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data-event-name="pinnable-header.vector-toc.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">hide</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">(Top)</div> </a> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Detecting_viruses" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Detecting_viruses"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Detecting viruses</span> </div> </a> <button 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class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Serology"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Serology</span> </div> </a> <ul id="toc-Serology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Polymerase_chain_reaction_(PCR)_and_other_nucleic_acid_detection_methods" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Polymerase_chain_reaction_(PCR)_and_other_nucleic_acid_detection_methods"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Polymerase chain reaction (PCR) and other nucleic acid detection methods</span> </div> </a> <ul id="toc-Polymerase_chain_reaction_(PCR)_and_other_nucleic_acid_detection_methods-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Diagnostic_tests" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Diagnostic_tests"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>Diagnostic tests</span> </div> </a> <ul id="toc-Diagnostic_tests-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Quantitation_and_viral_loads" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Quantitation_and_viral_loads"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Quantitation and viral loads</span> </div> </a> <button aria-controls="toc-Quantitation_and_viral_loads-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 Quantitation and viral loads subsection</span> </button> <ul id="toc-Quantitation_and_viral_loads-sublist" class="vector-toc-list"> <li id="toc-Infectivity_assays" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Infectivity_assays"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Infectivity assays</span> </div> </a> <ul id="toc-Infectivity_assays-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Viral_load_assays" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Viral_load_assays"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Viral load assays</span> </div> </a> <ul id="toc-Viral_load_assays-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Molecular_biology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Molecular_biology"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Molecular biology</span> </div> </a> <button aria-controls="toc-Molecular_biology-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 Molecular biology subsection</span> </button> <ul id="toc-Molecular_biology-sublist" class="vector-toc-list"> <li id="toc-Purifying_viruses_and_their_components" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Purifying_viruses_and_their_components"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Purifying viruses and their components</span> </div> </a> <ul id="toc-Purifying_viruses_and_their_components-sublist" class="vector-toc-list"> <li id="toc-Centrifugation" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Centrifugation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1.1</span> <span>Centrifugation</span> </div> </a> <ul id="toc-Centrifugation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Electrophoresis" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Electrophoresis"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1.2</span> <span>Electrophoresis</span> </div> </a> <ul id="toc-Electrophoresis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Sequencing_of_viral_genomes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Sequencing_of_viral_genomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Sequencing of viral genomes</span> </div> </a> <ul id="toc-Sequencing_of_viral_genomes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Phylogenetic_analysis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Phylogenetic_analysis"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Phylogenetic analysis</span> </div> </a> <ul id="toc-Phylogenetic_analysis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cloning" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cloning"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Cloning</span> </div> </a> <ul id="toc-Cloning-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Phage_virology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Phage_virology"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5</span> <span>Phage virology</span> </div> </a> <ul id="toc-Phage_virology-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genetics" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Genetics"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Genetics</span> </div> </a> <button aria-controls="toc-Genetics-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 Genetics subsection</span> </button> <ul id="toc-Genetics-sublist" class="vector-toc-list"> <li id="toc-Reassortment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Reassortment"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Reassortment</span> </div> </a> <ul id="toc-Reassortment-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Recombination" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Recombination"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Recombination</span> </div> </a> <ul id="toc-Recombination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Reverse_genetics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Reverse_genetics"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>Reverse genetics</span> </div> </a> <ul id="toc-Reverse_genetics-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Virus_classification" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Virus_classification"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Virus classification</span> </div> </a> <button aria-controls="toc-Virus_classification-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 Virus classification subsection</span> </button> <ul id="toc-Virus_classification-sublist" class="vector-toc-list"> <li id="toc-ICTV_classification" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#ICTV_classification"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>ICTV classification</span> </div> </a> <ul id="toc-ICTV_classification-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Baltimore_classification" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Baltimore_classification"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Baltimore classification</span> </div> </a> <ul id="toc-Baltimore_classification-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>References</span> </div> </a> <button aria-controls="toc-References-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle References subsection</span> </button> <ul id="toc-References-sublist" class="vector-toc-list"> <li id="toc-Bibliography" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bibliography"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Bibliography</span> </div> </a> <ul id="toc-Bibliography-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Virology</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 77 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-77" 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">77 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/%D8%B9%D9%84%D9%85_%D8%A7%D9%84%D9%81%D9%8A%D8%B1%D9%88%D8%B3%D8%A7%D8%AA" title="علم الفيروسات – Arabic" lang="ar" hreflang="ar" data-title="علم الفيروسات" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Virusologiya" title="Virusologiya – Azerbaijani" lang="az" hreflang="az" data-title="Virusologiya" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AD%E0%A6%BE%E0%A6%87%E0%A6%B0%E0%A6%BE%E0%A6%B8%E0%A6%AC%E0%A6%BF%E0%A6%A6%E0%A7%8D%E0%A6%AF%E0%A6%BE" title="ভাইরাসবিদ্যা – Bangla" lang="bn" hreflang="bn" data-title="ভাইরাসবিদ্যা" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-ba mw-list-item"><a href="https://ba.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Вирусология – Bashkir" lang="ba" hreflang="ba" data-title="Вирусология" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target"><span>Башҡортса</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%92%D1%96%D1%80%D1%83%D1%81%D0%B0%D0%BB%D0%BE%D0%B3%D1%96%D1%8F" title="Вірусалогія – Belarusian" lang="be" hreflang="be" data-title="Вірусалогія" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Вирусология – Bulgarian" lang="bg" hreflang="bg" data-title="Вирусология" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Virusologija" title="Virusologija – Bosnian" lang="bs" hreflang="bs" data-title="Virusologija" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Virologia" title="Virologia – Catalan" lang="ca" hreflang="ca" data-title="Virologia" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cv mw-list-item"><a href="https://cv.