="Viroide" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target">Asturianu</a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%92%D1%96%D1%80%D0%BE%D1%96%D0%B4%D1%8B" title="Віроіды – Belarusian" lang="be" hreflang="be" data-title="Віроіды" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target">Беларуская</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D0%BE%D0%B8%D0%B4" title="Вироид – Bulgarian" lang="bg" hreflang="bg" data-title="Вироид" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Viroide" title="Viroide – Catalan" lang="ca" hreflang="ca" data-title="Viroide" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Viroid" title="Viroid – Czech" lang="cs" hreflang="cs" data-title="Viroid" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Viroid" title="Viroid – German" lang="de" hreflang="de" data-title="Viroid" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Viroidid" title="Viroidid – Estonian" lang="et" hreflang="et" data-title="Viroidid" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%99%CE%BF%CE%B5%CE%B9%CE%B4%CE%AE" title="Ιοειδή – Greek" lang="el" hreflang="el" data-title="Ιοειδή" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Viroide" title="Viroide – Spanish" lang="es" hreflang="es" data-title="Viroide" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Biroide" title="Biroide – Basque" lang="eu" hreflang="eu" data-title="Biroide" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%88%DB%8C%D8%B1%D9%88%D8%A6%DB%8C%D8%AF" title="ویروئید – Persian" lang="fa" hreflang="fa" data-title="ویروئید" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Viro%C3%AFde" title="Viroïde – French" lang="fr" hreflang="fr" data-title="Viroïde" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-fy mw-list-item"><a href="https://fy.wikipedia.org/wiki/Firo%C3%AFde" title="Firoïde – Western Frisian" lang="fy" hreflang="fy" data-title="Firoïde" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target">Frysk</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/V%C3%ADor%C3%B3id" title="Víoróid – Irish" lang="ga" hreflang="ga" data-title="Víoróid" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Viroide" title="Viroide – Galician" lang="gl" hreflang="gl" data-title="Viroide" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%B0%94%EC%9D%B4%EB%A1%9C%EC%9D%B4%EB%93%9C" title="바이로이드 – Korean" lang="ko" hreflang="ko" data-title="바이로이드" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%B5%E0%A4%BE%E0%A4%AF%E0%A4%B0%E0%A5%8B%E0%A4%87%E0%A4%A1" title="वायरोइड – Hindi" lang="hi" hreflang="hi" data-title="वायरोइड" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Viroid" title="Viroid – Indonesian" lang="id" hreflang="id" data-title="Viroid" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Viroide" title="Viroide – Italian" lang="it" hreflang="it" data-title="Viroide" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%95%D7%99%D7%A8%D7%95%D7%90%D7%99%D7%93" title="וירואיד – Hebrew" lang="he" hreflang="he" data-title="וירואיד" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Viroid" title="Viroid – Javanese" lang="jv" hreflang="jv" data-title="Viroid" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target">Jawa</a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%B8%E0%B3%82%E0%B2%95%E0%B3%8D%E0%B2%B7%E0%B3%8D%E0%B2%AE_%E0%B2%B5%E0%B3%88%E0%B2%B0%E0%B2%BE%E0%B2%A3%E0%B3%81" title="ಸೂಕ್ಷ್ಮ ವೈರಾಣು – Kannada" lang="kn" hreflang="kn" data-title="ಸೂಕ್ಷ್ಮ ವೈರಾಣು" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target">ಕನ್ನಡ</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D0%BE%D0%B8%D0%B4%D1%82%D0%B0%D1%80" title="Вироидтар – Kazakh" lang="kk" hreflang="kk" data-title="Вироидтар" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Viroidas" title="Viroidas – Lithuanian" lang="lt" hreflang="lt" data-title="Viroidas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-lmo mw-list-item"><a href="https://lmo.wikipedia.org/wiki/Viroid" title="Viroid – Lombard" lang="lmo" hreflang="lmo" data-title="Viroid" data-language-autonym="Lombard" data-language-local-name="Lombard" class="interlanguage-link-target">Lombard</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Viroid" title="Viroid – Hungarian" lang="hu" hreflang="hu" data-title="Viroid" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D0%BE%D0%B8%D0%B4" title="Вироид – Macedonian" lang="mk" hreflang="mk" data-title="Вироид" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B5%E0%B5%88%E0%B4%B1%E0%B5%8B%E0%B4%AF%E0%B5%8D%E0%B4%A1%E0%B5%8D" title="വൈറോയ്ഡ് – Malayalam" lang="ml" hreflang="ml" data-title="വൈറോയ്ഡ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Viro%C3%AFde" title="Viroïde – Dutch" lang="nl" hreflang="nl" data-title="Viroïde" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%A6%E3%82%A4%E3%83%AD%E3%82%A4%E3%83%89" title="ウイロイド – Japanese" lang="ja" hreflang="ja" data-title="ウイロイド" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Viroid" title="Viroid – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Viroid" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Wiroid" title="Wiroid – Polish" lang="pl" hreflang="pl" data-title="Wiroid" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Viroide" title="Viroide – Portuguese" lang="pt" hreflang="pt" data-title="Viroide" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Viroid" title="Viroid – Romanian" lang="ro" hreflang="ro" data-title="Viroid" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-ru badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ru.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D0%BE%D0%B8%D0%B4%D1%8B" title="Вироиды – Russian" lang="ru" hreflang="ru" data-title="Вироиды" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Viroid" title="Viroid – Simple English" lang="en-simple" hreflang="en-simple" data-title="Viroid" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Viroid" title="Viroid – Slovak" lang="sk" hreflang="sk" data-title="Viroid" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%DA%A4%D8%A7%DB%8C%D8%B1%DB%86%DB%8C%D8%AF" title="ڤایرۆید – Central Kurdish" lang="ckb" hreflang="ckb" data-title="ڤایرۆید" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D0%BE%D0%B8%D0%B4" title="Вироид – Serbian" lang="sr" hreflang="sr" data-title="Вироид" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Viroidi" title="Viroidi – Finnish" lang="fi" hreflang="fi" data-title="Viroidi" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Viroid" title="Viroid – Swedish" lang="sv" hreflang="sv" data-title="Viroid" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%84%E0%B8%A7%E0%B8%A3%E0%B8%AD%E0%B8%A2%E0%B8%94%E0%B9%8C" title="ไวรอยด์ – Thai" lang="th" hreflang="th" data-title="ไวรอยด์" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target">ไทย</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Viroid" title="Viroid – Turkish" lang="tr" hreflang="tr" data-title="Viroid" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%92%D1%96%D1%80%D0%BE%D1%97%D0%B4" title="Віроїд – Ukrainian" lang="uk" hreflang="uk" data-title="Віроїд" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Viroid" title="Viroid – Vietnamese" lang="vi" hreflang="vi" data-title="Viroid" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target">Tiếng Việt</a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%B1%BB%E7%97%85%E6%AF%92" title="类病毒 – Chinese" lang="zh" hreflang="zh" data-title="类病毒" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target">中文</a></li> </ul> <div class="after-portlet after-portlet-lang"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q209917#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></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 id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet mw-portlet-associated-pages" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-nstab-main" class="selected vector-tab-noicon mw-list-item"><a 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searchaux" style="display:none">Pathogenic small single-stranded circular RNA</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">For viroid-like MGEs which are dependent on viruses, see <a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>.</div> <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(250,250,190)">Viroid </th></tr> <tr> <td colspan="2" style="text-align: center"><a href="/wiki/File:PSTviroid.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0f/PSTviroid.png/250px-PSTviroid.png" decoding="async" width="220" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0f/PSTviroid.png/330px-PSTviroid.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0f/PSTviroid.png/500px-PSTviroid.png 2x" data-file-width="1184" data-file-height="86" /></a> </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(250,250,190)"><a href="/wiki/Virus_classification" title="Virus classification">Virus classification</a> <a href="/wiki/Template:Taxonomy/Viroid" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a> </th></tr> <tr> <td>Informal group: </td> <td><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral agents</a> </td></tr> <tr> <td>(unranked): </td> <td>Viroid </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(250,250,190)">Families </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul><li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></li> <li><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li></ul> </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> Viroids are small single-stranded, <a href="/wiki/Circular_RNA" title="Circular RNA">circular RNAs</a> that are infectious pathogens.