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class="vector-toc-numb">2</span> <span>Genome organization</span> </div> </a> <button aria-controls="toc-Genome_organization-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Genome organization subsection</span> </button> <ul id="toc-Genome_organization-sublist" class="vector-toc-list"> <li id="toc-Proteins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Proteins</span> </div> </a> <ul id="toc-Proteins-sublist" class="vector-toc-list"> <li id="toc-Phosphoprotein" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Phosphoprotein"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.1</span> <span>Phosphoprotein</span> </div> </a> <ul id="toc-Phosphoprotein-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Large_structural_protein_(polymerase)" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Large_structural_protein_(polymerase)"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.2</span> <span>Large structural protein (polymerase)</span> </div> </a> <ul id="toc-Large_structural_protein_(polymerase)-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Life_cycle" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Life_cycle"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Life cycle</span> </div> </a> <ul id="toc-Life_cycle-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Infection" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Infection"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Infection</span> </div> </a> <ul id="toc-Infection-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Signs_and_symptoms" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Signs_and_symptoms"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Signs and symptoms</span> </div> </a> <ul id="toc-Signs_and_symptoms-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Classification_and_antigenic_types" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Classification_and_antigenic_types"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Classification and antigenic types</span> </div> </a> <ul id="toc-Classification_and_antigenic_types-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Antigenicity" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Antigenicity"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Antigenicity</span> </div> </a> <ul id="toc-Antigenicity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Application" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Application"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Application</span> </div> </a> <ul id="toc-Application-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> 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class="firstHeading mw-first-heading"><span class="mw-page-title-main">Rabies virus</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 27 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-27" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">27 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%81%D9%8A%D8%B1%D9%88%D8%B3_%D8%AF%D8%A7%D8%A1_%D8%A7%D9%84%D9%83%D9%84%D8%A8" title="فيروس داء الكلب – Arabic" lang="ar" hreflang="ar" data-title="فيروس داء الكلب" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%B0%E0%A7%87%E0%A6%AC%E0%A6%BF%E0%A6%9C_%E0%A6%AD%E0%A6%BE%E0%A6%87%E0%A6%B0%E0%A6%BE%E0%A6%B8" title="রেবিজ ভাইরাস – Bangla" lang="bn" hreflang="bn" data-title="রেবিজ ভাইরাস" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/Rabies_virus" title="Rabies virus – Bulgarian" lang="bg" hreflang="bg" data-title="Rabies virus" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Virus_de_la_r%C3%A0bia" title="Virus de la ràbia – Catalan" lang="ca" hreflang="ca" data-title="Virus de la ràbia" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Virus_vztekliny" title="Virus vztekliny – Czech" lang="cs" hreflang="cs" data-title="Virus vztekliny" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Rabiesvirus" title="Rabiesvirus – German" lang="de" hreflang="de" data-title="Rabiesvirus" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Virus_de_la_rabia" title="Virus de la rabia – Spanish" lang="es" hreflang="es" data-title="Virus de la rabia" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Virus_de_la_rage" title="Virus de la rage – French" lang="fr" hreflang="fr" data-title="Virus de la rage" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Virus_da_rabia" title="Virus da rabia – Galician" lang="gl" hreflang="gl" data-title="Virus da rabia" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EA%B4%91%EA%B2%AC%EB%B3%91%EB%B0%94%EC%9D%B4%EB%9F%AC%EC%8A%A4" title="광견병바이러스 – Korean" lang="ko" hreflang="ko" data-title="광견병바이러스" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Virus_rabies" title="Virus rabies – Indonesian" lang="id" hreflang="id" data-title="Virus rabies" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Virus_della_rabbia" title="Virus della rabbia – Italian" lang="it" hreflang="it" data-title="Virus della rabbia" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-lmo mw-list-item"><a href="https://lmo.wikipedia.org/wiki/Virus_de_la_rabia" title="Virus de la rabia – Lombard" lang="lmo" hreflang="lmo" data-title="Virus de la rabia" data-language-autonym="Lombard" data-language-local-name="Lombard" class="interlanguage-link-target"><span>Lombard</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Veszetts%C3%A9gv%C3%ADrus" title="Veszettségvírus – Hungarian" lang="hu" hreflang="hu" data-title="Veszettségvírus" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D9%81%D9%8A%D8%B1%D9%88%D8%B3_%D8%AF%D8%A7%D8%A1_%D8%A7%D9%84%D9%83%D9%84%D8%A8" title="فيروس داء الكلب – Egyptian Arabic" lang="arz" hreflang="arz" data-title="فيروس داء الكلب" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target"><span>مصرى</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Virus_penyakit_anjing_gila" title="Virus penyakit anjing gila – Malay" lang="ms" hreflang="ms" data-title="Virus penyakit anjing gila" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%8B%82%E7%8A%AC%E7%97%85%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9" title="狂犬病ウイルス – Japanese" lang="ja" hreflang="ja" data-title="狂犬病ウイルス" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Rabiesvirus" title="Rabiesvirus – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Rabiesvirus" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Wirus_w%C5%9Bcieklizny" title="Wirus wścieklizny – Polish" lang="pl" hreflang="pl" data-title="Wirus wścieklizny" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/V%C3%ADrus_da_raiva" title="Vírus da raiva – Portuguese" lang="pt" hreflang="pt" data-title="Vírus da raiva" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81_%D0%B1%D0%B5%D1%88%D0%B5%D0%BD%D1%81%D1%82%D0%B2%D0%B0" title="Вирус бешенства – Russian" lang="ru" hreflang="ru" data-title="Вирус бешенства" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Rabies_virus" title="Rabies virus – Simple English" lang="en-simple" hreflang="en-simple" data-title="Rabies virus" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Birus_ng_rabies" title="Birus ng rabies – Tagalog" lang="tl" hreflang="tl" data-title="Birus ng