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8" title="Вирусологи – Chuvash" lang="cv" hreflang="cv" data-title="Вирусологи" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target"><span>Чӑвашла</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Virologie" title="Virologie – Czech" lang="cs" hreflang="cs" data-title="Virologie" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-co mw-list-item"><a href="https://co.wikipedia.org/wiki/Virologia" title="Virologia – Corsican" lang="co" hreflang="co" data-title="Virologia" data-language-autonym="Corsu" data-language-local-name="Corsican" class="interlanguage-link-target"><span>Corsu</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Virologi" title="Virologi – Danish" lang="da" hreflang="da" data-title="Virologi" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Virologie" title="Virologie – German" lang="de" hreflang="de" data-title="Virologie" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Viroloogia" title="Viroloogia – Estonian" lang="et" hreflang="et" data-title="Viroloogia" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Virolog%C3%ADa" title="Virología – Spanish" lang="es" hreflang="es" data-title="Virología" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Virusscienco" title="Virusscienco – Esperanto" lang="eo" hreflang="eo" data-title="Virusscienco" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Birologia" title="Birologia – Basque" lang="eu" hreflang="eu" data-title="Birologia" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%88%DB%8C%D8%B1%D9%88%D8%B3%E2%80%8C%D8%B4%D9%86%D8%A7%D8%B3%DB%8C" title="ویروس‌شناسی – Persian" lang="fa" hreflang="fa" data-title="ویروس‌شناسی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Virologie" title="Virologie – French" lang="fr" hreflang="fr" data-title="Virologie" 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/Virolox%C3%ADa" title="Viroloxía – Galician" lang="gl" hreflang="gl" data-title="Viroloxía" 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/%EB%B0%94%EC%9D%B4%EB%9F%AC%EC%8A%A4%ED%95%99" title="바이러스학 – Korean" lang="ko" hreflang="ko" data-title="바이러스학" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%8E%D5%AB%D6%80%D5%B8%D6%82%D5%BD%D5%B8%D5%AC%D5%B8%D5%A3%D5%AB%D5%A1" title="Վիրուսոլոգիա – Armenian" lang="hy" hreflang="hy" data-title="Վիրուսոլոգիա" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%B5%E0%A4%BF%E0%A4%B7%E0%A4%BE%E0%A4%A3%E0%A5%81_%E0%A4%B5%E0%A4%BF%E0%A4%9C%E0%A5%8D%E0%A4%9E%E0%A4%BE%E0%A4%A8" title="विषाणु विज्ञान – Hindi" lang="hi" hreflang="hi" data-title="विषाणु विज्ञान" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Virologija" title="Virologija – Croatian" lang="hr" hreflang="hr" data-title="Virologija" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Virologio" title="Virologio – Ido" lang="io" hreflang="io" data-title="Virologio" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Virologi" title="Virologi – Indonesian" lang="id" hreflang="id" data-title="Virologi" 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-ie mw-list-item"><a href="https://ie.wikipedia.org/wiki/Virologie" title="Virologie – Interlingue" lang="ie" hreflang="ie" data-title="Virologie" data-language-autonym="Interlingue" data-language-local-name="Interlingue" class="interlanguage-link-target"><span>Interlingue</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Veirufr%C3%A6%C3%B0i" title="Veirufræði – Icelandic" lang="is" hreflang="is" data-title="Veirufræði" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Virologia" title="Virologia – Italian" lang="it" hreflang="it" data-title="Virologia" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></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%9C%D7%95%D7%92%D7%99%D7%94" title="וירולוגיה – Hebrew" lang="he" hreflang="he" data-title="וירולוגיה" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Virologi" title="Virologi – Javanese" lang="jv" hreflang="jv" data-title="Virologi" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%95%E1%83%98%E1%83%A0%E1%83%A3%E1%83%A1%E1%83%9D%E1%83%9A%E1%83%9D%E1%83%92%E1%83%98%E1%83%90" title="ვირუსოლოგია – Georgian" lang="ka" hreflang="ka" data-title="ვირუსოლოგია" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Вирусология – Kazakh" lang="kk" hreflang="kk" data-title="Вирусология" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Вирусология – Kyrgyz" lang="ky" hreflang="ky" data-title="Вирусология" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Virologia" title="Virologia – Latin" lang="la" hreflang="la" data-title="Virologia" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Virusolo%C4%A3ija" title="Virusoloģija – Latvian" lang="lv" hreflang="lv" data-title="Virusoloģija" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Virusologija" title="Virusologija – Lithuanian" lang="lt" hreflang="lt" data-title="Virusologija" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-li mw-list-item"><a href="https://li.wikipedia.org/wiki/Virologie" title="Virologie – Limburgish" lang="li" hreflang="li" data-title="Virologie" data-language-autonym="Limburgs" data-language-local-name="Limburgish" class="interlanguage-link-target"><span>Limburgs</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Virol%C3%B3gia" title="Virológia – Hungarian" lang="hu" hreflang="hu" data-title="Virológia" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></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%BB%D0%BE%D0%B3%D0%B8%D1%98%D0%B0" title="Вирологија – Macedonian" lang="mk" hreflang="mk" data-title="Вирологија" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></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%B3%E0%B4%9C%E0%B4%BF" title="വൈറോളജി – Malayalam" lang="ml" hreflang="ml" data-title="വൈറോളജി" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-xmf mw-list-item"><a href="https://xmf.wikipedia.org/wiki/%E1%83%95%E1%83%98%E1%83%A0%E1%83%A3%E1%83%A1%E1%83%9D%E1%83%9A%E1%83%9D%E1%83%92%E1%83%98%E1%83%90" title="ვირუსოლოგია – Mingrelian" lang="xmf" hreflang="xmf" data-title="ვირუსოლოგია" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target"><span>მარგალური</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Virologi" title="Virologi – Malay" lang="ms" hreflang="ms" data-title="Virologi" 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-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Virologie" title="Virologie – Dutch" lang="nl" hreflang="nl" data-title="Virologie" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%A6%E3%82%A4%E3%83%AB%E3%82%B9%E5%AD%A6" title="ウイルス学 – Japanese" lang="ja" hreflang="ja" data-title="ウイルス学" 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/Virologi" title="Virologi – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Virologi" 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-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Virologi" title="Virologi – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Virologi" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Virologia" title="Virologia – Occitan" lang="oc" hreflang="oc" data-title="Virologia" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Virusologiya" title="Virusologiya – Uzbek" lang="uz" hreflang="uz" data-title="Virusologiya" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target"><span>Oʻzbekcha / ўзбекча</span></a></li><li class="interlanguage-link interwiki-pnb mw-list-item"><a href="https://pnb.wikipedia.org/wiki/%D9%88%D8%A7%D8%A6%D8%B1%D9%88%D9%84%D9%88%D8%AC%DB%8C" title="وائرولوجی – Western Punjabi" lang="pnb" hreflang="pnb" data-title="وائرولوجی" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target"><span>پنجابی</span></a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Virologie" title="Virologie – Low German" lang="nds" hreflang="nds" data-title="Virologie" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target"><span>Plattdüütsch</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Wirusologia" title="Wirusologia – Polish" lang="pl" hreflang="pl" data-title="Wirusologia" 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/Virologia" title="Virologia – Portuguese" lang="pt" hreflang="pt" data-title="Virologia" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Virologie" title="Virologie – Romanian" lang="ro" hreflang="ro" data-title="Virologie" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</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%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Вирусология – Russian" lang="ru" hreflang="ru" data-title="Вирусология" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sah mw-list-item"><a href="https://sah.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Вирусология – Yakut" lang="sah" hreflang="sah" data-title="Вирусология" data-language-autonym="Саха тыла" data-language-local-name="Yakut" class="interlanguage-link-target"><span>Саха тыла</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Virologjia" title="Virologjia – Albanian" lang="sq" hreflang="sq" data-title="Virologjia" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B7%80%E0%B7%9B%E0%B6%BB%E0%B7%83%E0%B7%80%E0%B7%9A%E0%B6%AF%E0%B6%BA" title="වෛරසවේදය – Sinhala" lang="si" hreflang="si" data-title="වෛරසවේදය" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target"><span>සිංහල</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Virology" title="Virology – Simple English" lang="en-simple" hreflang="en-simple" data-title="Virology" 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-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Virol%C3%B3gia" title="Virológia – Slovak" lang="sk" hreflang="sk" data-title="Virológia" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%98%D0%B0" title="Вирусологија – Serbian" lang="sr" hreflang="sr" data-title="Вирусологија" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Virologija" title="Virologija – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Virologija" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Virologia" title="Virologia – Finnish" lang="fi" hreflang="fi" data-title="Virologia" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Virologi" title="Virologi – Swedish" lang="sv" hreflang="sv" data-title="Virologi" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Birolohiya" title="Birolohiya – Tagalog" lang="tl" hreflang="tl" data-title="Birolohiya" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-tt mw-list-item"><a href="https://tt.