<a href="#cite_note-Navarro-1">[1]</a><a href="#cite_note-2">[2]</a> Unlike <a href="/wiki/Virus" title="Virus">viruses</a>, they have no protein coating. All known viroids are inhabitants of <a href="/wiki/Angiosperms" class="mw-redirect" title="Angiosperms">angiosperms</a> (flowering plants),<a href="#cite_note-3">[3]</a> and most cause diseases whose <a href="/wiki/Agricultural_economics" title="Agricultural economics">economic</a> importance to humans varies widely.<a href="#cite_note-pmid32752288-4">[4]</a> A recent <a href="/wiki/Metatranscriptomics" title="Metatranscriptomics">metatranscriptomics</a> study suggests that the host diversity of viroids and viroid-like elements is broader than previously thought and may not be limited to plants, encompassing even <a href="/wiki/Prokaryotes" class="mw-redirect" title="Prokaryotes">prokaryotes</a>.<a href="#cite_note-Lee-5">[5]</a> The first discoveries of viroids in the 1970s triggered the historically third major extension of the <a href="/wiki/Biosphere" title="Biosphere">biosphere</a>—to include smaller lifelike entities—after the discoveries in 1675 by <a href="/wiki/Antonie_van_Leeuwenhoek" title="Antonie van Leeuwenhoek">Antonie van Leeuwenhoek</a> (of the "subvisible" microorganisms) and in 1892–1898 by <a href="/wiki/Dmitri_Iosifovich_Ivanovsky" class="mw-redirect" title="Dmitri Iosifovich Ivanovsky">Dmitri Iosifovich Ivanovsky</a> and <a href="/wiki/Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a> (of the "submicroscopic" viruses). The unique properties of viroids have been recognized by the <a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">International Committee on Taxonomy of Viruses</a>, in creating new two <a href="/wiki/Family_(biology)" title="Family (biology)">families</a> of <a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">subviral agents</a>.<a href="#cite_note-6">[6]</a> Although viroids are composed of nucleic acid, they do not code for any <a href="/wiki/Protein" title="Protein">protein</a>.<a href="#cite_note-7">[7]</a><a href="#cite_note-8">[8]</a> The viroid's replication mechanism uses <a href="/wiki/RNA_polymerase_II" title="RNA polymerase II">RNA polymerase II</a>, a host cell enzyme normally associated with synthesis of <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> from DNA, which instead catalyzes "<a href="/wiki/Rolling_circle_replication" title="Rolling circle replication">rolling circle replication</a>" of new RNA using the viroid's RNA as a template. Viroids are often <a href="/wiki/Ribozymes" class="mw-redirect" title="Ribozymes">ribozymes</a>, having <a href="/wiki/Catalytic" class="mw-redirect" title="Catalytic">catalytic</a> properties that allow self-cleavage and ligation of unit-size genomes from larger replication intermediates.<a href="#cite_note-pmid33800543-9">[9]</a> Diener initially hypothesized in 1989 that viroids may represent "living relics" from the widely assumed, ancient, and non-cellular <a href="/wiki/RNA_world" title="RNA world">RNA world</a>, and others have followed this conjecture.<a href="#cite_note-Diener_1989-10">[10]</a><a href="#cite_note-11">[11]</a> Following the discovery of <a href="/wiki/Retrozyme" title="Retrozyme">retrozymes</a>, it has been proposed that viroids and other viroid-like elements may derive from this newly found class of <a href="/wiki/Retrotransposon" title="Retrotransposon">retrotransposon</a>.<a href="#cite_note-:3-12">[12]</a><a href="#cite_note-:0-13">[13]</a><a href="#cite_note-:1-14">[14]</a> The human pathogen <a href="/wiki/Hepatitis_D_virus" class="mw-redirect" title="Hepatitis D virus">hepatitis D virus</a> is a subviral agent similar in structure to a viroid, as it is a hybrid particle enclosed by surface proteins from the <a href="/wiki/Hepatitis_B_virus" title="Hepatitis B virus">hepatitis B virus</a>.<a href="#cite_note-15">[15]</a> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Taxonomy">Taxonomy</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=1" title="Edit section: Taxonomy">edit</a>]</div> <figure class="mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:PSTviroid.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0f/PSTviroid.png/960px-PSTviroid.png" decoding="async" width="800" height="58" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/0/0f/PSTviroid.png 1.5x" data-file-width="1184" data-file-height="86" /></a><figcaption>Putative <a href="/wiki/Nucleic_acid_secondary_structure" title="Nucleic acid secondary structure">secondary structure</a> of the PSTVd viroid. The highlighted nucleotides are found in most other viroids.</figcaption></figure> As of 2024<a class="external text" href="https://en.wikipedia.org/w/index.php?title=Viroid&action=edit">[update]</a>:<a href="#cite_note-pmid16078879-16">[16]</a><a href="#cite_note-17">[17]</a> <ul><li>Family <a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a>: relies on host Rnase III<a href="#cite_note-Lee-5">[5]</a> <ul><li>Genus <a href="/wiki/Pospiviroid" title="Pospiviroid">Pospiviroid</a>; type species: <a href="/wiki/Potato_spindle_tuber_viroid" title="Potato spindle tuber viroid">Pospiviroid fusituberis</a> (former name Potato spindle tuber viroid<a href="#cite_note-18">[18]</a>); 356–361 nucleotides(nt)<a href="#cite_note-nt-19">[19]</a> <ul><li><a href="/w/index.php?title=Pospiviroid_chloronani&action=edit&redlink=1" class="new" title="Pospiviroid chloronani (page does not exist)">Pospiviroid chloronani</a> (former name Tomato chlorotic dwarf viroid<a href="#cite_note-20">[20]</a>); (TCDVd); accession AF162131, genome length 360nt</li> <li><a href="/w/index.php?title=Mexican_papita_viroid&action=edit&redlink=1" class="new" title="Mexican papita viroid (page does not exist)">Mexican papita viroid</a>; (MPVd); accession L78454, genome length 360nt</li> <li><a href="/w/index.php?title=Pospiviroid_machoplantae&action=edit&redlink=1" class="new" title="Pospiviroid machoplantae (page does not exist)">Pospiviroid machoplantae</a> (former name Tomato planta macho viroid<a href="#cite_note-21">[21]</a>); (TPMVd); accession K00817, genome length 360nt</li> <li><a href="/wiki/Citrus_exocortis" title="Citrus exocortis">Pospiviroid exocortiscitri</a> (former name Citrus exocortis viroid<a href="#cite_note-22">[22]</a>); 368–467 nt<a href="#cite_note-nt-19">[19]</a></li> <li><a href="/w/index.php?title=Pospiviroid_impedichrysanthemi&action=edit&redlink=1" class="new" title="Pospiviroid impedichrysanthemi (page does not exist)">Pospiviroid impedichrysanthemi</a> (former name Chrysanthemum stunt viroid<a href="#cite_note-23">[23]</a>); (CSVd); accession V01107, genome length 356nt</li> <li><a href="/w/index.php?title=Pospiviroid_apicimpeditum&action=edit&redlink=1" class="new" title="Pospiviroid apicimpeditum (page does not exist)">Pospiviroid apicimpeditum</a> (former name Tomato apical stunt viroid<a href="#cite_note-24">[24]</a>); (TASVd); accession K00818, genome length 360nt</li> <li><a href="/w/index.php?title=Pospiviroid_alphairesinis&action=edit&redlink=1" class="new" title="Pospiviroid alphairesinis (page does not exist)">Pospiviroid alphairesinis</a> (former name Iresine 1 viroid<a href="#cite_note-25">[25]</a>); (IrVd-1); accession X95734, genome length 370nt</li> <li><a href="/w/index.php?title=Pospiviroid_latenscolumneae&action=edit&redlink=1" class="new" title="Pospiviroid latenscolumneae (page does not exist)">Pospiviroid latenscolumneae</a> (former name Columnea latent viroid<a href="#cite_note-26">[26]</a>); (CLVd); accession X15663, genome length 370nt</li> <li><a href="/w/index.php?title=Pospiviroid_latensportulacae&action=edit&redlink=1" class="new" title="Pospiviroid latensportulacae (page does not exist)">Pospiviroid latensportulacae</a> (former name Pospiviroid plvd<a href="#cite_note-27">[27]</a>)</li> <li><a href="/w/index.php?title=Pospiviroid_parvicapsici&action=edit&redlink=1" class="new" title="Pospiviroid parvicapsici (page does not exist)">Pospiviroid parvicapsici</a> (former name Pepper chat fruit viroid<a href="#cite_note-28">[28]</a>)</li></ul></li> <li>Genus <a href="/wiki/Hostuviroid" title="Hostuviroid">Hostuviroid</a>; type species: <a href="/wiki/Hop_stunt_viroid" title="Hop stunt viroid">Hostuviroid impedihumuli</a> (former name Hop stunt viroid<a href="#cite_note-29">[29]</a>); 294–303 nt<a href="#cite_note-nt-19">[19]</a> <ul><li><a href="/w/index.php?title=Hostuviroid_latensdahliae&action=edit&redlink=1" class="new" title="Hostuviroid latensdahliae (page does not exist)">Hostuviroid latensdahliae</a> (former name Dahlia latent viroid<a href="#cite_note-30">[30]</a>)</li></ul></li> <li>Genus <a href="/wiki/Cocadviroid" class="mw-redirect" title="Cocadviroid">Cocadviroid</a>; type species: <a href="/wiki/Coconut_cadang-cadang_viroid" class="mw-redirect" title="Coconut cadang-cadang viroid">Cocadviroid cadangi</a> (former name Coconut cadang-cadang viroid<a href="#cite_note-31">[31]</a>); 246–247 nt<a href="#cite_note-nt-19">[19]</a> <ul><li><a href="/w/index.php?title=Cocadviroid_tinangajae&action=edit&redlink=1" class="new" title="Cocadviroid tinangajae (page does not exist)">Cocadviroid tinangajae</a> (former name Coconut tinangaja viroid<a href="#cite_note-32">[32]</a>); (CTiVd); accession M20731, genome length 254nt</li> <li><a href="/wiki/Hop_latent_viroid" title="Hop latent viroid">Cocadviroid latenshumuli</a> (former name Hop latent viroid<a href="#cite_note-33">[33]</a>); (HLVd); accession X07397, genome length 256nt</li> <li><a href="/w/index.php?title=Cocadviroid_rimocitri&action=edit&redlink=1" class="new" title="Cocadviroid rimocitri (page does not exist)">Cocadviroid rimocitri</a> (former names Citrus bark cracking viroid, Citrus IV viroid<a href="#cite_note-34">[34]</a>); (CVd-IV); accession X14638, genome length 284nt</li></ul></li> <li>Genus <a href="/wiki/Apscaviroid" title="Apscaviroid">Apscaviroid</a>; type species: <a href="/w/index.php?title=Apscaviroid_cicatricimali&action=edit&redlink=1" class="new" title="Apscaviroid