rabies" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></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%B1%E0%B8%AA%E0%B9%82%E0%B8%A3%E0%B8%84%E0%B8%9E%E0%B8%B4%E0%B8%A9%E0%B8%AA%E0%B8%B8%E0%B8%99%E0%B8%B1%E0%B8%82%E0%B8%9A%E0%B9%89%E0%B8%B2" title="ไวรัสโรคพิษสุนัขบ้า – Thai" lang="th" hreflang="th" data-title="ไวรัสโรคพิษสุนัขบ้า" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%92%D1%96%D1%80%D1%83%D1%81_%D1%81%D0%BA%D0%B0%D0%B7%D1%83" title="Вірус сказу – Ukrainian" lang="uk" hreflang="uk" data-title="Вірус сказу" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Virus_d%E1%BA%A1i" title="Virus dại – Vietnamese" lang="vi" hreflang="vi" data-title="Virus dại" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%8B%82%E7%8A%AC%E7%97%85%E7%97%85%E6%AF%92" title="狂犬病病毒 – Chinese" lang="zh" hreflang="zh" data-title="狂犬病病毒" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q698976#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet 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lyssavirus</a>)</span></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Species of virus</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">This article is about the virus. For the disease, see <a href="/wiki/Rabies" title="Rabies">Rabies</a>. 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width: 200px; font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(250,250,190)"><i>Rabies lyssavirus</i> </th></tr> <tr> <td colspan="2" style="text-align: center"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Rabies_Virus_(51706598561).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Rabies_Virus_%2851706598561%29.jpg/220px-Rabies_Virus_%2851706598561%29.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Rabies_Virus_%2851706598561%29.jpg/330px-Rabies_Virus_%2851706598561%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Rabies_Virus_%2851706598561%29.jpg/440px-Rabies_Virus_%2851706598561%29.jpg 2x" data-file-width="3510" data-file-height="3510" /></a></span> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%">Colorized transmission electron micrograph showing the rabies virus (in red) infecting cultured cells </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> <span class="plainlinks taxobox-edit-taxonomy skin-invert" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><span typeof="mw:File"><a href="/wiki/Template:Taxonomy/Lyssavirus" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></span> </th></tr> <tr> <td>(unranked): </td> <td><a href="/wiki/Virus" title="Virus">Virus</a> </td></tr> <tr> <td><i>Realm</i>: </td> <td><a href="/wiki/Riboviria" title="Riboviria"><i>Riboviria</i></a> </td></tr> <tr> <td>Kingdom: </td> <td><a href="/wiki/Orthornavirae" title="Orthornavirae"><i>Orthornavirae</i></a> </td></tr> <tr> <td>Phylum: </td> <td><a href="/wiki/Negarnaviricota" class="mw-redirect" title="Negarnaviricota"><i>Negarnaviricota</i></a> </td></tr> <tr> <td>Class: </td> <td><a href="/wiki/Monjiviricetes" title="Monjiviricetes"><i>Monjiviricetes</i></a> </td></tr> <tr> <td>Order: </td> <td><a href="/wiki/Mononegavirales" title="Mononegavirales"><i>Mononegavirales</i></a> </td></tr> <tr> <td>Family: </td> <td><a href="/wiki/Rhabdoviridae" title="Rhabdoviridae"><i>Rhabdoviridae</i></a> </td></tr> <tr> <td>Genus: </td> <td><a href="/wiki/Lyssavirus" title="Lyssavirus"><i>Lyssavirus</i></a> </td></tr> <tr> <td>Species: </td> <td><div style="display:inline" class="species"><i><b>Rabies lyssavirus</b></i></div> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(250,250,190)">Member viruses </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul><li>Rabies virus</li> <li><a href="/wiki/Arctic_rabies_virus" title="Arctic rabies virus">Arctic rabies virus</a></li></ul> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(250,250,190)"><a href="/wiki/Synonym_(taxonomy)" title="Synonym (taxonomy)">Synonyms</a><sup id="cite_ref-2015.006aM_1-0" class="reference"><a href="#cite_note-2015.006aM-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul><li><i>Rabies virus</i></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> <p><b>Rabies virus</b>, scientific name <i><b>Rabies lyssavirus</b></i>, is a <a href="/wiki/Neurotropic_virus" title="Neurotropic virus">neurotropic virus</a> that causes <a href="/wiki/Rabies" title="Rabies">rabies</a> in animals, including humans. It can cause violence, hydrophobia, and fever. <a href="/wiki/Rabies_transmission" class="mw-redirect" title="Rabies transmission">Rabies transmission</a> can also occur through the <a href="/wiki/Saliva" title="Saliva">saliva</a> of animals and less commonly through contact with human saliva. <i>Rabies lyssavirus</i>, like many <a href="/wiki/Rhabdoviridae" title="Rhabdoviridae">rhabdoviruses</a>, has an extremely wide host range. In the wild it has been found infecting many mammalian species, while in the laboratory it has been found that birds can be infected, as well as cell cultures from mammals, birds, reptiles and insects.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Rabies is reported in more than 150 countries and on all continents except Antarctica.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The main burden of disease is reported in Asia and Africa, but some cases have been reported also in Europe in the past 10 years, especially in returning travellers.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p><i>Rabies lyssavirus</i> has a cylindrical morphology and is a member of the <i><a href="/wiki/Lyssavirus" title="Lyssavirus">Lyssavirus</a></i> <a href="/wiki/Genus" title="Genus">genus</a> of the <i><a href="/wiki/Rhabdoviridae" title="Rhabdoviridae">Rhabdoviridae</a></i> family. These viruses are <a href="/wiki/Viral_envelope" title="Viral envelope">enveloped</a> and have a <a href="/wiki/Negative-sense_single-stranded_RNA_virus" class="mw-redirect" title="Negative-sense single-stranded RNA virus">single stranded RNA genome with negative-sense</a>. The genetic information is packaged as a <a href="/wiki/Ribonucleoprotein" class="mw-redirect" title="Ribonucleoprotein">ribonucleoprotein</a> complex in which <a href="/wiki/RNA" title="RNA">RNA</a> is tightly bound by the viral nucleoprotein. The RNA genome of the virus encodes five genes whose order is highly conserved. These genes code for nucleoprotein (N), phosphoprotein (P), matrix protein (M), glycoprotein (G) and the viral RNA polymerase (L).<sup id="cite_ref-pmid15885837_5-0" class="reference"><a href="#cite_note-pmid15885837-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The complete genome sequences range from 11,615 to 11,966 nt in length.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>All transcription and replication events take place in the cytoplasm inside a specialized "virus factory", the <i><a href="/wiki/Negri_bodies" class="mw-redirect" title="Negri bodies">Negri body</a></i> (named after <a href="/wiki/Adelchi_Negri" title="Adelchi Negri">Adelchi Negri</a><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>). These are 2–10 <a href="/wiki/Micrometre" title="Micrometre">μm</a> in diameter and are typical for a rabies infection and thus have been used as <a href="/wiki/Pathognomonic" title="Pathognomonic">definite histological proof of such infection</a>.<sup id="cite_ref-pmid17938861_8-0" class="reference"><a href="#cite_note-pmid17938861-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Rabies_virus_structure.