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Вирусология – Tatar" lang="tt" hreflang="tt" data-title="Вирусология" data-language-autonym="Татарча / tatarça" data-language-local-name="Tatar" class="interlanguage-link-target"><span>Татарча / tatarça</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%A7%E0%B8%B4%E0%B8%97%E0%B8%A2%E0%B8%B2%E0%B9%84%E0%B8%A7%E0%B8%A3%E0%B8%B1%E0%B8%AA" title="วิทยาไวรัส – Thai" lang="th" hreflang="th" data-title="วิทยาไวรัส" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tg mw-list-item"><a href="https://tg.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D1%88%D0%B8%D0%BD%D0%BE%D1%81%D3%A3" title="Вирусшиносӣ – Tajik" lang="tg" hreflang="tg" data-title="Вирусшиносӣ" data-language-autonym="Тоҷикӣ" data-language-local-name="Tajik" class="interlanguage-link-target"><span>Тоҷикӣ</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Viroloji" title="Viroloji – Turkish" lang="tr" hreflang="tr" data-title="Viroloji" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</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%D0%BE%D0%BB%D0%BE%D0%B3%D1%96%D1%8F" title="Вірусологія – Ukrainian" lang="uk" hreflang="uk" data-title="Вірусологія" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%B9%D9%84%D9%85_%D8%A7%D9%84%D9%88%D8%A7%D8%A6%D8%B1%D8%B3" title="علم الوائرس – Urdu" lang="ur" hreflang="ur" data-title="علم الوائرس" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Virus_h%E1%BB%8Dc" title="Virus học – Vietnamese" lang="vi" hreflang="vi" data-title="Virus học" 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-war mw-list-item"><a href="https://war.wikipedia.org/wiki/Virolohiya" title="Virolohiya – Waray" lang="war" hreflang="war" data-title="Virolohiya" data-language-autonym="Winaray" data-language-local-name="Waray" class="interlanguage-link-target"><span>Winaray</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E7%97%85%E6%AF%92%E5%AD%A6" title="病毒学 – Wu" lang="wuu" hreflang="wuu" data-title="病毒学" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-yi mw-list-item"><a href="https://yi.wikipedia.org/wiki/%D7%95%D7%95%D7%99%D7%A8%D7%90%D7%9C%D7%90%D7%92%D7%99%D7%A2" title="וויראלאגיע – Yiddish" lang="yi" hreflang="yi" data-title="וויראלאגיע" data-language-autonym="ייִדיש" data-language-local-name="Yiddish" class="interlanguage-link-target"><span>ייִדיש</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%97%85%E6%AF%92%E5%AD%B8" title="病毒學 – Cantonese" lang="yue" hreflang="yue" data-title="病毒學" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh 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id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Study of viruses</div> <p class="mw-empty-elt"> </p> <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 the journals, see <a href="/wiki/Virology_(journal)" title="Virology (journal)">Virology (journal)</a> and <a href="/wiki/Virology_Journal" title="Virology Journal">Virology Journal</a>.</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Gamma_phage.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Gamma_phage.png/220px-Gamma_phage.png" decoding="async" width="220" height="301" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Gamma_phage.png/330px-Gamma_phage.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Gamma_phage.png/440px-Gamma_phage.png 2x" data-file-width="1290" data-file-height="1763" /></a><figcaption>Gamma phage, an example of virus particles (visualised by electron microscopy)</figcaption></figure> <p><b>Virology</b> is the <a href="/wiki/Scientific_method" title="Scientific method">scientific study</a> of biological <a href="/wiki/Virus" title="Virus">viruses</a>. It is a subfield of <a href="/wiki/Microbiology" title="Microbiology">microbiology</a> that focuses on their detection, structure, classification and evolution, their methods of infection and exploitation of <a href="/wiki/Host_(biology)" title="Host (biology)">host</a> <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a> for reproduction, their interaction with host organism physiology and immunity, the diseases they cause, the techniques to isolate and culture them, and their use in research and therapy. </p><p>The identification of the causative agent of <a href="/wiki/Tobacco_mosaic_disease" class="mw-redirect" title="Tobacco mosaic disease">tobacco mosaic disease</a> (TMV) as a novel <a href="/wiki/Pathogen" title="Pathogen">pathogen</a> by <a href="/wiki/Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a> (1898) is now acknowledged as being the <a href="/wiki/History_of_virology" title="History of virology">official beginning of the field of virology as a discipline</a> distinct from <a href="/wiki/Bacteriology" title="Bacteriology">bacteriology</a>. He realized the source was neither a <a href="/wiki/Bacterial" class="mw-redirect" title="Bacterial">bacterial</a> nor a <a href="/wiki/Fungal" class="mw-redirect" title="Fungal">fungal</a> <a href="/wiki/Infection" title="Infection">infection</a>, but something completely different. Beijerinck used the word "virus" to describe the mysterious agent in his '<a href="/wiki/Contagium_vivum_fluidum" title="Contagium vivum fluidum">contagium vivum fluidum</a>' ('contagious living fluid'). <a href="/wiki/Rosalind_Franklin" title="Rosalind Franklin">Rosalind Franklin</a> proposed the full structure of the tobacco mosaic virus in 1955. </p><p>One main motivation for the study of viruses is because they cause many infectious diseases of plants and animals.<sup id="cite_ref-pmid22408703_1-0" class="reference"><a href="#cite_note-pmid22408703-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The study of the manner in which viruses cause disease is <a href="/wiki/Viral_pathogenesis" title="Viral pathogenesis">viral pathogenesis</a>. The degree to which a virus causes disease is its <a href="/wiki/Virulence" title="Virulence">virulence</a>.<sup id="cite_ref-pmid25462446_2-0" class="reference"><a href="#cite_note-pmid25462446-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> These fields of study are called <a href="/wiki/Plant_virology" class="mw-redirect" title="Plant virology">plant virology</a>, <a href="/wiki/Animal_virology" class="mw-redirect" title="Animal virology">animal virology</a> and human or <a href="/wiki/Medical_virology" class="mw-redirect" title="Medical virology">medical virology</a>.<sup id="cite_ref-ReferenceA_3-0" class="reference"><a href="#cite_note-ReferenceA-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Virology began when there were no methods for propagating or visualizing viruses or specific laboratory tests for viral infections. The methods for separating viral nucleic acids (<a href="/wiki/RNA" title="RNA">RNA</a> and <a href="/wiki/DNA" title="DNA">DNA</a>) and <a href="/wiki/Protein" title="Protein">proteins</a>, which are now the mainstay of virology, did not exist. Now there are many methods for observing the structure and functions of viruses and their component parts. Thousands of different viruses are now known about and virologists often specialize in either the viruses that infect plants, or <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and other <a href="/wiki/Microorganism" title="Microorganism">microorganisms</a>, or animals. Viruses that infect humans are now studied by medical virologists. Virology is a broad subject covering biology, health, animal welfare, agriculture and ecology. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/History_of_virology" title="History of virology">History of virology</a> and <a href="/wiki/Social_history_of_viruses" title="Social history of viruses">Social history of viruses</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Martinus_Willem_Beijerinck_in_his_laboratory.jpg" class="mw-file-description"><img alt="An old, bespectacled man wearing a suit and sitting at a bench by a large window. The bench is covered with small bottles and test tubes. On the wall behind him is a large old-fashioned clock below which are four small enclosed shelves on which sit many neatly labelled bottles." src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Martinus_Willem_Beijerinck_in_his_laboratory.jpg/220px-Martinus_Willem_Beijerinck_in_his_laboratory.jpg" decoding="async" width="220" height="307" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Martinus_Willem_Beijerinck_in_his_laboratory.jpg/330px-Martinus_Willem_Beijerinck_in_his_laboratory.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Martinus_Willem_Beijerinck_in_his_laboratory.jpg/440px-Martinus_Willem_Beijerinck_in_his_laboratory.jpg 2x" data-file-width="1312" data-file-height="1832" /></a><figcaption><a href="/wiki/Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a> in his laboratory in 1921</figcaption></figure> <p><a href="/wiki/Louis_Pasteur" title="Louis Pasteur">Louis Pasteur</a> was unable to find a causative agent for <a href="/wiki/Rabies" title="Rabies">rabies</a> and speculated about a pathogen too small to be detected by microscopes.