cicatricimali (page does not exist)">Apscaviroid cicatricimali</a> (former name Apple scar skin viroid<a href="#cite_note-35">[35]</a>); 329–334 nt<a href="#cite_note-nt-19">[19]</a> <ul><li><a href="/w/index.php?title=Citrus_III_viroid&action=edit&redlink=1" class="new" title="Citrus III viroid (page does not exist)">Citrus III viroid</a>; (CVd-III); accession AF184147, genome length 294nt</li> <li><a href="/w/index.php?title=Apscaviroid_fossulamali&action=edit&redlink=1" class="new" title="Apscaviroid fossulamali (page does not exist)">Apscaviroid fossulamali</a> (former name Apple dimple fruit viroid<a href="#cite_note-36">[36]</a>); (ADFVd); accession X99487, genome length 306nt</li> <li><a href="/wiki/Grapevine_yellow_speckle_viroid_1" title="Grapevine yellow speckle viroid 1">Apscaviroid alphaflavivitis</a> (former name Grapevine yellow speckle viroid 1<a href="#cite_note-37">[37]</a>); (GVYSd-1); accession X06904, genome length 367nt</li> <li><a href="/w/index.php?title=Apscaviroid_betaflavivitis&action=edit&redlink=1" class="new" title="Apscaviroid betaflavivitis (page does not exist)">Apscaviroid betaflavivitis</a> (former name Grapevine yellow speckle viroid 2<a href="#cite_note-38">[38]</a>); (GVYSd-2); accession J04348, genome length 363nt</li> <li><a href="/w/index.php?title=Apscaviroid_curvifoliumcitri&action=edit&redlink=1" class="new" title="Apscaviroid curvifoliumcitri (page does not exist)">Apscaviroid curvifoliumcitri</a> (former name Citrus bent leaf viroid<a href="#cite_note-39">[39]</a>); (CBLVd); accession M74065, genome length 318nt</li> <li><a href="/w/index.php?title=Apscaviroid_pustulapyri&action=edit&redlink=1" class="new" title="Apscaviroid pustulapyri (page does not exist)">Apscaviroid pustulapyri</a> (former name Pear blister canker viroid<a href="#cite_note-40">[40]</a>); (PBCVd); accession D12823, genome length 315nt</li> <li><a href="/wiki/Australian_grapevine_viroid" title="Australian grapevine viroid">Apscaviroid austravitis</a> (former name Australian grapevine viroid<a href="#cite_note-41">[41]</a>); (AGVd); accession X17101, genome length 369nt</li> <li><a href="/w/index.php?title=Apscaviroid_maculamali&action=edit&redlink=1" class="new" title="Apscaviroid maculamali (page does not exist)">Apscaviroid maculamali</a> (former name Apscaviroid aclsvd<a href="#cite_note-42">[42]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_etacitri&action=edit&redlink=1" class="new" title="Apscaviroid etacitri (page does not exist)">Apscaviroid etacitri</a> (former name Apscaviroid cvd-VII<a href="#cite_note-43">[43]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_dendrobii&action=edit&redlink=1" class="new" title="Apscaviroid dendrobii (page does not exist)">Apscaviroid dendrobii</a> (former name Apscaviroid dvd<a href="#cite_note-44">[44]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_latensvitis&action=edit&redlink=1" class="new" title="Apscaviroid latensvitis (page does not exist)">Apscaviroid latensvitis</a> (former name Apscaviroid glvd<a href="#cite_note-45">[45]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_litchis&action=edit&redlink=1" class="new" title="Apscaviroid litchis (page does not exist)">Apscaviroid litchis</a> (former name Apscaviroid lvd<a href="#cite_note-46">[46]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_latenspruni&action=edit&redlink=1" class="new" title="Apscaviroid latenspruni (page does not exist)">Apscaviroid latenspruni</a> (former name Apscaviroid plvd-I<a href="#cite_note-47">[47]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_diospyri&action=edit&redlink=1" class="new" title="Apscaviroid diospyri (page does not exist)">Apscaviroid diospyri</a> (former name Apscaviroid pvd<a href="#cite_note-48">[48]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_betadiospyri&action=edit&redlink=1" class="new" title="Apscaviroid betadiospyri (page does not exist)">Apscaviroid betadiospyri</a> (former name Apscaviroid pvd-2<a href="#cite_note-49">[49]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_nanocitri&action=edit&redlink=1" class="new" title="Apscaviroid nanocitri (page does not exist)">Apscaviroid nanocitri</a> (former name Citrus dwarfing viroid<a href="#cite_note-50">[50]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_epsiloncitri&action=edit&redlink=1" class="new" title="Apscaviroid epsiloncitri (page does not exist)">Apscaviroid epsiloncitri</a> (former name Citrus viroid V<a href="#cite_note-51">[51]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_zetacitri&action=edit&redlink=1" class="new" title="Apscaviroid zetacitri (page does not exist)">Apscaviroid zetacitri</a> (former name Citrus viroid VI<a href="#cite_note-52">[52]</a>)</li> <li><a href="/w/index.php?title=Apscaviroid_japanvitis&action=edit&redlink=1" class="new" title="Apscaviroid japanvitis (page does not exist)">Apscaviroid japanvitis</a><a href="#cite_note-53">[53]</a></li></ul></li> <li>Genus <a href="/wiki/Coleviroid" class="mw-redirect" title="Coleviroid">Coleviroid</a>; type species: <a href="/w/index.php?title=Coleviroid_alphacolei&action=edit&redlink=1" class="new" title="Coleviroid alphacolei (page does not exist)">Coleviroid alphacolei</a> (former name Coleus blumei viroid 1<a href="#cite_note-54">[54]</a>); (CbVd-1); 248–251 nt<a href="#cite_note-nt-19">[19]</a> <ul><li><a href="/w/index.php?title=Coleviroid_betacolei&action=edit&redlink=1" class="new" title="Coleviroid betacolei (page does not exist)">Coleviroid betacolei</a> (former name Coleus blumei viroid 2<a href="#cite_note-55">[55]</a>); (CbVd-2); accession X95365, genome length 301nt</li> <li><a href="/w/index.php?title=Coleviroid_gammacolei&action=edit&redlink=1" class="new" title="Coleviroid gammacolei (page does not exist)">Coleviroid gammacolei</a> (former name Coleus blumei viroid 3<a href="#cite_note-56">[56]</a>); (CbVd-3); accession X95364, genome length 361nt</li> <li><a href="/w/index.php?title=Coleviroid_epsiloncolei&action=edit&redlink=1" class="new" title="Coleviroid epsiloncolei (page does not exist)">Coleviroid epsiloncolei</a> (former name Coleviroid cbvd-5<a href="#cite_note-57">[57]</a>)</li> <li><a href="/w/index.php?title=Coleviroid_zetacolei&action=edit&redlink=1" class="new" title="Coleviroid zetacolei (page does not exist)">Coleviroid zetacolei</a> (former name Coleviroid cbvd-6<a href="#cite_note-58">[58]</a>)</li></ul></li></ul></li> <li>Family <a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a>: autocatalytic clevage<a href="#cite_note-Lee-5">[5]</a> <ul><li>Genus <a href="/wiki/Avsunviroid" class="mw-redirect" title="Avsunviroid">Avsunviroid</a>; type species: <a href="/wiki/Avocado_sunblotch_viroid" title="Avocado sunblotch viroid">Avsunviroid albamaculaperseae</a> (former name Avocado sunblotch viroid<a href="#cite_note-59">[59]</a>); 246–251 nt<a href="#cite_note-nt-19">[19]</a></li> <li>Genus <a href="/wiki/Pelamoviroid" class="mw-redirect" title="Pelamoviroid">Pelamoviroid</a>; type species: <a href="/wiki/Peach_latent_mosaic_viroid" title="Peach latent mosaic viroid">Pelamoviroid latenspruni </a> (former name Peach latent mosaic viroid<a href="#cite_note-60">[60]</a>); 335–351 nt<a href="#cite_note-nt-19">[19]</a> <ul><li><a href="/w/index.php?title=Pelamoviroid_maculachrysanthemi&action=edit&redlink=1" class="new" title="Pelamoviroid maculachrysanthemi (page does not exist)">Pelamoviroid maculachrysanthemi</a> (former name Chrysanthemum chlorotic mottle viroid<a href="#cite_note-61">[61]</a>)</li> <li><a href="/w/index.php?title=Pelamoviroid_malleusmali&action=edit&redlink=1" class="new" title="Pelamoviroid malleusmali (page does not exist)">Pelamoviroid malleusmali</a> (former name Apple hammerhead viroid<a href="#cite_note-62">[62]</a>)</li></ul></li> <li>Genus <a href="/wiki/Elaviroid" class="mw-redirect" title="Elaviroid">Elaviroid</a>; type species: <a href="/w/index.php?title=Elaviroid_latensmelongenae&action=edit&redlink=1" class="new" title="Elaviroid latensmelongenae (page does not exist)">Elaviroid latensmelongenae</a> (former name Eggplant latent viroid<a href="#cite_note-63">[63]</a>); 332–335 nt<a href="#cite_note-nt-19">[19]</a></li></ul></li></ul> <div class="mw-heading mw-heading2"><h2 id="Transmission_and_replication">Transmission and replication</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=2" title="Edit section: Transmission and replication">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Viroids-_how_it_do.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Viroids-_how_it_do.gif/250px-Viroids-_how_it_do.gif" decoding="async" width="220" height="124" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Viroids-_how_it_do.gif/330px-Viroids-_how_it_do.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/05/Viroids-_how_it_do.gif/500px-Viroids-_how_it_do.gif 2x" data-file-width="1500" data-file-height="843" /></a><figcaption>The reproduction mechanism of a typical viroid. Leaf contact transmits the viroid. The viroid enters the cell via its <a href="/wiki/Plasmodesmata" class="mw-redirect" title="Plasmodesmata">plasmodesmata</a>. RNA polymerase II catalyzes rolling-circle synthesis of new viroids.</figcaption></figure> Viroids are only known to infect plants, and infectious viroids can be transmitted to new plant hosts by <a href="/wiki/Aphid" title="Aphid">aphids</a>, by cross contamination following mechanical damage to plants as a result of horticultural or agricultural practices, or from plant to plant by leaf contact.