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8c/Rabies_virus_structure.jpg/220px-Rabies_virus_structure.jpg" decoding="async" width="220" height="124" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8c/Rabies_virus_structure.jpg/330px-Rabies_virus_structure.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8c/Rabies_virus_structure.jpg/440px-Rabies_virus_structure.jpg 2x" data-file-width="1920" data-file-height="1080" /></a><figcaption>3D still showing rabies virus structure.</figcaption></figure> <p>Rhabdoviruses have <a href="/wiki/Helix" title="Helix">helical</a> symmetry, so their infectious particles are approximately cylindrical in shape. They are characterized by an extremely broad host spectrum ranging from plants <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (July 2022)">citation needed</span></a></i>]</sup> to insects <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (July 2022)">citation needed</span></a></i>]</sup> and mammals; human-infecting viruses more commonly have icosahedral symmetry and take shapes approximating <a href="/wiki/Regular_polyhedron" title="Regular polyhedron">regular polyhedra</a>. </p><p>The rabies genome encodes five proteins: nucleoprotein (N), phosphoprotein (P), matrix protein (M), glycoprotein (G) and polymerase (L). All rhabdoviruses have two major structural components: a helical ribonucleoprotein core (RNP) and a surrounding envelope. In the RNP, genomic RNA is tightly encased by the nucleoprotein. Two other viral proteins, the phosphoprotein and the large protein (L-protein or polymerase) are associated with the RNP. The glycoprotein forms approximately 400 trimeric spikes which are tightly arranged on the surface of the virus. The M protein is associated both with the envelope and the RNP and may be the central protein of rhabdovirus assembly.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p><i>Rabies lyssavirus</i> has a bullet-like shape with a length of about 180 <a href="/wiki/Nanometer" class="mw-redirect" title="Nanometer">nm</a> and a cross-sectional diameter of about 75 nm. One end is rounded or conical and the other end is planar or concave. The <a href="/wiki/Lipoprotein" title="Lipoprotein">lipoprotein</a> envelope carries knob-like spikes composed of <a href="/wiki/Glycoprotein" title="Glycoprotein">Glycoprotein</a> G. Spikes do not cover the planar end of the virion (virus particle). Beneath the envelope is the membrane or matrix (M) protein layer which may be <a href="/wiki/Invagination" title="Invagination">invaginated</a> at the planar end. The core of the virion consists of helically arranged <a href="/wiki/Ribonucleoprotein" class="mw-redirect" title="Ribonucleoprotein">ribonucleoprotein</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (April 2023)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genome_organization">Genome organization</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=2" title="Edit section: Genome organization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The rhabdovirus virion is an enveloped, rod- or bullet-shaped structure containing five protein species: </p> <ul><li>The nucleoprotein (N) coats the RNA at the rate of one monomer of protein to nine nucleotides, forming a nucleocapsid with helical symmetry. Associated with the nucleocapsid are copies of P (phosphoprotein) and L (large) protein.</li> <li>The L protein is well named, its gene taking up about half of the genome. Its large size is justified by the fact that it is a multifunctional protein.</li> <li>The M (matrix) protein forms a layer between the nucleocapsid and the envelope</li> <li>Trimers of G (glycoprotein) form spikes that protrude from the envelope.</li></ul> <p>The genomes of most rhabdoviruses possess the five genes encoding these proteins,<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 5">: 5 </span></sup> as in the case of <i>Rhabdovirus lyssae</i>.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Proteins">Proteins</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=3" title="Edit section: Proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table class="wikitable sortable"> <caption>List of Rabies virus ORFs (3′ to 5′) </caption> <tbody><tr> <th>Symbol</th> <th>Name</th> <th>UniProt</th> <th>Function </th></tr> <tr> <td>N</td> <td>Nucleoprotein</td> <td><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P16285">P16285</a></span></td> <td>Coats the RNA. </td></tr> <tr> <td>P</td> <td>Phosphoprotein</td> <td><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P16286">P16286</a></span></td> <td>L cofactor and various regulatory functions. Has many isoforms from multiple initiation. </td></tr> <tr> <td>M</td> <td>Matrix</td> <td><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P16287">P16287</a></span></td> <td>Keeps nucleoprotein condensed. Important for assembly; has roles in regulation. </td></tr> <tr> <td>G</td> <td>Glycoprotein</td> <td><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P16288">P16288</a></span></td> <td>Spike. Uses muscular <a href="/wiki/NAChR" class="mw-redirect" title="NAChR">nAChR</a>, <a href="/wiki/NCAM" class="mw-redirect" title="NCAM">NCAM</a>, and <a href="/wiki/P75NTR" class="mw-redirect" title="P75NTR">p75NTR</a> as receptors. </td></tr> <tr> <td>L</td> <td>Large structural protein</td> <td><span class="plainlinks"><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprot/P16289">P16289</a></span></td> <td><a href="/wiki/RNA_replicase" class="mw-redirect" title="RNA replicase">RNA replicase</a> of the <i>Mononegavirales</i> type. </td></tr></tbody></table> <p>Rabies virus is estimated to cause around 55,000 deaths per year across the world<sup id="cite_ref-:0_13-0" class="reference"><a href="#cite_note-:0-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> and has a death rate of nearly 100%.<sup id="cite_ref-:1_14-0" class="reference"><a href="#cite_note-:1-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> These statistics coupled with the fact that there is currently no specific treatment, or antiviral drug<sup id="cite_ref-:0_13-1" class="reference"><a href="#cite_note-:0-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> makes research on the virus of vital importance for the scientific community in order to possibly lower the current death rate. The rabies virus phosphoprotein and polymerase are both important targets for <a href="/wiki/Antiviral" class="mw-redirect" title="Antiviral">antivirals</a> and are currently used to create the vaccine used for domestic and wild animals. A lot of research is being done to better understand the specific roles and functions of the L-P protein because there is significant evidence already that it could be one of the most important proteins to target for future drugs.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (April 2023)">citation needed</span></a></i>]</sup> </p><p>There are five proteins that are coded for by the rabies virus genome—phosphoprotein (P), polymerase (L), matrix protein (M), nucleoprotein (N), and glycoprotein (G). These five proteins are transcribed into mRNA in different quantities. The protein transcribed the most is the nucleoprotein, then the phosphoprotein, then the matrix protein, then the glycoprotein and finally the polymerase.