<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> In 1884, the French <a href="/wiki/Microbiologist" title="Microbiologist">microbiologist</a> <a href="/wiki/Charles_Chamberland" title="Charles Chamberland">Charles Chamberland</a> invented the <a href="/wiki/Chamberland_filter" title="Chamberland filter">Chamberland filter</a> (or Pasteur-Chamberland filter) with pores small enough to remove all bacteria from a solution passed through it.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In 1892, the Russian biologist <a href="/wiki/Dmitri_Ivanovsky" title="Dmitri Ivanovsky">Dmitri Ivanovsky</a> used this filter to study what is now known as the <a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">tobacco mosaic virus</a>: crushed leaf extracts from infected tobacco plants remained infectious even after filtration to remove bacteria. Ivanovsky suggested the infection might be caused by a <a href="/wiki/Toxin" title="Toxin">toxin</a> produced by bacteria, but he did not pursue the idea.<sup id="cite_ref-Collier3_6-0" class="reference"><a href="#cite_note-Collier3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> At the time it was thought that all infectious agents could be retained by filters and grown on a nutrient medium—this was part of the <a href="/wiki/Germ_theory_of_disease" title="Germ theory of disease">germ theory of disease</a>.<sup id="cite_ref-Dimmock_7-0" class="reference"><a href="#cite_note-Dimmock-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>In 1898, the Dutch microbiologist <a href="/wiki/Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a> repeated the experiments and became convinced that the filtered solution contained a new form of infectious agent.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> He observed that the agent multiplied only in cells that were dividing, but as his experiments did not show that it was made of particles, he called it a <i><a href="/wiki/Contagium_vivum_fluidum" title="Contagium vivum fluidum">contagium vivum fluidum</a></i> (soluble living germ) and reintroduced the word <i>virus</i>. Beijerinck maintained that viruses were liquid in nature, a theory later discredited by <a href="/wiki/Wendell_Stanley" class="mw-redirect" title="Wendell Stanley">Wendell Stanley</a>, who proved they were particulate.<sup id="cite_ref-Collier3_6-1" class="reference"><a href="#cite_note-Collier3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> In the same year, <a href="/wiki/Friedrich_Loeffler" title="Friedrich Loeffler">Friedrich Loeffler</a> and Paul Frosch passed the first animal virus, <a href="/wiki/Aphthovirus" title="Aphthovirus">aphthovirus</a> (the agent of <a href="/wiki/Foot-and-mouth_disease" title="Foot-and-mouth disease">foot-and-mouth disease</a>), through a similar filter.<sup id="cite_ref-isbn0-12-375146-2_9-0" class="reference"><a href="#cite_note-isbn0-12-375146-2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>In the early 20th century, the English bacteriologist <a href="/wiki/Frederick_Twort" title="Frederick Twort">Frederick Twort</a> discovered a group of viruses that infect bacteria, now called <a href="/wiki/Bacteriophages" class="mw-redirect" title="Bacteriophages">bacteriophages</a><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> (or commonly 'phages'), and the French-Canadian microbiologist <a href="/wiki/F%C3%A9lix_d%27Herelle" class="mw-redirect" title="Félix d'Herelle">Félix d'Herelle</a> described viruses that, when added to bacteria on an <a href="/wiki/Agar_plate" title="Agar plate">agar plate</a>, would produce areas of dead bacteria. He accurately diluted a suspension of these viruses and discovered that the highest dilutions (lowest virus concentrations), rather than killing all the bacteria, formed discrete areas of dead organisms. Counting these areas and multiplying by the dilution factor allowed him to calculate the number of viruses in the original suspension.<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> Phages were heralded as a potential treatment for diseases such as <a href="/wiki/Typhoid" class="mw-redirect" title="Typhoid">typhoid</a> and <a href="/wiki/Cholera" title="Cholera">cholera</a>, but their promise was forgotten with the development of <a href="/wiki/Penicillin" title="Penicillin">penicillin</a>. The development of <a href="/wiki/Antimicrobial_resistance" title="Antimicrobial resistance">bacterial resistance to antibiotics</a> has renewed interest in the therapeutic use of bacteriophages.<sup id="cite_ref-pmid26891965_12-0" class="reference"><a href="#cite_note-pmid26891965-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>By the end of the 19th century, viruses were defined in terms of their <a href="/wiki/Infectivity" title="Infectivity">infectivity</a>, their ability to pass filters, and their requirement for living hosts. Viruses had been grown only in plants and animals. In 1906 <a href="/wiki/Ross_Granville_Harrison" title="Ross Granville Harrison">Ross Granville Harrison</a> invented a method for <a href="/wiki/Tissue_culture" title="Tissue culture">growing tissue</a> in <a href="/wiki/Lymph" title="Lymph">lymph</a>, and in 1913 E. Steinhardt, C. Israeli, and R.A. Lambert used this method to grow <a href="/wiki/Vaccinia" title="Vaccinia">vaccinia</a> virus in fragments of guinea pig corneal tissue.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> In 1928, H. B. Maitland and M. C. Maitland grew vaccinia virus in suspensions of minced hens' kidneys. Their method was not widely adopted until the 1950s when <a href="/wiki/Poliovirus" title="Poliovirus">poliovirus</a> was grown on a large scale for vaccine production.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>Another breakthrough came in 1931 when the American pathologist <a href="/wiki/Ernest_William_Goodpasture" title="Ernest William Goodpasture">Ernest William Goodpasture</a> and <a href="/wiki/Alice_Miles_Woodruff" title="Alice Miles Woodruff">Alice Miles Woodruff</a> grew influenza and several other viruses in fertilised chicken eggs.<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> In 1949, <a href="/wiki/John_Franklin_Enders" title="John Franklin Enders">John Franklin Enders</a>, <a href="/wiki/Thomas_Huckle_Weller" title="Thomas Huckle Weller">Thomas Weller</a>, and <a href="/wiki/Frederick_Robbins" class="mw-redirect" title="Frederick Robbins">Frederick Robbins</a> grew poliovirus in cultured cells from aborted human embryonic tissue,<sup id="cite_ref-WellerLibrary2004_16-0" class="reference"><a href="#cite_note-WellerLibrary2004-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> the first virus to be grown without using solid animal tissue or eggs. This work enabled <a href="/wiki/Hilary_Koprowski" title="Hilary Koprowski">Hilary Koprowski</a>, and then <a href="/wiki/Jonas_Salk" title="Jonas Salk">Jonas Salk</a>, to make an effective <a href="/wiki/Polio_vaccine" title="Polio vaccine">polio vaccine</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>The first images of viruses were obtained upon the invention of <a href="/wiki/Electron_microscopy" class="mw-redirect" title="Electron microscopy">electron microscopy</a> in 1931 by the German engineers <a href="/wiki/Ernst_Ruska" title="Ernst Ruska">Ernst Ruska</a> and <a href="/wiki/Max_Knoll" title="Max Knoll">Max Knoll</a>.<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> In 1935, American biochemist and virologist <a href="/wiki/Wendell_Meredith_Stanley" title="Wendell Meredith Stanley">Wendell Meredith Stanley</a> examined the tobacco mosaic virus and found it was mostly made of protein.<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> A short time later, this virus was separated into protein and RNA parts.<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 tobacco mosaic virus was the first to be <a href="/wiki/Crystal" title="Crystal">crystallised</a> and its structure could, therefore, be elucidated in detail. The first <a href="/wiki/X-ray_diffraction" title="X-ray diffraction">X-ray diffraction</a> pictures of the crystallised virus were obtained by Bernal and Fankuchen in 1941. Based on her X-ray crystallographic pictures, <a href="/wiki/Rosalind_Franklin" title="Rosalind Franklin">Rosalind Franklin</a> discovered the full structure of the virus in 1955.<sup id="cite_ref-pmid18702397_21-0" class="reference"><a href="#cite_note-pmid18702397-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> In the same year, <a href="/wiki/Heinz_Fraenkel-Conrat" title="Heinz Fraenkel-Conrat">Heinz Fraenkel-Conrat</a> and <a href="/wiki/Robley_Williams" class="mw-redirect" title="Robley Williams">Robley Williams</a> showed that purified tobacco mosaic virus RNA and its protein coat can assemble by themselves to form functional viruses, suggesting that this simple mechanism was probably the means through which viruses were created within their host cells.<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> </p><p>The second half of the 20th century was the golden age of virus discovery, and most of the documented species of animal, plant, and bacterial viruses were discovered during these years.<sup id="cite_ref-pmid18446425_23-0" class="reference"><a href="#cite_note-pmid18446425-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> In 1957 <a href="/wiki/Arterivirus" class="mw-redirect" title="Arterivirus">equine arterivirus</a> and the cause of <a href="/wiki/Bovine_virus_diarrhea" class="mw-redirect" title="Bovine virus diarrhea">bovine virus diarrhoea</a> (a <a href="/wiki/Pestivirus" title="Pestivirus">pestivirus</a>) were discovered. In 1963 the <a href="/wiki/Hepatitis_B" title="Hepatitis B">hepatitis B virus</a> was discovered by <a href="/wiki/Baruch_Blumberg" class="mw-redirect" title="Baruch Blumberg">Baruch Blumberg</a>,<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> and in 1965 <a href="/wiki/Howard_Temin" class="mw-redirect" title="Howard Temin">Howard Temin</a> described the first <a href="/wiki/Retrovirus" title="Retrovirus">retrovirus</a>. <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">Reverse transcriptase</a>, the <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> that retroviruses use to make DNA copies of their RNA, was first described in 1970 by Temin and <a href="/wiki/David_Baltimore" title="David Baltimore">David Baltimore</a> independently.