<a href="#cite_note-nt-19">[19]</a><a href="#cite_note-64">[64]</a> Upon infection, viroids replicate in the nucleus (Pospiviroidae) or <a href="/wiki/Chloroplasts" class="mw-redirect" title="Chloroplasts">chloroplasts</a> (Avsunviroidae) of plant cells in three steps(what are the steps?) through an RNA-based mechanism. They require <a href="/wiki/RNA_polymerase_II" title="RNA polymerase II">RNA polymerase II</a>, a host cell enzyme normally associated with synthesis of <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> from DNA, which instead catalyzes "<a href="/wiki/Rolling_circle" class="mw-redirect" title="Rolling circle">rolling circle</a>" synthesis of new RNA using the viroid as template.<a href="#cite_note-pmid22719735-65">[65]</a> Unlike plant viruses which produce <a href="/wiki/Movement_proteins" class="mw-redirect" title="Movement proteins">movement proteins</a>, viroids are entirely passive, relying entirely on the host. This is useful in the study of RNA kinetics in plants.<a href="#cite_note-pmid16078879-16">[16]</a> <div class="mw-heading mw-heading2"><h2 id="RNA_silencing">RNA silencing</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=3" title="Edit section: RNA silencing">edit</a>]</div> There has long been uncertainty over how viroids induce <a href="/wiki/Symptom" class="mw-redirect" title="Symptom">symptoms</a> in plants without encoding any <a href="/wiki/Protein" title="Protein">protein</a> products within their sequences.<a href="#cite_note-pmid28738223-66">[66]</a> Evidence suggests that <a href="/wiki/RNA_silencing" title="RNA silencing">RNA silencing</a> is involved in the process. First, changes to the viroid <a href="/wiki/Genome" title="Genome">genome</a> can dramatically alter its <a href="/wiki/Virulence" title="Virulence">virulence</a>.<a href="#cite_note-pmid1546460-67">[67]</a> This reflects the fact that any <a href="/wiki/SiRNA" class="mw-redirect" title="SiRNA">siRNAs</a> produced would have less complementary <a href="/wiki/Base_pairing" class="mw-redirect" title="Base pairing">base pairing</a> with target <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a>. Secondly, <a href="/wiki/SiRNA" class="mw-redirect" title="SiRNA">siRNAs</a> corresponding to sequences from viroid genomes have been isolated from infected plants. Finally, <a href="/wiki/Transgenic" class="mw-redirect" title="Transgenic">transgenic</a> expression of the noninfectious <a href="/wiki/Hairpin_RNA" class="mw-redirect" title="Hairpin RNA">hpRNA</a> of <a href="/wiki/Potato_spindle_tuber_viroid" title="Potato spindle tuber viroid">potato spindle tuber viroid</a> develops all the corresponding viroid-like symptoms.<a href="#cite_note-68">[68]</a> This indicates that when viroids replicate via a double stranded intermediate <a href="/wiki/RNA" title="RNA">RNA</a>, they are targeted by a <a href="/wiki/Dicer" title="Dicer">dicer</a> enzyme and cleaved into siRNAs that are then loaded onto the <a href="/wiki/RNA-induced_silencing_complex" title="RNA-induced silencing complex">RNA-induced silencing complex</a>. The viroid siRNAs contain sequences capable of complementary base pairing with the plant's own messenger RNAs, and induction of degradation or inhibition of translation causes the classic viroid symptoms.<a href="#cite_note-69">[69]</a> <div class="mw-heading mw-heading2"><h2 id="Viroid-like_elements">Viroid-like elements</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=4" title="Edit section: Viroid-like elements">edit</a>]</div> "Viroid-like elements" refer to pieces of covalently closed circular (ccc) RNA molecules that do not share the viroid's lifecycle. The category encompasses satellite RNAs (including small plant satRNAs "<a href="/wiki/Virusoids" class="mw-redirect" title="Virusoids">virusoids</a>", fungal "<a href="/wiki/Ambivirus" class="mw-redirect" title="Ambivirus">ambivirus</a>", and the much larger <a href="/wiki/Hepatitis_D" title="Hepatitis D">HDV</a>-like <a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a>) and "retroviroids". Most of them also carry some type of a <a href="/wiki/Ribozyme" title="Ribozyme">ribozyme</a>.<a href="#cite_note-Lee-5">[5]</a> <div class="mw-heading mw-heading3"><h3 id="Viroid-like_satellite_RNAs">Viroid-like satellite RNAs</h3>[<a href="/w/index.php?title=Viroid&action=edit&section=5" title="Edit section: Viroid-like satellite RNAs">edit</a>]</div> Viroid-like <a href="/wiki/Satellite_RNA" class="mw-redirect" title="Satellite RNA">satellite RNAs</a> are infectious circular RNA molecules that depend on a carrier virus to reproduce, being carried in their <a href="/wiki/Capsid" title="Capsid">capsids</a>. Like Avsunviroidae, however, they are capable of self-clevage.<a href="#cite_note-pmid35183574-70">[70]</a> <div class="mw-heading mw-heading3"><h3 id="Ambiviruses">Ambiviruses</h3>[<a href="/w/index.php?title=Viroid&action=edit&section=6" title="Edit section: Ambiviruses">edit</a>]</div> "Ambiviruses" are <a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">mobile genetic elements</a> that were recently (2020s) discovered in <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a>. Their RNA <a href="/wiki/Genome" title="Genome">genomes</a> are circular, circa 5 kb in length. One of at least two open reading frames encodes a viral RNA-directed <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a>, that firmly places "ambiviruses" into <a href="/wiki/Riboviria" title="Riboviria">ribovirian</a> kingdom <a href="/wiki/Orthornavirae" title="Orthornavirae">Orthornavirae</a>; a separate phylum <a href="/wiki/Ambiviricota" title="Ambiviricota">Ambiviricota</a> has been established since the 2023 ICTV Virus Taxonomy Release because of the unique features of encoding RNA-directed RNA polymerases but also having divergent <a href="/wiki/Ribozyme" title="Ribozyme">ribozymes</a> in various combinations in both sense and antisense orientation – the detection of circular forms in both sense orientations suggest that "ambiviruses" use rolling circle replication for propagation.<a href="#cite_note-71">[71]</a><a href="#cite_note-72">[72]</a><a href="#cite_note-:2-73">[73]</a> <div class="mw-heading mw-heading3"><h3 id="Retroviroids">Retroviroids</h3>[<a href="/w/index.php?title=Viroid&action=edit&section=7" title="Edit section: Retroviroids">edit</a>]</div> "Retroviroids", more formally "retroviroid-like elements", are viroid-like circular RNA sequences that are also found with homologous copies in the <a href="/wiki/DNA" title="DNA">DNA</a> genome of the host.<a href="#cite_note-daros-74">[74]</a> The only types found are closely related to the original "carnation small viroid-like RNA" (CarSV).<a href="#cite_note-pmid11172112-75">[75]</a><a href="#cite_note-hegedus-76">[76]</a> These elements may act as a <a href="/wiki/Sequence_homology" title="Sequence homology">homologous</a> substrate upon which <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a> may occur and are linked to <a href="/wiki/DNA_repair#Double-strand_breaks" title="DNA repair">double-stranded break repair</a>.<a href="#cite_note-hegedus-76">[76]</a><a href="#cite_note-Truong-77">[77]</a> These elements are dubbed retroviroids as the homologous DNA is generated by <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a> that is encoded by <a href="/wiki/Retrovirus" title="Retrovirus">retroviruses</a>.<a href="#cite_note-Flores-2014-78">[78]</a><a href="#cite_note-79">[79]</a> They are neither true viroids nor viroid-like <a href="/wiki/Satellite_RNA" class="mw-redirect" title="Satellite RNA">satellite RNAs</a>: there is no extracellular form of these elements; instead, they are spread only through pollen or egg-cells.<a href="#cite_note-pmid35183574-70">[70]</a> They appear to co-occur with a <a href="/wiki/Pararetrovirus" class="mw-redirect" title="Pararetrovirus">pararetrovirus</a>.<a href="#cite_note-80">[80]</a> <div class="mw-heading mw-heading3"><h3 id="Obelisks">Obelisks</h3>[<a href="/w/index.php?title=Viroid&action=edit&section=8" title="Edit section: Obelisks">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Obelisk_(biology)" title="Obelisk (biology)">Obelisk (biology)</a></div> After applying <a href="/wiki/Metatranscriptomics" title="Metatranscriptomics">metatranscriptomics</a> – the computer-aided search for RNA sequences and their analysis – biologists reported in January 2024 the discovery of "<a href="/wiki/Obelisk_(life_form)" class="mw-redirect" title="Obelisk (life form)">obelisks</a>", a new class of viroid-like elements, and "oblins", their related group of proteins, in the <a href="/wiki/Human_microbiome" title="Human microbiome">human microbiome</a>. Given that the RNA sequences recovered do not have <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologies</a> in any other known life form, the researchers suggest that the obelisks are distinct from viruses, viroids and viroid-like entities, and thus form an entirely new class of organisms.<a href="#cite_note-SA-20240129-81">[81]</a><a href="#cite_note-BRX-20240121-82">[82]</a> <div class="mw-heading mw-heading2"><h2 id="RNA_world_hypothesis">RNA world hypothesis</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=9" title="Edit section: RNA world hypothesis">edit</a>]</div> Diener's 1989 hypothesis<a href="#cite_note-83">[83]</a> had proposed that the unique properties of viroids make them more plausible macromolecules than <a href="/wiki/Intron" title="Intron">introns</a>, or other RNAs considered in the past as possible "living relics" of a hypothetical, pre-cellular <a href="/wiki/RNA_world" title="RNA world">RNA world</a>. If so, viroids have assumed significance beyond plant virology for evolutionary theory, because their properties make them more plausible candidates than other RNAs to perform crucial steps in the evolution of life from inanimate matter (abiogenesis). Diener's hypothesis was mostly forgotten until 2014, when it was resurrected in a review article by Flores et al.,<a href="#cite_note-Flores-2014-78">[78]</a> in which the authors summarized Diener's evidence supporting his hypothesis as: <ol><li>Viroids' small size, imposed by error-prone replication.