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Of those proteins, the ones that may be the most important for the functions of the virus are the L-P protein complex. These two proteins are required for the production of all of the proteins utilized by the rabies virus and they interact with many of the other proteins to complete the functions needed by the virus to infect cells, replicate and complete other vital functions.<sup id="cite_ref-:1_14-1" class="reference"><a href="#cite_note-:1-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> When the structure of the L-P protein was analyzed using UCSF Chimera, it was found that it contained two zinc molecules as well as 2 five-membered rings. The secondary structures were also analyzed, and it was found that there were three different kinds-coil, helix and strand. </p> <div class="mw-heading mw-heading4"><h4 id="Phosphoprotein">Phosphoprotein</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=4" title="Edit section: Phosphoprotein"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The phosphoprotein (P) consists of 297 amino acids,<sup id="cite_ref-:2_16-0" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> has a total mass of 33 k-Da<sup id="cite_ref-:0_13-2" class="reference"><a href="#cite_note-:0-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> and is a type I interferon antagonist. It is in a viral ribonucleoprotein made up of RNA surrounded by nucleoproteins (N),<sup id="cite_ref-:1_14-2" class="reference"><a href="#cite_note-:1-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> which form a structure that is ring-like.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Each nucleoprotein surrounds nine nucleotides of the RNA<sup id="cite_ref-:0_13-3" class="reference"><a href="#cite_note-:0-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> and the nucleoproteins in conjunction with the phosphoprotein are important for initiating synthesis of the RNA. The phosphoprotein is largely responsible for preventing the immune system from destroying the virus before it can take hold. It has been found that it stops production of interferons by disrupting the interferon regulatory factor-3 (IRF-3) and, therefore, prevents the normal function of the interferons, which allows it to continue to infect the host.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The phosphoprotein is also very important for replication. It is a required cofactor of the viral polymerase (L) and is also a chaperone for the nucleoprotein (N), which is responsible for the encapsidation of viral RNA.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> It is thought that the P protein consists of only a few repeating units. </p> <div class="mw-heading mw-heading4"><h4 id="Large_structural_protein_(polymerase)"><span id="Large_structural_protein_.28polymerase.29"></span>Large structural protein (polymerase)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=5" title="Edit section: Large structural protein (polymerase)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The polymerase (L) consists of 2,127 amino acids,<sup id="cite_ref-:2_16-1" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> has a total mass of 242 k-Da<sup id="cite_ref-:0_13-4" class="reference"><a href="#cite_note-:0-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> and interacts with the phosphoprotein to form the RNA-dependent RNA polymerase complex. L and P bind to each other and according to current research, the major L binding site is somewhere in the first 19 residues of the P protein.<sup id="cite_ref-:2_16-2" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Other research found that the amino acid sequence numbered 1929 to 1933 is the sequence that is important for the L protein to bind with the P protein. It has been found that the L protein initiates synthesis and works optimally if it works together with the P protein.<sup id="cite_ref-:0_13-5" class="reference"><a href="#cite_note-:0-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The region from 1900 to 1930, which contains part of the sequence that is important for L and P protein binding, and the region from 1870 to 1890 are hydrophilic regions and are also highly conserved among rabies virus strains.<sup id="cite_ref-:1_14-3" class="reference"><a href="#cite_note-:1-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The 1929 to 1933 sequence is also of importance because it is involved with RNA synthesis of the virus and thus could possibly be a target for new drugs. The carboxy-terminal domain of L must be present in order for L to bind to P. The L protein also has a catalytic center while the P protein is believed to not have a catalytic center.<sup id="cite_ref-:2_16-3" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> There is evidence that for the rabies virus, the L protein can be expressed even if the P protein is not expressed. This is different from many other related viruses, which require both to be expressed and present.<sup id="cite_ref-:2_16-4" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> One of the domains in the L protein is responsible for speeding up the reaction that results in the capping of the RNA and thus could possibly be another good target for an antiviral drug.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> The nucleoproteins (N), which it was previously mentioned aid the P protein, also interact with the L protein and this is necessary for initiation.<sup id="cite_ref-:0_13-6" class="reference"><a href="#cite_note-:0-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The interactions between the L protein and the M protein have also been noted to be of importance to its function. Research has shown that there are interactions between the two proteins in relation to the balance of transcription and translation.<sup id="cite_ref-:1_14-4" class="reference"><a href="#cite_note-:1-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The polymerase is an enzyme and can aid in initiation, elongation, capping, methylation, polyadenylation<sup id="cite_ref-:2_16-5" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> and RNA polymerization.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Life_cycle">Life cycle</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=6" title="Edit section: Life cycle"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>After receptor binding, <i>Rabies lyssavirus</i> enters its host cells through the <a href="/wiki/Endosome" title="Endosome">endosomal</a> transport pathway. Inside the endosome, the low <a href="/wiki/PH" title="PH">pH</a> value induces the membrane fusion process, thus enabling the viral genome to reach the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>. Both processes, receptor binding and membrane fusion, are catalyzed by the glycoprotein G which plays a critical role in pathogenesis (mutant virus without G proteins cannot propagate).