<sup id="cite_ref-pmid4348509_25-0" class="reference"><a href="#cite_note-pmid4348509-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> In 1983 <a href="/wiki/Luc_Montagnier" title="Luc Montagnier">Luc Montagnier</a>'s team at the <a href="/wiki/Pasteur_Institute" title="Pasteur Institute">Pasteur Institute</a> in France, first isolated the retrovirus now called HIV.<sup id="cite_ref-Barre_26-0" class="reference"><a href="#cite_note-Barre-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> In 1989 <a href="/wiki/Michael_Houghton_(virologist)" class="mw-redirect" title="Michael Houghton (virologist)">Michael Houghton</a>'s team at <a href="/wiki/Chiron_Corporation" title="Chiron Corporation">Chiron Corporation</a> discovered <a href="/wiki/Hepatitis_C" title="Hepatitis C">hepatitis C</a>.<sup id="cite_ref-choo_27-0" class="reference"><a href="#cite_note-choo-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid19781804_28-0" class="reference"><a href="#cite_note-pmid19781804-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Detecting_viruses">Detecting viruses</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=2" title="Edit section: Detecting viruses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Jeol_Transmission_and_scanning_EM.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Jeol_Transmission_and_scanning_EM.jpg/220px-Jeol_Transmission_and_scanning_EM.jpg" decoding="async" width="220" height="320" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Jeol_Transmission_and_scanning_EM.jpg/330px-Jeol_Transmission_and_scanning_EM.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/39/Jeol_Transmission_and_scanning_EM.jpg/440px-Jeol_Transmission_and_scanning_EM.jpg 2x" data-file-width="1682" data-file-height="2443" /></a><figcaption>An electron microscope</figcaption></figure> <p>There are several approaches to detecting viruses and these include the detection of virus particles (virions) or their <a href="/wiki/Antigens" class="mw-redirect" title="Antigens">antigens</a> or nucleic acids and infectivity assays. </p> <div class="mw-heading mw-heading3"><h3 id="Electron_microscopy">Electron microscopy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=3" title="Edit section: Electron microscopy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Gastroenteritis_viruses.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/71/Gastroenteritis_viruses.jpg/220px-Gastroenteritis_viruses.jpg" decoding="async" width="220" height="172" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/71/Gastroenteritis_viruses.jpg/330px-Gastroenteritis_viruses.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/7/71/Gastroenteritis_viruses.jpg 2x" data-file-width="419" data-file-height="328" /></a><figcaption>Electron micrographs of viruses. A, rotavirus; B, adenovirus; C, norovirus; and D, astrovirus.</figcaption></figure> <p>Viruses were seen for the first time in the 1930s when electron microscopes were invented. These microscopes use beams of <a href="/wiki/Electron" title="Electron">electrons</a> instead of light, which have a much shorter wavelength and can detect objects that cannot be seen using light microscopes. The highest magnification obtainable by electron microscopes is up to 10,000,000 times<sup id="cite_ref-Payne_29-0" class="reference"><a href="#cite_note-Payne-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> whereas for light microscopes it is around 1,500 times.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>Virologists often use <a href="/wiki/Negative_staining#Transmission_electron_microscopy" class="mw-redirect" title="Negative staining">negative staining</a> to help visualise viruses. In this procedure, the viruses are suspended in a solution of metal salts such as uranium acetate. The atoms of metal are opaque to electrons and the viruses are seen as suspended in a dark background of metal atoms.<sup id="cite_ref-Payne_29-1" class="reference"><a href="#cite_note-Payne-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> This technique has been in use since the 1950s.<sup id="cite_ref-pmid13804200_31-0" class="reference"><a href="#cite_note-pmid13804200-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Many viruses were discovered using this technique and negative staining electron microscopy is still a valuable weapon in a virologist's arsenal.<sup id="cite_ref-pmid19822888_32-0" class="reference"><a href="#cite_note-pmid19822888-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>Traditional electron microscopy has disadvantages in that viruses are damaged by drying in the high vacuum inside the electron microscope and the electron beam itself is destructive.<sup id="cite_ref-Payne_29-2" class="reference"><a href="#cite_note-Payne-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Cryogenic_electron_microscopy" title="Cryogenic electron microscopy">cryogenic electron microscopy</a> the structure of viruses is preserved by embedding them in an environment of <a href="/wiki/Vitreous_water" class="mw-redirect" title="Vitreous water">vitreous water</a>.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> This allows the determination of biomolecular structures at near-atomic resolution,<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> and has attracted wide attention to the approach as an alternative to <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a> or <a href="/wiki/NMR_spectroscopy" class="mw-redirect" title="NMR spectroscopy">NMR spectroscopy</a> for the determination of the structure of viruses.<sup id="cite_ref-Stoddart_35-0" class="reference"><a href="#cite_note-Stoddart-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Rotavirus_Reconstruction.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/220px-Rotavirus_Reconstruction.jpg" decoding="async" width="220" height="193" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/330px-Rotavirus_Reconstruction.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/440px-Rotavirus_Reconstruction.jpg 2x" data-file-width="1986" data-file-height="1739" /></a><figcaption>Cryoelectron micrograph of a rotavirus</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Growth_in_cultures">Growth in cultures</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=4" title="Edit section: Growth in cultures"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Viruses are obligate intracellular parasites and because they only reproduce inside the living cells of a host these cells are needed to grow them in the laboratory. For viruses that infect animals (usually called "animal viruses") cells grown in laboratory <a href="/wiki/Cell_culture" title="Cell culture">cell cultures</a> are used. In the past, fertile hens' eggs were used and the viruses were grown on the membranes surrounding the embryo. This method is still used in the manufacture of some vaccines. For the viruses that infect bacteria, the <a href="/wiki/Bacteriophages" class="mw-redirect" title="Bacteriophages">bacteriophages</a>, the bacteria growing in test tubes can be used directly. For plant viruses, the natural host plants can be used or, particularly when the infection is not obvious, so-called indicator plants, which show signs of infection more clearly.<sup id="cite_ref-pmid19653216_36-0" class="reference"><a href="#cite_note-pmid19653216-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid35050076_37-0" class="reference"><a href="#cite_note-pmid35050076-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:CPE_rounding.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/25/CPE_rounding.jpg/220px-CPE_rounding.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/25/CPE_rounding.jpg/330px-CPE_rounding.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/25/CPE_rounding.jpg/440px-CPE_rounding.jpg 2x" data-file-width="1200" data-file-height="900" /></a><figcaption>Cytopathic effect of herpes simplex virus. The infected cells have become round and balloon-like.</figcaption></figure> <p>Viruses that have grown in cell cultures can be indirectly detected by the detrimental effect they have on the host cell. These <a href="/wiki/Cytopathic_effect" title="Cytopathic effect">cytopathic effects</a> are often characteristic of the type of virus. For instance, <a href="/wiki/Herpes_simplex_virus" title="Herpes simplex virus">herpes simplex viruses</a> produce a characteristic "ballooning" of the cells, typically human <a href="/wiki/Fibroblast" title="Fibroblast">fibroblasts</a>. Some viruses, such as <a href="/wiki/Mumps_virus" title="Mumps virus">mumps virus</a> cause red blood cells from chickens to firmly attach to the infected cells. This is called "haemadsorption" or "hemadsorption". Some viruses produce localised "lesions" in cell layers called <a href="/wiki/Viral_plaque" title="Viral plaque">plaques</a>, which are useful in quantitation assays and in identifying the species of virus by <a href="/wiki/Plaque_forming_units" class="mw-redirect" title="Plaque forming units">plaque reduction assays</a>.