</li> <li>Their high <a href="/wiki/Guanine" title="Guanine">guanine</a> and <a href="/wiki/Cytosine" title="Cytosine">cytosine</a> content, which increases stability and replication fidelity.</li> <li>Their circular structure, which assures complete replication without genomic tags.</li> <li>Existence of structural periodicity, which permits modular assembly into enlarged genomes.</li> <li>Their lack of protein-coding ability, consistent with a <a href="/wiki/Ribosome" title="Ribosome">ribosome</a>-free habitat.</li> <li>Replication mediated in some by <a href="/wiki/Ribozymes" class="mw-redirect" title="Ribozymes">ribozymes</a>—the fingerprint of the RNA world.</li></ol> The presence, in extant cells, of RNAs with molecular properties predicted for RNAs of the RNA world constitutes another powerful argument supporting the RNA world hypothesis. However, the origins of viroids themselves from this RNA world has been cast into doubt by several factors, including the discovery of <a href="/wiki/Retrozyme" title="Retrozyme">retrozymes</a> (a family of <a href="/wiki/Retrotransposon" title="Retrotransposon">retrotransposon</a> likely representing their ancestors) and their complete absence from organisms outside of the <a href="/wiki/Plant" title="Plant">plants</a> (especially their complete absence from <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a> including <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>).<a href="#cite_note-:3-12">[12]</a><a href="#cite_note-:0-13">[13]</a><a href="#cite_note-:1-14">[14]</a> However, recent studies suggest that the diversity of viroids and others viroid-like elements is broader than previously thought and that it would not be limited to plants, encompassing even the <a href="/wiki/Prokaryotes" class="mw-redirect" title="Prokaryotes">prokaryotes</a>. Matches between viroid cccRNAs and <a href="/wiki/CRISPR" title="CRISPR">CRISPR</a> spacers suggest that some of them might replicate in prokaryotes.<a href="#cite_note-Lee-5">[5]</a> <div class="mw-heading mw-heading2"><h2 id="Control">Control</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=10" title="Edit section: Control">edit</a>]</div> The development of tests based on <a href="/wiki/ELISA" title="ELISA">ELISA</a>, <a href="/wiki/Polymerase_chain_reaction" title="Polymerase chain reaction">PCR</a>, and <a href="/wiki/Nucleic_acid_hybridization" title="Nucleic acid hybridization">nucleic acid hybridization</a> has allowed for rapid and inexpensive detection of known viroids in <a href="/wiki/Agricultural_biosecurity_inspection" class="mw-redirect" title="Agricultural biosecurity inspection">biosecurity inspections</a>, <a href="/wiki/Phytosanitary_inspection" class="mw-redirect" title="Phytosanitary inspection">phytosanitary inspections</a>, and <a href="/wiki/Agricultural_quarantine" class="mw-redirect" title="Agricultural quarantine">quarantine</a>.<a href="#cite_note-pmid26047558-84">[84]</a> <div class="mw-heading mw-heading2"><h2 id="History">History</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=11" title="Edit section: History">edit</a>]</div> In the 1920s, symptoms of a previously unknown potato disease were noticed in New York and New Jersey fields. Because tubers on affected plants become elongated and misshapen, they named it the potato spindle tuber disease.<a href="#cite_note-85">[85]</a> The symptoms appeared on plants onto which pieces from affected plants had been budded—indicating that the disease was caused by a transmissible pathogenic agent. A <a href="/wiki/Fungus" title="Fungus">fungus</a> or <a href="/wiki/Bacteria" title="Bacteria">bacterium</a> could not be found consistently associated with symptom-bearing plants, however, and therefore, it was assumed the disease was caused by a virus. Despite numerous attempts over the years to isolate and purify the assumed virus, using increasingly sophisticated methods, these were unsuccessful when applied to extracts from potato spindle tuber disease-afflicted plants.<a href="#cite_note-ARS_timeline-86">[86]</a> The first recognized viroid, the pathogenic agent of the <a href="/wiki/Potato_spindle_tuber_viroid" title="Potato spindle tuber viroid">potato spindle tuber disease</a>, was discovered, initially molecularly characterized, and named by <a href="/wiki/Theodor_Otto_Diener" title="Theodor Otto Diener">Theodor Otto Diener</a>, a plant pathologist at the <a href="/wiki/Beltsville_Agricultural_Research_Center" class="mw-redirect" title="Beltsville Agricultural Research Center">U.S. Department of Agriculture's Research Center in Beltsville, Maryland</a>, in 1971.<a href="#cite_note-pmid5095900-87">[87]</a><a href="#cite_note-ARS_timeline-86">[86]</a> He showed that the agent was not a virus, but a totally unexpected novel type of pathogen, 1/80th the size of typical viruses, for which he proposed the term "viroid".<a href="#cite_note-pmid5095900-87">[87]</a> This agent is now called the potato spindle tuber viroid (PSTVd). The <a href="/wiki/Citrus_exocortis" title="Citrus exocortis">Citrus exocortis viroid</a> (CEVd) was discovered soon thereafter, and together understanding of PSTVd and CEVd shaped the concept of the viroid.<a href="#cite_note-pmid16078879-16">[16]</a> Parallel to agriculture-directed studies, more basic scientific research elucidated many of viroids' physical, chemical, and macromolecular properties. Viroids were shown to consist of short stretches (a few hundred nucleotides) of single-stranded RNA and, unlike viruses, did not have a protein coat. Viroids are extremely small, from 246 to 467 nucleotides, smaller than other infectious plant pathogens; they thus consist of fewer than 10,000 atoms. In comparison, the genomes of the smallest known viruses capable of causing an infection by themselves are around 2,000 nucleotides long.<a href="#cite_note-Pommerville-88">[88]</a> In 1976, Sanger et al.<a href="#cite_note-pmid1069269-89">[89]</a> presented evidence that potato spindle tuber viroid is a "single-stranded, covalently closed, circular RNA molecule, existing as a highly base-paired rod-like structure"—believed to be the first such molecule described. Circular RNA, unlike linear RNA, forms a covalently closed continuous loop, in which the 3' and 5' ends present in linear RNA molecules have been joined. Sanger et al. also provided evidence for the true circularity of viroids by finding that the RNA could not be phosphorylated at the 5' terminus. In other tests, they failed to find even one free 3' end, which ruled out the possibility of the molecule having two 3' ends. Viroids thus are true circular RNAs.<a href="#cite_note-pmid33460914-90">[90]</a> The single-strandedness and circularity of viroids was confirmed by electron microscopy,<a href="#cite_note-pmid4728831-91">[91]</a> The complete nucleotide sequence of potato spindle tuber viroid was determined in 1978.<a href="#cite_note-92">[92]</a> PSTVd was the first pathogen of a eukaryotic organism for which the complete molecular structure has been established. Over thirty plant diseases have since been identified as viroid-, not virus-caused, as had been assumed.<a href="#cite_note-Pommerville-88">[88]</a><a href="#cite_note-Hammond_2006-93">[93]</a> Four additional viroids or viroid-like RNA particles were discovered between 2009 and 2015.<a href="#cite_note-pmid26047558-84">[84]</a> In 2014, <a href="/wiki/New_York_Times" class="mw-redirect" title="New York Times">New York Times</a> science writer Carl Zimmer published a popularized piece that mistakenly credited Flores et al. with the virioid - RNA world hypothesis' original conception.<a href="#cite_note-Zimmer-94">[94]</a> In January 2024, biologists reported the discovery of "<a href="/wiki/Obelisk_(life_form)" class="mw-redirect" title="Obelisk (life form)">obelisks</a>", a new class of viroid-like elements, and "oblins", their related group of proteins, in the <a href="/wiki/Human_microbiome" title="Human microbiome">human microbiome</a>.<a href="#cite_note-SA-20240129-81">[81]</a><a href="#cite_note-BRX-20240121-82">[82]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=12" title="Edit section: See also">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1266661725">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output .portalbox-link{display:table-cell;padding:0.2em 0.2em 0.2em 0.3em;vertical-align:middle}@media(min-width:720px){.mw-parser-output .portalleft{margin:0.5em 1em 0.5em 0}.mw-parser-output .portalright{clear:right;float:right;margin:0.5em 0 0.5em 1em}}</style><ul role="navigation" aria-label="Portals" class="noprint portalbox portalborder portalright"> <li class="portalbox-entry"><a href="/wiki/File:Issoria_lathonia.jpg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/32px-Issoria_lathonia.jpg" decoding="async" width="32" height="23" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/48px-Issoria_lathonia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/64px-Issoria_lathonia.jpg 2x" data-file-width="629" data-file-height="445" /></a><a href="/wiki/Portal:Biology" title="Portal:Biology">Biology portal</a></li><li class="portalbox-entry"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/20px-WHO_Rod.svg.png" decoding="async" width="12" height="28" class="mw-file-element" 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hypothesis</a></li> <li><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite (biology)</a></li> <li><a href="/wiki/Virus" title="Virus">Virus</a></li> <li><a href="/wiki/Virus_classification" title="Virus classification">Virus classification</a></li> <li><a href="/wiki/Virusoid" title="Virusoid">Virusoid</a></li></ul></div> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=13" title="Edit section: References">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 35em;"> <ol class="references"> <li id="cite_note-Navarro-1"><a href="#cite_ref-Navarro_1-0">^</a> <style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFNavarroFloresDi_Serio2021" class="citation journal cs1">Navarro B, Flores R, Di Serio F (29 September 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Annual Review of Virology. 