<sup id="cite_ref-pmid15885837_5-1" class="reference"><a href="#cite_note-pmid15885837-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>The next step after its entry is the <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> of the viral genome by the P-L polymerase (P is an essential cofactor for the L polymerase) in order to make new viral protein. The viral polymerase can only recognize <a href="/wiki/Ribonucleoprotein" class="mw-redirect" title="Ribonucleoprotein">ribonucleoprotein</a> and cannot use free RNA as template. Transcription is regulated by <a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element"><i>cis</i>-acting sequences</a> on the virus genome and by protein M which is not only essential for virus budding but also regulates the fraction of mRNA production to replication. Later in infection, the activity of the polymerase switches to replication in order to produce full-length positive-strand RNA copies. These complementary RNAs are used as templates to make new negative-strand RNA genomes. They are packaged together with protein N to form <a href="/wiki/Ribonucleoprotein" class="mw-redirect" title="Ribonucleoprotein">ribonucleoprotein</a> which then can form new viruses.<sup id="cite_ref-pmid17938861_8-1" class="reference"><a href="#cite_note-pmid17938861-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Infection">Infection</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=7" title="Edit section: Infection"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Rabies_Virus_EM_PHIL_1876.JPG" class="mw-file-description"><img alt="TEM micrograph with numerous rabies virions (small dark-grey rod-like particles) and Negri bodies (the larger pathognomonic cellular inclusions of rabies infection)" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/Rabies_Virus_EM_PHIL_1876.JPG/220px-Rabies_Virus_EM_PHIL_1876.JPG" decoding="async" width="220" height="287" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/Rabies_Virus_EM_PHIL_1876.JPG/330px-Rabies_Virus_EM_PHIL_1876.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/57/Rabies_Virus_EM_PHIL_1876.JPG/440px-Rabies_Virus_EM_PHIL_1876.JPG 2x" data-file-width="1835" data-file-height="2392" /></a><figcaption><a href="/wiki/Transmission_electron_microscopy" title="Transmission electron microscopy">TEM</a> <a href="/wiki/Micrograph" title="Micrograph">micrograph</a> with numerous rabies <a href="/wiki/Virion" class="mw-redirect" title="Virion">virions</a> (small dark-grey rod-like particles) and <a href="/wiki/Negri_bodies" class="mw-redirect" title="Negri bodies">Negri bodies</a> (the larger <a href="/wiki/Pathognomonic" title="Pathognomonic">pathognomonic</a> cellular inclusions of rabies infection)</figcaption></figure> <p>In September 1931, <a href="/wiki/Joseph_Lennox_Pawan" title="Joseph Lennox Pawan">Joseph Lennox Pawan</a> of <a href="/wiki/Trinidad" title="Trinidad">Trinidad</a> found <a href="/wiki/Negri_bodies" class="mw-redirect" title="Negri bodies">Negri bodies</a> in the brain of a bat with unusual habits. In 1932, Pawan first discovered that infected <a href="/wiki/Vampire_bat" title="Vampire bat">vampire bats</a> could transmit rabies to humans and other animals.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Rabies virus initially replicates in muscle tissue following a bite before entering neurons through their nerve endings and spreading to the nervous system.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Axoplasmic_transport" class="mw-redirect" title="Axoplasmic transport">retrograde axonal transport</a> of <i>Rabies lyssavirus</i> to the <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a> (CNS) is the key step of pathogenesis during natural infection. The exact molecular mechanism of this transport is unknown although binding of the P protein from <i>Rabies lyssavirus</i> to the <a href="/wiki/Dynein" title="Dynein">dynein</a> light chain protein <a href="/wiki/DYNLL1" title="DYNLL1">DYNLL1</a> has been shown.<sup id="cite_ref-pmid11024151_28-0" class="reference"><a href="#cite_note-pmid11024151-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> P also acts as an <a href="/wiki/Interferon" title="Interferon">interferon</a> antagonist, thus decreasing the <a href="/wiki/Immune" class="mw-redirect" title="Immune">immune</a> response of the host.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (April 2023)">citation needed</span></a></i>]</sup> </p><p>From the CNS, the virus further spreads to other organs. The salivary glands located in the tissues of the mouth and cheeks receive high concentrations of the virus, thus allowing it to be further transmitted due to projectile salivation. Fatality can occur from two days to five years from the time of initial infection.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> This however depends largely on the species of animal acting as a <a href="/wiki/Natural_reservoir" title="Natural reservoir">reservoir</a>. Most infected mammals die within weeks, while strains of a species such as the African <a href="/wiki/Yellow_mongoose" title="Yellow mongoose">yellow mongoose</a> (<i>Cynictis penicillata</i>) might survive an infection asymptomatically for years.<sup id="cite_ref-pmid7777324_30-0" class="reference"><a href="#cite_note-pmid7777324-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Signs_and_symptoms">Signs and symptoms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=8" title="Edit section: Signs and symptoms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first symptoms of rabies may be very similar to those of the flu, including general weakness or discomfort, fever, or headache. These symptoms may last for days. There may be also discomfort or a prickling or itching sensation at the site of bite, progressing within days to symptoms of cerebral dysfunction, anxiety, confusion, and agitation. At the onset of these progressive symptoms, the disease begins to worsen. As the disease progresses, the person may experience delirium, abnormal behavior, hallucinations, and insomnia. <i>Rabies lyssavirus</i> may also be inactive in its host's body and become active after a long period of time.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>The incubation period for this virus, in some cases, can last anywhere from weeks to months. This does not depend on specific form of the virus. This long incubation period is most likely due to the presence of <a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a>, which slow down viral replication in the muscles. However, when the patient starts to exhibit prodromal symptoms (fever, flu) and gastrointestinal symptoms, the virus has widely spread.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Prodromal symptoms are noted within the first 2–10 days after incubation and include <a href="/wiki/Fever" title="Fever">fever</a> and fatigue. They can also involve the <a href="/wiki/Respiratory_system" title="Respiratory system">respiratory system</a>, causing sore throat and cough, the gastrointestinal system, causing anorexia, nausea, vomiting, abdominal pain, diarrhea, and central nervous system. The central nervous system is the most prevalent in some cases and includes symptoms like headache, vertigo, anxiety, nightmares, depression, and more. The neurological dysfunction starts when the central nervous system begins to slow and not function properly.