<sup id="cite_ref-pmid24899440_38-0" class="reference"><a href="#cite_note-pmid24899440-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid32367358_39-0" class="reference"><a href="#cite_note-pmid32367358-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p><p>Viruses growing in cell cultures are used to measure their susceptibility to validated and novel <a href="/wiki/Antiviral_drug" title="Antiviral drug">antiviral drugs</a>.<sup id="cite_ref-pmid32056049_40-0" class="reference"><a href="#cite_note-pmid32056049-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Serology">Serology</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=5" title="Edit section: Serology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Viruses are <a href="/wiki/Antigens" class="mw-redirect" title="Antigens">antigens</a> that induce the production of <a href="/wiki/Antibodies" class="mw-redirect" title="Antibodies">antibodies</a> and these antibodies can be used in laboratories to study viruses. Related viruses often react with each other's antibodies and some viruses can be named based on the antibodies they react with. The use of the antibodies which were once exclusively derived from the serum (blood fluid) of animals is called <a href="/wiki/Serology" title="Serology">serology</a>.<sup id="cite_ref-pmid32342927_41-0" class="reference"><a href="#cite_note-pmid32342927-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Once an antibody–reaction has taken place in a test, other methods are needed to confirm this. Older methods included <a href="/wiki/Complement_fixation_test" title="Complement fixation test">complement fixation tests</a>,<sup id="cite_ref-pmid1365549_42-0" class="reference"><a href="#cite_note-pmid1365549-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Hemagglutination_assay" title="Hemagglutination assay">hemagglutination inhibition</a> and <a href="/wiki/Neutralizing_antibody" title="Neutralizing antibody">virus neutralisation</a>.<sup id="cite_ref-pmid21887208_43-0" class="reference"><a href="#cite_note-pmid21887208-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Newer methods use <a href="/wiki/ELISA" title="ELISA">enzyme immunoassays</a> (EIA).<sup id="cite_ref-pmid22185616_44-0" class="reference"><a href="#cite_note-pmid22185616-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>In the years before <a href="/wiki/Polymerase_chain_reaction" title="Polymerase chain reaction">PCR</a> was invented <a href="/wiki/Immunofluorescence" title="Immunofluorescence">immunofluorescence</a> was used to quickly confirm viral infections. It is an infectivity assay that is virus species specific because antibodies are used. The antibodies are tagged with a dye that is luminescencent and when using an optical microscope with a modified light source, infected cells glow in the dark.<sup id="cite_ref-pmid32197545_45-0" class="reference"><a href="#cite_note-pmid32197545-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Polymerase_chain_reaction_(PCR)_and_other_nucleic_acid_detection_methods"><span id="Polymerase_chain_reaction_.28PCR.29_and_other_nucleic_acid_detection_methods"></span>Polymerase chain reaction (PCR) and other nucleic acid detection methods</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=6" title="Edit section: Polymerase chain reaction (PCR) and other nucleic acid detection methods"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>PCR is a mainstay method for detecting viruses in all species including plants and animals. It works by detecting traces of virus specific RNA or DNA. It is very sensitive and specific, but can be easily compromised by contamination. Most of the tests used in veterinary virology and medical virology are based on PCR or similar methods such as <a href="/wiki/Transcription_mediated_amplification" class="mw-redirect" title="Transcription mediated amplification">transcription mediated amplification</a>. When a novel virus emerges, such as the covid coronavirus, a specific test can be devised quickly so long as the viral genome has been sequenced and unique regions of the viral DNA or RNA identified.<sup id="cite_ref-pmid32815744_46-0" class="reference"><a href="#cite_note-pmid32815744-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> The invention of <a href="/wiki/Microfluidic" class="mw-redirect" title="Microfluidic">microfluidic</a> tests as allowed for most of these tests to be automated,<sup id="cite_ref-pmid35300739_47-0" class="reference"><a href="#cite_note-pmid35300739-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> Despite its specificity and sensitivity, PCR has a disadvantage in that it does not differentiate infectious and non-infectious viruses and "tests of cure" have to be delayed for up to 21 days to allow for residual viral nucleic acid to clear from the site of the infection.<sup id="cite_ref-pmid33219449_48-0" class="reference"><a href="#cite_note-pmid33219449-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Diagnostic_tests">Diagnostic tests</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=7" title="Edit section: Diagnostic tests"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In laboratories many of the diagnostic test for detecting viruses are nucleic acid amplification methods such as PCR. Some tests detect the viruses or their components as these include electron microscopy and <a href="/wiki/ELISA" title="ELISA">enzyme-immunoassays</a>. The so-called "home" or "self"-testing gadgets are usually <a href="/wiki/COVID-19_testing#Antigen_tests" title="COVID-19 testing">lateral flow tests</a>, which detect the virus using a tagged <a href="/wiki/Monoclonal_antibody" title="Monoclonal antibody">monoclonal antibody</a>.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> These are also used in agriculture, food and environmental sciences.<sup id="cite_ref-pmid27365041_50-0" class="reference"><a href="#cite_note-pmid27365041-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Quantitation_and_viral_loads">Quantitation and viral loads</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=8" title="Edit section: Quantitation and viral loads"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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/Virus_quantification" title="Virus quantification">Virus quantification</a></div> <p>Counting viruses (quantitation) has always had an important role in virology and has become central to the control of some infections of humans where the <a href="/wiki/Viral_load" title="Viral load">viral load</a> is measured.<sup id="cite_ref-pmid34369836_51-0" class="reference"><a href="#cite_note-pmid34369836-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> There are two basic methods: those that count the fully infective virus particles, which are called infectivity assays, and those that count all the particles including the defective ones.<sup id="cite_ref-Payne_29-3" class="reference"><a href="#cite_note-Payne-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Infectivity_assays">Infectivity assays</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=9" title="Edit section: Infectivity assays"><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:Plaque_assay_macro.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/Plaque_assay_macro.jpg/220px-Plaque_assay_macro.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/76/Plaque_assay_macro.jpg/330px-Plaque_assay_macro.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/76/Plaque_assay_macro.jpg/440px-Plaque_assay_macro.jpg 2x" data-file-width="600" data-file-height="450" /></a><figcaption>Plaques in cells caused herpes simplex virus. The cells have been fixed and stained blue.</figcaption></figure> <p>Infectivity assays measure the amount (concentration) of infective viruses in a sample of known volume.<sup id="cite_ref-pmid29925033_52-0" class="reference"><a href="#cite_note-pmid29925033-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> For host cells, plants or cultures of bacterial or animal cells are used. Laboratory animals such as mice have also been used particularly in veterinary virology.<sup id="cite_ref-pmid15585191_53-0" class="reference"><a href="#cite_note-pmid15585191-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> These are assays are either quantitative where the results are on a continuous scale or quantal, where an event either occurs or it does not. Quantitative assays give <a href="/wiki/Absolute_value" title="Absolute value">absolute values</a> and quantal assays give a statistical probability such as the volume of the test sample needed to ensure 50% of the hosts cells, plants or animals are infected. This is called the median infectious dose or <a href="/wiki/Minimal_infective_dose" title="Minimal infective dose">ID <sub>50</sub></a>.<sup id="cite_ref-pmid21474258_54-0" class="reference"><a href="#cite_note-pmid21474258-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> Infective bacteriophages can be counted by seeding them onto "lawns" of bacteria in culture dishes. When at low concentrations, the viruses form holes in the lawn that can be counted. The number of viruses is then expressed as <a href="/wiki/Plaque_forming_units" class="mw-redirect" title="Plaque forming units">plaque forming units</a>. For the bacteriophages that reproduce in bacteria that cannot be grown in cultures, viral load assays are used.<sup id="cite_ref-pmid33624268_55-0" class="reference"><a href="#cite_note-pmid33624268-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Virus_Infected_Cells.