8 (1): 305–325. <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-virology-091919-092331">10.1146/annurev-virology-091919-092331</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/2327-056X">2327-056X</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/34255541">34255541</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+Virology&rft.atitle=Advances+in+Viroid-Host+Interactions&rft.volume=8&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E305-%3C%2Fspan%3E325&rft.date=2021-09-29&rft.issn=2327-056X&rft_id=info%3Apmid%2F34255541&rft_id=info%3Adoi%2F10.1146%2Fannurev-virology-091919-092331&rft.aulast=Navarro&rft.aufirst=Beatriz&rft.au=Flores%2C+Ricardo&rft.au=Di+Serio%2C+Francesco&rft_id=https%3A%2F%2Fdoi.org%2F10.1146%252Fannurev-virology-091919-092331&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-2"><a href="#cite_ref-2">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFDi_SerioOwensLiMatoušek2020" class="citation web cs1">Di Serio F, Owens RA, Li SF, Matoušek J, Pallás V, Randles JW, Sano T, Verhoeven JT, Vidalakis G, Flores R (November 2020). 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Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Pospiviroid+latenscolumneae%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304524%26taxon_name%3DPospiviroid%2520latenscolumneae&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-27"><a href="#cite_ref-27">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313877&taxon_name=Pospiviroid%20latensportulacae">"Pospiviroid latensportulacae: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Pospiviroid+latensportulacae%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313877%26taxon_name%3DPospiviroid%2520latensportulacae&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-28"><a href="#cite_ref-28">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304526&taxon_name=Pospiviroid%20parvicapsici">"Pospiviroid parvicapsici: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Pospiviroid+parvicapsici%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304526%26taxon_name%3DPospiviroid%2520parvicapsici&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-29"><a href="#cite_ref-29">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304520&taxon_name=Hostuviroid%20impedihumuli">"Hostuviroid impedihumuli: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Hostuviroid+impedihumuli%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304520%26taxon_name%3DHostuviroid%2520impedihumuli&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-30"><a href="#cite_ref-30">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304519&taxon_name=Hostuviroid%20latensdahliae">"Hostuviroid latensdahliae: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Hostuviroid+latensdahliae%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304519%26taxon_name%3DHostuviroid%2520latensdahliae&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-31"><a href="#cite_ref-31">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304511&taxon_name=Cocadviroid%20cadangi">"Cocadviroid cadangi: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Cocadviroid+cadangi%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304511%26taxon_name%3DCocadviroid%2520cadangi&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-32"><a href="#cite_ref-32">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304512&taxon_name=Cocadviroid%20tinangajae">"Cocadviroid tinangajae: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Cocadviroid+tinangajae%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304512%26taxon_name%3DCocadviroid%2520tinangajae&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-33"><a href="#cite_ref-33">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304513&taxon_name=Cocadviroid%20latenshumuli">"Cocadviroid latenshumuli: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Cocadviroid+latenshumuli%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304513%26taxon_name%3DCocadviroid%2520latenshumuli&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-34"><a href="#cite_ref-34">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304510&taxon_name=Cocadviroid%20rimocitri">"Cocadviroid rimocitri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Cocadviroid+rimocitri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304510%26taxon_name%3DCocadviroid%2520rimocitri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-35"><a href="#cite_ref-35">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304500&taxon_name=Apscaviroid%20cicatricimali">"Apscaviroid cicatricimali: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+cicatricimali%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304500%26taxon_name%3DApscaviroid%2520cicatricimali&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-36"><a href="#cite_ref-36">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304499&taxon_name=Apscaviroid%20fossulamali">"Apscaviroid fossulamali: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+fossulamali%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304499%26taxon_name%3DApscaviroid%2520fossulamali&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-37"><a href="#cite_ref-37">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304506&taxon_name=Apscaviroid%20alphaflavivitis">"Apscaviroid alphaflavivitis: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+alphaflavivitis%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304506%26taxon_name%3DApscaviroid%2520alphaflavivitis&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-38"><a href="#cite_ref-38">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304507&taxon_name=Apscaviroid%20betaflavivitis">"Apscaviroid betaflavivitis: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+betaflavivitis%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304507%26taxon_name%3DApscaviroid%2520betaflavivitis&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-39"><a href="#cite_ref-39">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304502&taxon_name=Apscaviroid%20curvifoliumcitri">"Apscaviroid curvifoliumcitri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+curvifoliumcitri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304502%26taxon_name%3DApscaviroid%2520curvifoliumcitri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-40"><a href="#cite_ref-40">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304508&taxon_name=Apscaviroid%20pustulapyri">"Apscaviroid pustulapyri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+pustulapyri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304508%26taxon_name%3DApscaviroid%2520pustulapyri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-41"><a href="#cite_ref-41">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304501&taxon_name=Apscaviroid%20austravitis">"Apscaviroid austravitis: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+austravitis%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304501%26taxon_name%3DApscaviroid%2520austravitis&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-42"><a href="#cite_ref-42">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313867&taxon_name=Apscaviroid%20maculamali">"Apscaviroid maculamali: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+maculamali%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313867%26taxon_name%3DApscaviroid%2520maculamali&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-43"><a href="#cite_ref-43">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313868&taxon_name=Apscaviroid%20etacitri">"Apscaviroid etacitri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+etacitri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313868%26taxon_name%3DApscaviroid%2520etacitri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-44"><a href="#cite_ref-44">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313869&taxon_name=Apscaviroid%20dendrobii">"Apscaviroid dendrobii: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+dendrobii%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313869%26taxon_name%3DApscaviroid%2520dendrobii&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-45"><a href="#cite_ref-45">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313870&taxon_name=Apscaviroid%20latensvitis">"Apscaviroid latensvitis: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+latensvitis%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313870%26taxon_name%3DApscaviroid%2520latensvitis&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-46"><a href="#cite_ref-46">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313871&taxon_name=Apscaviroid%20litchis">"Apscaviroid litchis: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+litchis%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313871%26taxon_name%3DApscaviroid%2520litchis&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-47"><a href="#cite_ref-47">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313874&taxon_name=Apscaviroid%20latenspruni">"Apscaviroid latenspruni: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+latenspruni%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313874%26taxon_name%3DApscaviroid%2520latenspruni&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-48"><a