<sup id="cite_ref-Rupprecht_1996_33-0" class="reference"><a href="#cite_note-Rupprecht_1996-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Classification_and_antigenic_types">Classification and antigenic types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=9" title="Edit section: Classification and antigenic types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <i><a href="/wiki/Lyssavirus" title="Lyssavirus">Lyssavirus</a></i> genome includes a variety of viruses ranging from the rabies virus to genetically and anti-genetically rabies-like related viruses. These viruses include <a href="/wiki/Lagos_bat_lyssavirus" title="Lagos bat lyssavirus">Lagos Bat</a>, <a href="/wiki/Mokola_lyssavirus" title="Mokola lyssavirus">Mokola</a>, and <a href="/wiki/Duvenhage_lyssavirus" title="Duvenhage lyssavirus">Duvenhage</a> viruses, as well as European bat viruses.<sup id="cite_ref-Rupprecht_1996_33-1" class="reference"><a href="#cite_note-Rupprecht_1996-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Through other anti-genetic characterization, over 350 rabies viruses of just the Americas were discovered. Four phylogenetic groups were associated with these viruses. Additionally, other areas of ciruculation for the virus, such as <i><a href="/wiki/Big_brown_bat" title="Big brown bat">E. fuscus</a></i>, were found through this discovery.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Although these viruses are from the same genome, cross-protection studies show that animals that had taken the <a href="/wiki/Lyssavirus" title="Lyssavirus"><i>Lyssavirus</i> vaccine</a> for traditional rabies were not fully protected when exposed to other types of <i><a href="/wiki/Lyssavirus" title="Lyssavirus">Lyssavirus</a></i>. </p><p>This leads to different categorization of these Rabies viruses. A fixed classification denotes that the virus was adapted by passage in animals or cell culture. Wild type classification, more generally known as street type, implies the virus was adapted through other means, such as a bite. To differentiate street rabies variants, <a href="/wiki/Monoclonal_antibody" title="Monoclonal antibody">monoclonal antibodies</a> identified origins in host reservoirs throughout the world.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> This suggested sources of exposure even when the bite was missing from patient history. </p> <div class="mw-heading mw-heading2"><h2 id="Antigenicity">Antigenicity</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=10" title="Edit section: Antigenicity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Upon viral entry into the body and also after <a href="/wiki/Rabies_vaccine" title="Rabies vaccine">vaccination</a>, the body produces virus neutralizing antibodies which bind and inactivate the virus. Specific regions of the G protein have been shown to be most antigenic in leading to the production of virus neutralizing antibodies. These antigenic sites, or epitopes, are categorized into regions I–IV and minor site a. Previous work has demonstrated that antigenic sites II and III are most commonly targeted by natural neutralizing antibodies.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Additionally, a monoclonal antibody with neutralizing functionality has been demonstrated to target antigenic site I.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Other proteins, such as the nucleoprotein, have been shown to be unable to elicit production of virus neutralizing antibodies.<sup id="cite_ref-Wiktor-1973_38-0" class="reference"><a href="#cite_note-Wiktor-1973-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> The epitopes which bind neutralizing antibodies are both linear and conformational.<sup id="cite_ref-Bakker-2005_39-0" class="reference"><a href="#cite_note-Bakker-2005-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=11" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>All extant rabies viruses appear to have evolved within the last 1500 years.<sup id="cite_ref-Nadin-Davis2011_40-0" class="reference"><a href="#cite_note-Nadin-Davis2011-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> There are seven genotypes of <i>Rabies lyssavirus</i>. In Eurasia, cases are due to three of these: genotype 1 (classical rabies) and to a lesser extent genotypes 5 and 6 (European bat lyssaviruses type-1 and -2).<sup id="cite_ref-McElhinney2008_41-0" class="reference"><a href="#cite_note-McElhinney2008-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Genotype 1 evolved in Europe in the 17th century and spread to Asia, Africa, and the Americas as a result of European exploration and colonization.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (April 2023)">citation needed</span></a></i>]</sup> </p><p>Bat rabies in North America appears to have been present since 1281 AD (95% <a href="/wiki/Confidence_interval" title="Confidence interval">confidence interval</a>: 906–1577 AD).<sup id="cite_ref-Kuzmina2013_42-0" class="reference"><a href="#cite_note-Kuzmina2013-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>The rabies virus appears to have undergone an evolutionary shift in hosts from Chiroptera (<a href="/wiki/Bat" title="Bat">bats</a>) to a species of <a href="/wiki/Carnivora" title="Carnivora">Carnivora</a> (i.e. <a href="/wiki/Raccoon" title="Raccoon">raccoon</a> or <a href="/wiki/Skunk" title="Skunk">skunk</a>) as a result of an <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombination</a> event that occurred hundreds of years ago.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> This recombination event altered the gene that encodes the virus <a href="/wiki/Glycoprotein" title="Glycoprotein">glycoprotein</a> that is necessary for receptor recognition and binding. </p> <div class="mw-heading mw-heading2"><h2 id="Application">Application</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=12" title="Edit section: Application"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i>Rabies lyssavirus</i> is used in research for <a href="/wiki/Viral_neuronal_tracing" title="Viral neuronal tracing">viral neuronal tracing</a> to establish synaptic connections and directionality of synaptic transmission.<sup id="cite_ref-Ginger_44-0" class="reference"><a href="#cite_note-Ginger-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>Interestingly, the rabies virus vaccine that was created using the SAD-B19 complex, which includes the L-P protein, was utilized in the creation of a vaccine for SARS-CoV-2. The S1 protein from SARS-CoV-2 was inserted into the rabies virus vaccine vector to create a new vaccine that was shown to be effective in protecting against COVID-19. Rabies virus has also been used to create a vaccine against Ebola, called FiloRab1, and it was found to be 100% effective for nonhuman primates.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=13" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1259569809">.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 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srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/42px-Sida-aids.