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Virus_Infected_Cells.jpg/220px-Virus_Infected_Cells.jpg" decoding="async" width="220" height="292" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Virus_Infected_Cells.jpg/330px-Virus_Infected_Cells.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/07/Virus_Infected_Cells.jpg/440px-Virus_Infected_Cells.jpg 2x" data-file-width="1834" data-file-height="2433" /></a><figcaption>Immunoflourescence: Cells infected by <a href="/wiki/Rotavirus" title="Rotavirus">rotavirus</a> (top) and uninfected cells (bottom)</figcaption></figure> <p>The focus forming assay (FFA) is a variation of the plaque assay, but instead of relying on cell lysis in order to detect plaque formation, the FFA employs <a href="/wiki/Immunostaining" title="Immunostaining">immunostaining</a> techniques using fluorescently labeled <a href="/wiki/Antibodies" class="mw-redirect" title="Antibodies">antibodies</a> specific for a viral <a href="/wiki/Antigen" title="Antigen">antigen</a> to detect infected host cells and infectious virus particles before an actual plaque is formed. The FFA is particularly useful for quantifying classes of viruses that do not lyse the cell membranes, as these viruses would not be amenable to the plaque assay. Like the plaque assay, host cell monolayers are infected with various dilutions of the virus sample and allowed to incubate for a relatively brief incubation period (e.g., 24–72 hours) under a semisolid overlay medium that restricts the spread of infectious virus, creating localized clusters (foci) of infected cells. Plates are subsequently probed with fluorescently labeled antibodies against a viral antigen, and fluorescence microscopy is used to count and quantify the number of foci. The FFA method typically yields results in less time than plaque or fifty-percent-tissue-culture-infective-dose (TCID<sub>50</sub>) assays, but it can be more expensive in terms of required reagents and equipment. Assay completion time is also dependent on the size of area that the user is counting. A larger area will require more time but can provide a more accurate representation of the sample. Results of the FFA are expressed as focus forming units per milliliter, or FFU/<sup id="cite_ref-pmid28701394_56-0" class="reference"><a href="#cite_note-pmid28701394-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Viral_load_assays">Viral load assays</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=10" title="Edit section: Viral load assays"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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/Viral_load" title="Viral load">Viral load</a></div> <p>When an assay for measuring the infective virus particle is done (Plaque assay, Focus assay), viral titre often refers to the concentration of infectious viral particles, which is different from the total viral particles. Viral load assays usually count the number of viral genomes present rather than the number of particles and use methods similar to <a href="/wiki/Polymerase_chain_reaction" title="Polymerase chain reaction">PCR</a>.<sup id="cite_ref-pmid33627730_57-0" class="reference"><a href="#cite_note-pmid33627730-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Viral load tests are an important in the control of infections by HIV.<sup id="cite_ref-pmid31515967_58-0" class="reference"><a href="#cite_note-pmid31515967-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> This versatile method can be used for plant viruses.<sup id="cite_ref-pmid28315718_59-0" class="reference"><a href="#cite_note-pmid28315718-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid32765569_60-0" class="reference"><a href="#cite_note-pmid32765569-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Molecular_biology">Molecular biology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=11" title="Edit section: Molecular biology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Molecular virology is the study of viruses at the level of nucleic acids and proteins. The methods invented by <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biologists</a> have all proven useful in virology. Their small sizes and relatively simple structures make viruses an ideal candidate for study by these techniques. </p> <div class="mw-heading mw-heading3"><h3 id="Purifying_viruses_and_their_components">Purifying viruses and their components</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=12" title="Edit section: Purifying viruses and their components"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:CsCl_density_gradient_centrifugation.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/CsCl_density_gradient_centrifugation.jpg/220px-CsCl_density_gradient_centrifugation.jpg" decoding="async" width="220" height="516" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/CsCl_density_gradient_centrifugation.jpg/330px-CsCl_density_gradient_centrifugation.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/11/CsCl_density_gradient_centrifugation.jpg/440px-CsCl_density_gradient_centrifugation.jpg 2x" data-file-width="1217" data-file-height="2857" /></a><figcaption>Caesium chloride (CsCl) solution and two morphological types of <a href="/wiki/Rotavirus" title="Rotavirus">rotavirus</a>. Following centrifugation at 100 g a density gradient forms in the CsCl solution and the virus particles separate according to their densities. The tube is 10 cm tall. The viruses are the two "milky" zones close together.<sup id="cite_ref-pmid6290520_61-0" class="reference"><a href="#cite_note-pmid6290520-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup></figcaption></figure><p>For further study, viruses grown in the laboratory need purifying to remove contaminants from the host cells. The methods used often have the advantage of concentrating the viruses, which makes it easier to investigate them. </p><div class="mw-heading mw-heading4"><h4 id="Centrifugation">Centrifugation</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=13" title="Edit section: Centrifugation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Centrifuge" title="Centrifuge">Centrifuges</a> are often used to purify viruses. Low speed centrifuges, i.e. those with a top speed of 10,000 <a href="/wiki/Revolutions_per_minute" title="Revolutions per minute">revolutions per minute</a> (rpm) are not powerful enough to concentrate viruses, but <a href="/wiki/Ultracentrifuge" title="Ultracentrifuge">ultracentrifuges</a> with a top speed of around 100,000 rpm, are and this difference is used in a method called <a href="/wiki/Differential_centrifugation" title="Differential centrifugation">differential centrifugation</a>. In this method the larger and heavier contaminants are removed from a virus mixture by low speed centrifugation. The viruses, which are small and light and are left in suspension, are then concentrated by high speed centrifugation.<sup id="cite_ref-pmid32825599_62-0" class="reference"><a href="#cite_note-pmid32825599-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p><p>Following differential centrifugation, virus suspensions often remain contaminated with debris that has the same <a href="/wiki/Sedimentation_coefficient" title="Sedimentation coefficient">sedimentation coefficient</a> and are not removed by the procedure. In these cases a modification of centrifugation, called <a href="/wiki/Buoyant_density_centrifugation" title="Buoyant density centrifugation">buoyant density centrifugation</a>, is used. In this method the viruses recovered from differential centrifugation are centrifuged again at very high speed for several hours in dense solutions of sugars or salts that form a density gradient, from low to high, in the tube during the centrifugation. In some cases, preformed gradients are used where solutions of steadily decreasing density are carefully overlaid on each other. Like an object in the <a href="/wiki/Dead_Sea" title="Dead Sea">Dead Sea</a>, despite the centrifugal force the virus particles cannot sink into solutions that are more dense than they are and they form discrete layers of, often visible, concentrated viruses in the tube. Caesium chloride is often used for these solutions as it is relatively inert but easily self-forms a gradient when centrifuged at high speed in an ultracentrifuge.<sup id="cite_ref-pmid6290520_61-1" class="reference"><a href="#cite_note-pmid6290520-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> Buoyant density centrifugation can also be used to purify the components of viruses such as their nucleic acids or proteins.<sup id="cite_ref-pmid31043560_63-0" class="reference"><a href="#cite_note-pmid31043560-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Electrophoresis">Electrophoresis</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=14" title="Edit section: Electrophoresis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Coomassie_blue_stained_gel.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Coomassie_blue_stained_gel.png/200px-Coomassie_blue_stained_gel.png" decoding="async" width="200" height="435" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/Coomassie_blue_stained_gel.png/300px-Coomassie_blue_stained_gel.