href="#cite_ref-48">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313872&taxon_name=Apscaviroid%20diospyri">"Apscaviroid diospyri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+diospyri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313872%26taxon_name%3DApscaviroid%2520diospyri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-49"><a href="#cite_ref-49">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313873&taxon_name=Apscaviroid%20betadiospyri">"Apscaviroid betadiospyri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+betadiospyri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313873%26taxon_name%3DApscaviroid%2520betadiospyri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-50"><a href="#cite_ref-50">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304503&taxon_name=Apscaviroid%20nanocitri">"Apscaviroid nanocitri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+nanocitri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304503%26taxon_name%3DApscaviroid%2520nanocitri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-51"><a href="#cite_ref-51">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304504&taxon_name=Apscaviroid%20epsiloncitri">"Apscaviroid epsiloncitri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+epsiloncitri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304504%26taxon_name%3DApscaviroid%2520epsiloncitri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-52"><a href="#cite_ref-52">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304505&taxon_name=Apscaviroid%20zetacitri">"Apscaviroid zetacitri: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+zetacitri%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304505%26taxon_name%3DApscaviroid%2520zetacitri&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-53"><a href="#cite_ref-53">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202319071&taxon_name=Apscaviroid%20japanvitis">"Apscaviroid japanvitis: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Apscaviroid+japanvitis%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202319071%26taxon_name%3DApscaviroid%2520japanvitis&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-54"><a href="#cite_ref-54">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304515&taxon_name=Coleviroid%20alphacolei">"Coleviroid alphacolei: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Coleviroid+alphacolei%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304515%26taxon_name%3DColeviroid%2520alphacolei&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-55"><a href="#cite_ref-55">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304516&taxon_name=Coleviroid%20betacolei">"Coleviroid betacolei: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Coleviroid+betacolei%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304516%26taxon_name%3DColeviroid%2520betacolei&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-56"><a href="#cite_ref-56">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202304517&taxon_name=Coleviroid%20gammacolei">"Coleviroid gammacolei: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Coleviroid+gammacolei%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202304517%26taxon_name%3DColeviroid%2520gammacolei&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-57"><a href="#cite_ref-57">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313875&taxon_name=Coleviroid%20epsiloncolei">"Coleviroid epsiloncolei: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Coleviroid+epsiloncolei%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313875%26taxon_name%3DColeviroid%2520epsiloncolei&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-58"><a href="#cite_ref-58">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202313876&taxon_name=Coleviroid%20zetacolei">"Coleviroid zetacolei: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Coleviroid+zetacolei%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202313876%26taxon_name%3DColeviroid%2520zetacolei&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-59"><a href="#cite_ref-59">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202302648&taxon_name=Avsunviroid%20albamaculaperseae">"Avsunviroid albamaculaperseae: Taxon Details"</a>. ICTV Taxonomy Browser. Retrieved 9 December 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=ICTV+Taxonomy+Browser&rft.atitle=Avsunviroid+albamaculaperseae%3A+Taxon+Details&rft.au=ICTV&rft_id=https%3A%2F%2Fictv.global%2Ftaxonomy%2Ftaxondetails%3Ftaxnode_id%3D202302648%26taxon_name%3DAvsunviroid%2520albamaculaperseae&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-60"><a href="#cite_ref-60">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFICTV" class="citation web cs1"><a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">ICTV</a>. <a rel="nofollow" class="external text" href="https://ictv.global/taxonomy/taxondetails?taxnode_id=202302653&taxon_name=Pelamoviroid%20latenspruni">"Pelamoviroid latenspruni: Taxon Details"</a>. ICTV Taxonomy Browser. 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"Nucleotide sequence and secondary structure of potato spindle tuber viroid". Nature. 273 (5659): 203–8. <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/1978Natur.273..203G">1978Natur.273..203G</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%2F273203a0">10.1038/273203a0</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/643081">643081</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:19398777">19398777</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Nucleotide+sequence+and+secondary+structure+of+potato+spindle+tuber+viroid&rft.volume=273&rft.issue=5659&rft.pages=%3Cspan+class%3D%22nowrap%22%3E203-%3C%2Fspan%3E8&rft.date=1978-05&rft_id=info%3Adoi%2F10.1038%2F273203a0&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A19398777%23id-name%3DS2CID&rft_id=info%3Apmid%2F643081&rft_id=info%3Abibcode%2F1978Natur.273..203G&rft.aulast=Gross&rft.aufirst=HJ&rft.au=Domdey%2C+H&rft.au=Lossow%2C+C&rft.au=Jank%2C+P&rft.au=Raba%2C+M&rft.au=Alberty%2C+H&rft.au=S%C3%A4nger%2C+HL&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-Hammond_2006-93"><a href="#cite_ref-Hammond_2006_93-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFHammondOwens2006" class="citation journal cs1">Hammond RW, Owens RA (2006). "Viroids: New and Continuing Risks for Horticultural and Agricultural Crops". APSnet Feature Articles. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1094%2FAPSnetFeature-2006-1106">10.1094/APSnetFeature-2006-1106</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=APSnet+Feature+Articles&rft.atitle=Viroids%3A+New+and+Continuing+Risks+for+Horticultural+and+Agricultural+Crops.&rft.date=2006&rft_id=info%3Adoi%2F10.1094%2FAPSnetFeature-2006-1106&rft.aulast=Hammond&rft.aufirst=RW&rft.au=Owens%2C+RA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> <li id="cite_note-Zimmer-94"><a href="#cite_ref-Zimmer_94-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFZimmer2014" class="citation news cs1"><a href="/wiki/Carl_Zimmer" title="Carl Zimmer">Zimmer C</a> (September 25, 2014). <a rel="nofollow" class="external text" href="https://www.nytimes.com/2014/09/25/science/a-tiny-emissary-from-the-ancient-past.html">"A Tiny Emissary From the Ancient Past"</a>. <a href="/wiki/New_York_Times" class="mw-redirect" title="New York Times">New York Times</a>. Retrieved November 22, 2014.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=New+York+Times&rft.atitle=A+Tiny+Emissary+From+the+Ancient+Past&rft.date=2014-09-25&rft.aulast=Zimmer&rft.aufirst=C&rft_id=https%3A%2F%2Fwww.nytimes.com%2F2014%2F09%2F25%2Fscience%2Fa-tiny-emissary-from-the-ancient-past.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AViroid" class="Z3988"> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=Viroid&action=edit&section=14" title="Edit section: External links">edit</a>]</div> <ul><li><a rel="nofollow" class="external text" href="http://www.atsu.edu/faculty/chamberlain/Website/Lects/Prions.htm">Viroids/ATSU</a></li> <li><a rel="nofollow" class="external text" href="https://viroids.org/">ViroidDB</a>, a database of viroids and viroid-like circular RNAs</li></ul> <div class="navbox-styles"><style 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href="/wiki/Self-replication" title="Self-replication">Self-replicating</a> organic structures</div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Life" title="Life">Cellular life</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a></li> <li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li><a href="/wiki/Protist" title="Protist">Protista</a></li></ul></li> <li><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a> <ul><li><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></li> <li><a href="/wiki/Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Virus" title="Virus">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a> <ul><li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li></ul></li> <li><a href="/wiki/DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li> <li><a href="/wiki/Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li> <li><a href="/wiki/Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li> <li><a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a class="mw-selflink selflink">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li> <li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Helper_virus" title="Helper virus">Helper-virus dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;"><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>ssRNA satellite virus</li> <li>dsDNA satellite virus (<a href="/wiki/Virophage" title="Virophage">Virophage</a>)</li> <li>ssDNA satellite virus</li> <li>ssDNA satellite</li> <li>dsRNA