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/56px-Sida-aids.png 2x" data-file-width="290" data-file-height="290" /></a></span></span><span class="portalbox-link"><a href="/wiki/Portal:Viruses" title="Portal:Viruses">Viruses portal</a></span></li></ul> <ul><li><a href="/wiki/Cryptic_bat_rabies" class="mw-redirect" title="Cryptic bat rabies">Cryptic bat rabies</a></li> <li><a href="/wiki/Rabies_vaccine" title="Rabies vaccine">Rabies vaccine</a></li> <li><a href="/wiki/Duck_embryo_vaccine" title="Duck embryo vaccine">Duck embryo vaccine</a></li> <li><a href="/wiki/Arctic_rabies_virus" title="Arctic rabies virus">Arctic rabies virus</a></li> <li><a href="/wiki/Bat-borne_virus" class="mw-redirect" title="Bat-borne virus">Bat-borne virus</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Rabies_virus&action=edit&section=14" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-2015.006aM-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-2015.006aM_1-0">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFWalker2015" class="citation web cs1">Walker, Peter (15 June 2015). <a rel="nofollow" class="external text" href="https://ictv.global/ictv/proposals/2015.006aM.A.v3.Rhabdoviridae_spren.pdf">"mplementation of taxon-wide non-Latinized binomial species names in the family <i>Rhabdoviridae</i>"</a> <span class="cs1-format">(PDF)</span>. <i>International Committee on Taxonomy of Viruses (ICTV)</i><span class="reference-accessdate">. Retrieved <span class="nowrap">11 February</span> 2019</span>. <q><i>Rabies virus</i> <i>Rabies lyssavirus</i> rabies virus (RABV)[M13215]</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=International+Committee+on+Taxonomy+of+Viruses+%28ICTV%29&rft.atitle=mplementation+of+taxon-wide+non-Latinized+binomial+species+names+in+the+family+Rhabdoviridae&rft.date=2015-06-15&rft.aulast=Walker&rft.aufirst=Peter&rft_id=https%3A%2F%2Fictv.global%2Fictv%2Fproposals%2F2015.006aM.A.v3.Rhabdoviridae_spren.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARabies+virus" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarterSaunders2007" class="citation book cs1">Carter, John; Saunders, Venetia (2007). <i>Virology: Principles and Applications</i>. Wiley. p. 175. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-470-02386-0" title="Special:BookSources/978-0-470-02386-0"><bdi>978-0-470-02386-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Virology%3A+Principles+and+Applications&rft.pages=175&rft.pub=Wiley&rft.date=2007&rft.isbn=978-0-470-02386-0&rft.aulast=Carter&rft.aufirst=John&rft.au=Saunders%2C+Venetia&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARabies+virus" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.who.int/news-room/fact-sheets/detail/rabies">"Rabies"</a>. <i>www.who.int</i><span class="reference-accessdate">. Retrieved <span class="nowrap">2021-09-10</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=www.who.int&rft.atitle=Rabies&rft_id=https%3A%2F%2Fwww.who.int%2Fnews-room%2Ffact-sheets%2Fdetail%2Frabies&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARabies+virus" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRiccardiGiacomelliAntonelloGobbi2021" class="citation journal cs1">Riccardi, Niccolò; Giacomelli, Andrea; Antonello, Roberta Maria; Gobbi, Federico; Angheben, Andrea (June 2021). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.ejim.2021.04.010">"Rabies in Europe: An epidemiological and clinical update"</a>. <i>European Journal of Internal Medicine</i>. <b>88</b>: 15–20. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.ejim.2021.04.010">10.1016/j.ejim.2021.04.010</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/33934971">33934971</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=European+Journal+of+Internal+Medicine&rft.atitle=Rabies+in+Europe%3A+An+epidemiological+and+clinical+update&rft.volume=88&rft.pages=15-20&rft.date=2021-06&rft_id=info%3Adoi%2F10.1016%2Fj.ejim.2021.04.010&rft_id=info%3Apmid%2F33934971&rft.aulast=Riccardi&rft.aufirst=Niccol%C3%B2&rft.au=Giacomelli%2C+Andrea&rft.au=Antonello%2C+Roberta+Maria&rft.au=Gobbi%2C+Federico&rft.au=Angheben%2C+Andrea&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.ejim.2021.04.010&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARabies+virus" class="Z3988"></span></span> </li> <li id="cite_note-pmid15885837-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-pmid15885837_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-pmid15885837_5-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFinkeConzelmann2005" class="citation journal cs1">Finke S, Conzelmann KK (August 2005). "Replication strategies of rabies virus". <i>Virus Res</i>. <b>111</b> (2): 120–131. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.virusres.2005.04.004">10.1016/j.virusres.2005.04.004</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/15885837">15885837</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Virus+Res.&rft.atitle=Replication+strategies+of+rabies+virus&rft.volume=111&rft.issue=2&rft.pages=120-131&rft.date=2005-08&rft_id=info%3Adoi%2F10.1016%2Fj.virusres.2005.04.004&rft_id=info%3Apmid%2F15885837&rft.aulast=Finke&rft.aufirst=S&rft.au=Conzelmann%2C+KK&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARabies+virus" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/nuccore/?term=rabies+complete+genome">"Rabies complete genome"</a>. <a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a> Nucleotide Database<span class="reference-accessdate">. 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(identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/33088593">33088593</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=npj+Vaccines&rft.atitle=Rabies+virus-based+COVID-19+vaccine+CORAVAX%E2%84%A2+induces+high+levels+of+neutralizing+antibodies+against+SARS-CoV-2&rft.volume=5&rft.issue=1&rft.pages=98&rft.date=2020-10-16&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7568577%23id-name%3DPMC&rft.issn=2059-0105&rft_id=info%3Apmid%2F33088593&rft_id=info%3Adoi%2F10.1038%2Fs41541-020-00248-6&rft.aulast=Kurup&rft.aufirst=Drishya&rft.au=Wirblich%2C+Christoph&rft.au=Ramage%2C+Holly&rft.au=Schnell%2C+Matthias+J.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7568577&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARabies+virus" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External 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eye</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cardiovascular_disease" title="Cardiovascular disease">Cardiovascular</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><i><a href="/wiki/Coxsackie_B_virus" title="Coxsackie B virus">CBV</a></i> <ul><li><a href="/wiki/Pericarditis" title="Pericarditis">Pericarditis</a></li> <li><a href="/wiki/Myocarditis" title="Myocarditis">Myocarditis</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Respiratory_system" title="Respiratory system">Respiratory system</a>/<br /><a href="/wiki/Common_cold" title="Common cold">acute viral<br />nasopharyngitis</a>/<br /><a href="/wiki/Viral_pneumonia" title="Viral pneumonia">viral pneumonia</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:1%"><a href="/wiki/DNA_virus" title="DNA virus">DNA virus</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><i><a href="/wiki/Epstein%E2%80%93Barr_virus" title="Epstein–Barr virus">Epstein–Barr virus</a></i> <ul><li><a href="/wiki/Epstein%E2%80%93Barr_virus_infection" title="Epstein–Barr virus infection">EBV infection</a>/<a href="/wiki/Infectious_mononucleosis" title="Infectious mononucleosis">Infectious mononucleosis</a></li></ul></li> <li><i><a href="/wiki/Cytomegalovirus" title="Cytomegalovirus">Cytomegalovirus</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/RNA_virus" title="RNA virus">RNA virus</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/RNA_virus#Group_IV_–_positive-sense_ssRNA_viruses" title="RNA virus">IV</a>: <i><a href="/wiki/Human_coronavirus" class="mw-redirect" title="Human coronavirus">Human coronavirus</a> <a href="/wiki/Human_coronavirus_229E" title="Human coronavirus 229E">229E</a>/<a href="/wiki/Human_coronavirus_NL63" title="Human coronavirus NL63">NL63</a>/<a href="/wiki/Human_coronavirus_HKU1" title="Human coronavirus HKU1">HKU1</a>/<a href="/wiki/Human_coronavirus_OC43" title="Human coronavirus OC43">OC43</a></i> <ul><li><a href="/wiki/Common_cold" title="Common cold">Common cold</a></li></ul></li> <li><i><a href="/wiki/Middle_East_respiratory_syndrome%E2%80%93related_coronavirus" class="mw-redirect" title="Middle East respiratory syndrome–related coronavirus">MERS coronavirus</a></i> <ul><li><a href="/wiki/Middle_East_respiratory_syndrome" class="mw-redirect" title="Middle East respiratory syndrome">Middle East respiratory syndrome</a></li></ul></li> <li><i><a href="/wiki/Severe_acute_respiratory_syndrome_coronavirus" class="mw-redirect" title="Severe acute respiratory syndrome coronavirus">SARS coronavirus</a></i> <ul><li><a href="/wiki/Severe_acute_respiratory_syndrome" class="mw-redirect" title="Severe acute respiratory syndrome">Severe acute respiratory syndrome</a></li></ul></li> <li><i><a href="/wiki/SARS-CoV-2" title="SARS-CoV-2">SARS-CoV-2</a></i> <ul><li><a href="/wiki/COVID-19" title="COVID-19">COVID-19</a></li></ul></li></ul> <ul><li><a href="/wiki/RNA_virus#Group_V_–_negative-sense_ssRNA_viruses" title="RNA virus">V</a>, <i><a href="/wiki/Orthomyxoviridae" title="Orthomyxoviridae">Orthomyxoviridae</a></i>: <i><a href="/wiki/Influenza_A_virus" title="Influenza A virus">Influenza virus A</a>/<a href="/wiki/Influenza_B_virus" title="Influenza B virus">B</a>/<a href="/wiki/Influenza_C_virus" title="Influenza C virus">C</a>/<a href="/wiki/Influenza_D_virus" title="Influenza D virus">D</a></i> <ul><li><a href="/wiki/Influenza" title="Influenza">Influenza</a>/<a href="/wiki/Avian_influenza" title="Avian influenza">Avian influenza</a></li></ul></li></ul> <ul><li>V, <i><a href="/wiki/Paramyxoviridae" title="Paramyxoviridae">Paramyxoviridae</a></i>: <i><a href="/wiki/Human_parainfluenza_viruses" title="Human parainfluenza viruses">Human parainfluenza viruses</a></i> <ul><li><a href="/wiki/Human_parainfluenza_viruses" title="Human parainfluenza viruses">Parainfluenza</a></li></ul></li> <li><i><a href="/wiki/Human_orthopneumovirus" class="mw-redirect" title="Human orthopneumovirus">Human orthopneumovirus</a></i></li> <li><i><a href="/wiki/Human_metapneumovirus" title="Human metapneumovirus">hMPV</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Human_digestive_system" title="Human digestive system">Human<br /> digestive system</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:1%"><a href="/wiki/Pharynx" title="Pharynx">Pharynx</a>/<a href="/wiki/Esophagus" title="Esophagus">Esophagus</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><i><a href="/wiki/Mumps_virus" title="Mumps virus">MuV</a></i> <ul><li><a href="/wiki/Mumps" title="Mumps">Mumps</a></li></ul></li> <li><i><a href="/wiki/Cytomegalovirus" title="Cytomegalovirus">Cytomegalovirus</a></i> <ul><li><a href="/wiki/Cytomegalovirus_esophagitis" title="Cytomegalovirus esophagitis">Cytomegalovirus esophagitis</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Gastroenteritis#Viral" title="Gastroenteritis">Gastroenteritis</a>/<br /><a href="/wiki/Gastroenteritis#Virus" title="Gastroenteritis">diarrhea</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <dl><dt><a href="/wiki/DNA_virus" title="DNA virus">DNA virus</a></dt> <dd><i><a href="/wiki/Adenoviridae" title="Adenoviridae">Adenovirus</a></i> <dl><dd><a href="/wiki/Adenovirus_infection" title="Adenovirus infection">Adenovirus infection</a></dd></dl></dd></dl> <dl><dt><a href="/wiki/RNA_virus" title="RNA virus">RNA virus</a></dt> <dd><i><a href="/wiki/Rotavirus" title="Rotavirus">Rotavirus</a></i> (<a href="/wiki/Rotaviral_gastroenteritis" class="mw-redirect" title="Rotaviral gastroenteritis">Gastroenteritis</a>)</dd> <dd><i><a href="/wiki/Norovirus" title="Norovirus">Norovirus</a></i></dd> <dd><i><a href="/wiki/Astrovirus" title="Astrovirus">Astrovirus</a></i></dd> <dd><i><a href="/wiki/Coronavirus" title="Coronavirus">Coronavirus</a></i></dd></dl> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Viral_hepatitis" title="Viral hepatitis">Hepatitis</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <dl><dt><a href="/wiki/DNA_virus" title="DNA virus">DNA virus</a></dt> <dd><i><a href="/wiki/Hepatitis_B_virus" title="Hepatitis B virus">HBV</a></i> (<a href="/wiki/Hepatitis_B" title="Hepatitis B">B</a>)</dd></dl> <dl><dt><a href="/wiki/RNA_virus" title="RNA virus">RNA virus</a></dt> <dd><i><a href="/wiki/Coxsackie_B_virus" title="Coxsackie B virus">CBV</a></i></dd> <dd><i><a href="/wiki/Hepatitis_A#Virology" title="Hepatitis A">HAV</a></i> (<a href="/wiki/Hepatitis_A" title="Hepatitis A">A</a>)</dd> <dd><i><a href="/wiki/Hepatitis_C_virus" title="Hepatitis C virus">HCV</a></i> (<a href="/wiki/Hepatitis_C" title="Hepatitis C">C</a>)</dd> <dd><i><a href="/wiki/Hepatitis_D" title="Hepatitis D">HDV</a></i> (<a href="/wiki/Hepatitis_D" title="Hepatitis D">D</a>)</dd> <dd><i><a href="/wiki/Orthohepevirus_A" title="Orthohepevirus A">HEV</a></i> (<a href="/wiki/Hepatitis_E" title="Hepatitis E">E</a>)</dd></dl> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Pancreatitis" title="Pancreatitis">Pancreatitis</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><i><a href="/wiki/Coxsackie_B_virus" title="Coxsackie B virus">CBV</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genitourinary_system" title="Genitourinary system">Urogenital</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><i><a href="/wiki/BK_virus" title="BK virus">BK virus</a></i></li> <li><i><a href="/wiki/Mumps_virus" title="Mumps virus">MuV</a></i> <ul><li><a href="/wiki/Mumps" title="Mumps">Mumps</a></li></ul></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_identifiers" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Taxon_identifiers" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Taxon_identifiers" title="Help:Taxon identifiers">Taxon identifiers</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;"><i>Rabies virus</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><span style="white-space:nowrap;"><a href="/wiki/Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q698976" class="extiw" title="wikidata:Q698976">Q698976</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Wikispecies" title="Wikispecies">Wikispecies</a>: <span class="uid"><span class="external"><a href="https://species.wikimedia.org/wiki/Rabies_lyssavirus" class="extiw" title="wikispecies:Rabies lyssavirus">Rabies lyssavirus</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://eol.org/pages/46700042">46700042</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.gbif.org/species/10111380">10111380</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&id=11460641">11460641</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/wiki/Invasive_Species_Compendium" title="Invasive Species Compendium">ISC</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.cabi.org/isc/datasheet/66456">66456</a></span></span></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 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