satellite</li> <li>ssRNA satellite (<a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>)</li> <li>Satellite-like nucleic acids <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">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/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a> <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Prion" title="Prion">Mammalian prion</a></li> <li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a> self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mobilome" title="Mobilome">Mobilome</a> <ul><li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Genomic_island" title="Genomic island">Genomic island</a></li></ul></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li> <li><a href="/wiki/DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a> <ul><li><a href="/wiki/Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li> <li><a href="/wiki/R-factor" class="mw-redirect" title="R-factor">Resistance</a></li> <li><a href="/wiki/Colicin" title="Colicin">Col</a></li> <li>Degradative</li> <li><a href="/wiki/Virulence_factor" title="Virulence factor">Virulence</a>/<a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti</a></li> <li>Cryptic</li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a> <ul><li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li></ul></li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li> <li><a href="/wiki/Group_II_intron" title="Group II intron">Group II intron</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a> <ul><li><a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear</a></li> <li><a href="/wiki/Circular_chromosome" title="Circular chromosome">Circular</a></li> <li><a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li> <li><a href="/wiki/Genome" title="Genome">Genome</a> <ul><li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Prophage" title="Prophage">Prophage</a></li> <li><a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a> <ul><li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li> <li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Kappa_organism" title="Kappa organism">Kappa organism</a></li> <li>Organs <ul><li><a href="/wiki/Bacteriome" title="Bacteriome">Bacteriome</a></li> <li><a href="/wiki/Trophosome" title="Trophosome">Trophosome</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li>?<a href="/wiki/RNA_world" title="RNA world">RNA life</a> <ul><li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li> <li>†<a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Coacervate" title="Coacervate">Coacervate</a></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a></li> <li><a href="/wiki/Sulphobes" title="Sulphobes">Sulphobe</a></li> <li>Research <ul><li><a href="/wiki/Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Organism" title="Organism">Organism</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a> <ul><li><a href="/wiki/Cell_division" title="Cell division">Cell division</a></li> <li><a href="/wiki/Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Synthetic_virology" title="Synthetic virology">Synthetic virus</a> <ul><li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a></li> <li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li>?<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></li> <li>?<a href="/wiki/Nanobe" title="Nanobe">Nanobe</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235" /></div><div role="navigation" class="navbox" aria-labelledby="Life,_non-cellular_life,_and_comparable_structures112" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231" /><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Life,_non-cellular_life,_and_comparable_structures" title="Template:Life, non-cellular life, and comparable structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Life,_non-cellular_life,_and_comparable_structures" title="Template talk:Life, non-cellular life, and comparable structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Life,_non-cellular_life,_and_comparable_structures" title="Special:EditPage/Template:Life, non-cellular life, and comparable structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Life,_non-cellular_life,_and_comparable_structures112" style="font-size:114%;margin:0 4em"><a href="/wiki/Life" title="Life">Life</a>, <a href="/wiki/Non-cellular_life" title="Non-cellular life">non-cellular life</a>, and comparable structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">"<a href="/wiki/Prokaryote" title="Prokaryote">Prokaryota</a>"</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li></ul></li> <li><a href="/wiki/LUCA" class="mw-redirect" title="LUCA">LUCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li>'<a href="/wiki/Protist" title="Protist">Protista</a>'</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></li> <li><a href="/wiki/Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Virus" title="Virus">Virus</a> <hr /><a class="mw-selflink selflink">Viroid</a> <hr /><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Realms</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Adnaviria" title="Adnaviria">Adnaviria</a></li> <li><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></li> <li><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></li> <li><a href="/wiki/Riboviria" title="Riboviria">Riboviria</a></li> <li><a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a></li> <li><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Unassigned</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Classes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><a href="/wiki/Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Families</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></li> <li><a href="/wiki/Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></li> <li><a href="/wiki/Anelloviridae" title="Anelloviridae">Anelloviridae</a></li> <li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></li> <li><a href="/wiki/Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></li> <li><a href="/wiki/Clavaviridae" title="Clavaviridae">Clavaviridae</a></li> <li><a href="/wiki/Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></li> <li><a href="/wiki/Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></li> <li><a href="/wiki/Globuloviridae" title="Globuloviridae">Globuloviridae</a></li> <li><a href="/wiki/Guttaviridae" title="Guttaviridae">Guttaviridae</a></li> <li><a href="/wiki/Halspiviridae" title="Halspiviridae">Halspiviridae</a></li> <li><a href="/wiki/Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></li> <li><a href="/wiki/Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></li> <li><a href="/wiki/Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></li> <li><a href="/wiki/Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></li> <li><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li> <li><a href="/wiki/Spiraviridae" title="Spiraviridae">Spiraviridae</a></li> <li><a href="/wiki/Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></li> <li><a href="/wiki/Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genera</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/Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></li> <li><a href="/wiki/Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nanobe" title="Nanobe">Nanobe</a> (?)</li> <li><a href="/wiki/Obelisk_(biology)" title="Obelisk (biology)">Obelisk</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Comparable structures</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bio-like_structure" title="Bio-like structure">Bio-like structure</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Integrative_and_conjugative_element" title="Integrative and conjugative element">Integrative and conjugative element</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li> <li>"<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a>"</li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Prion" title="Prion">Prion</a> <ul><li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a> <a href="/wiki/Microparticle#Biological_protocells" title="Microparticle">microsphere</a></li> <li>Retroelements not elsewhere classified <ul><li><a href="/wiki/Retron" title="Retron">Retron</a></li> <li><a href="/wiki/Diversity-generating_retroelement" title="Diversity-generating retroelement">Diversity-generating retroelement</a></li> <li><a href="/wiki/Telomerase_reverse_transcriptase" title="Telomerase reverse transcriptase">Telomerase reverse transcriptase</a></li> <li><a href="/wiki/Reverse_transcriptase-related_cellular_gene" class="mw-redirect" title="Reverse transcriptase-related cellular gene">Reverse transcriptase-related cellular gene</a></li></ul></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Spiegelman%27s_Monster" title="Spiegelman's Monster">Spiegelman's Monster</a></li> <li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retroposon" title="Retroposon">Retroposon</a></li></ul></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li> <li><a href="/wiki/Xenobot" title="Xenobot">Xenobot</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235" /></div><div role="navigation" class="navbox" aria-labelledby="Taxon_identifiers852" 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_identifiers852" 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href="https://gd.eppo.int/taxon/1VIRDD">1VIRDD</a></li> <li><a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=12884">12884</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235" /><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319" /></div><div role="navigation" class="navbox authority-control" aria-label="Navbox1095" style="padding:3px"><table class="nowraplinks hlist navbox-inner" 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