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/46/Coomassie_blue_stained_gel.png/400px-Coomassie_blue_stained_gel.png 2x" data-file-width="459" data-file-height="998" /></a><figcaption>Polyacrylamide gel electrophoresis of <a href="/wiki/Rotavirus" title="Rotavirus">rotavirus</a> proteins stained with Coomassie blue</figcaption></figure> <p>The separation of molecules based on their electric charge is called <a href="/wiki/Electrophoresis" title="Electrophoresis">electrophoresis</a>. Viruses and all their components can be separated and purified using this method. This is usually done in a supporting medium such as <a href="/wiki/Agarose" title="Agarose">agarose</a> and <a href="/wiki/Polyacrylamide_gel" class="mw-redirect" title="Polyacrylamide gel">polyacrylamide gels</a>. The separated molecules are revealed using stains such as <a href="/wiki/Coomassie_brilliant_blue" title="Coomassie brilliant blue">coomasie blue</a>, for proteins, or <a href="/wiki/Ethidium_bromide" title="Ethidium bromide">ethidium bromide</a> for nucleic acids. In some instances the viral components are rendered radioactive before electrophoresis and are revealed using photographic film in a process known as <a href="/wiki/Autoradiography" class="mw-redirect" title="Autoradiography">autoradiography</a>.<sup id="cite_ref-pmid20127703_64-0" class="reference"><a href="#cite_note-pmid20127703-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Sequencing_of_viral_genomes">Sequencing of viral genomes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=15" title="Edit section: Sequencing of viral genomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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/DNA_sequencing" title="DNA sequencing">DNA sequencing</a></div> <p>As most viruses are too small to be seen by a light microscope, sequencing is one of the main tools in virology to identify and study the virus. Traditional Sanger sequencing and next-generation sequencing (NGS) are used to sequence viruses in basic and clinical research, as well as for the diagnosis of emerging viral infections, molecular epidemiology of viral pathogens, and drug-resistance testing. There are more than 2.3 million unique viral sequences in GenBank.<sup id="cite_ref-:0_65-0" class="reference"><a href="#cite_note-:0-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> NGS has surpassed traditional Sanger as the most popular approach for generating viral genomes.<sup id="cite_ref-:0_65-1" class="reference"><a href="#cite_note-:0-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> Viral genome sequencing as become a central method in viral <a href="/wiki/Epidemiology" title="Epidemiology">epidemiology</a> and <a href="/wiki/Virus_classification" title="Virus classification">viral classification</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Phylogenetic_analysis">Phylogenetic analysis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=16" title="Edit section: Phylogenetic analysis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Data from the sequencing of viral genomes can be used to determine evolutionary relationships and this is called <a href="/wiki/Phylogenetic_analysis" class="mw-redirect" title="Phylogenetic analysis">phylogenetic analysis</a>.<sup id="cite_ref-pmid30531947_66-0" class="reference"><a href="#cite_note-pmid30531947-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Software, such as <a href="/wiki/PHYLIP" title="PHYLIP">PHYLIP</a>, is used to draw <a href="/wiki/Phylogenetic_trees" class="mw-redirect" title="Phylogenetic trees">phylogenetic trees</a>. This analysis is also used in studying the spread of viral infections in communities (<a href="/wiki/Epidemiology" title="Epidemiology">epidemiology</a>).<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cloning">Cloning</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=17" title="Edit section: Cloning"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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/Cloning_vector" title="Cloning vector">Cloning vector</a></div> <p>When purified viruses or viral components are needed for diagnostic tests or vaccines, cloning can be used instead of growing the viruses.<sup id="cite_ref-pmid32404960_68-0" class="reference"><a href="#cite_note-pmid32404960-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> At the start of the <a href="/wiki/COVID-19_pandemic" title="COVID-19 pandemic">COVID-19 pandemic</a> the availability of the <a href="/wiki/Severe_acute_respiratory_syndrome_coronavirus_2" class="mw-redirect" title="Severe acute respiratory syndrome coronavirus 2">severe acute respiratory syndrome coronavirus 2</a> RNA sequence enabled tests to be manufactured quickly.<sup id="cite_ref-pmid32365353_69-0" class="reference"><a href="#cite_note-pmid32365353-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> There are several proven methods for cloning viruses and their components. Small pieces of DNA called <a href="/wiki/Cloning_vector" title="Cloning vector">cloning vectors</a> are often used and the most common ones are laboratory modified <a href="/wiki/Plasmids" class="mw-redirect" title="Plasmids">plasmids</a> (small circular molecules of DNA produced by bacteria). The viral nucleic acid, or a part of it, is inserted in the plasmid, which is the copied many times over by bacteria. This <a href="/wiki/Recombinant_DNA" title="Recombinant DNA">recombinant DNA</a> can then be used to produce viral components without the need for native viruses.<sup id="cite_ref-pmid31219641_70-0" class="reference"><a href="#cite_note-pmid31219641-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Phage_virology">Phage virology</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=18" title="Edit section: Phage virology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The viruses that reproduce in bacteria, archaea and fungi are informally called "phages",<sup id="cite_ref-pmid16791793_71-0" class="reference"><a href="#cite_note-pmid16791793-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> and the ones that infect bacteria – <a href="/wiki/Bacteriophages" class="mw-redirect" title="Bacteriophages">bacteriophages</a> – in particular are useful in virology and biology in general.<sup id="cite_ref-pmid29853167_72-0" class="reference"><a href="#cite_note-pmid29853167-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> Bacteriophages were some of the first viruses to be discovered, early in the twentieth century,<sup id="cite_ref-pmid31216787_73-0" class="reference"><a href="#cite_note-pmid31216787-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> and because they are relatively easy to grow quickly in laboratories, much of our understanding of viruses originated by studying them.<sup id="cite_ref-pmid31216787_73-1" class="reference"><a href="#cite_note-pmid31216787-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> Bacteriophages, long known for their positive effects in the environment, are used in <a href="/wiki/Phage_display" title="Phage display">phage display</a> techniques for screening proteins DNA sequences. They are a powerful tool in molecular biology.<sup id="cite_ref-pmid33504115_74-0" class="reference"><a href="#cite_note-pmid33504115-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genetics">Genetics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=19" title="Edit section: Genetics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>All viruses have <a href="/wiki/Gene" title="Gene">genes</a> which are studied using <a href="/wiki/Genetics" title="Genetics">genetics</a>.<sup id="cite_ref-pmid29751021_75-0" class="reference"><a href="#cite_note-pmid29751021-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> All the techniques used in molecular biology, such as cloning, creating mutations <a href="/wiki/RNA_silencing" title="RNA silencing">RNA silencing</a> are used in viral genetics.<sup id="cite_ref-Bamford_Zuckerman_p._76-0" class="reference"><a href="#cite_note-Bamford_Zuckerman_p.-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Reassortment">Reassortment</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=20" title="Edit section: Reassortment"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Reassortment" title="Reassortment">Reassortment</a> is the switching of genes from different parents and it is particularly useful when studying the genetics of viruses that have segmented genomes (fragmented into two or more nucleic acid molecules) such as <a href="/wiki/Influenza_virus" class="mw-redirect" title="Influenza virus">influenza viruses</a> and <a href="/wiki/Rotavirus" title="Rotavirus">rotaviruses</a>. The genes that encode properties such as <a href="/wiki/Serotype" title="Serotype">serotype</a> can be identified in this way.<sup id="cite_ref-pmid27211789_77-0" class="reference"><a href="#cite_note-pmid27211789-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Recombination">Recombination</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=21" title="Edit section: Recombination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Often confused with reassortment, recombination is also the mixing of genes but the mechanism differs in that <i>stretches</i> of DNA or RNA molecules, as opposed to the full molecules, are joined during the RNA or DNA replication cycle. Recombination is not as common as reassortment in nature but it is a powerful tool in laboratories for studying the structure and functions of viral genes.<sup id="cite_ref-pmid22922205_78-0" class="reference"><a href="#cite_note-pmid22922205-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Reverse_genetics">Reverse genetics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=22" title="Edit section: Reverse genetics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Reverse_genetics" title="Reverse genetics">Reverse genetics</a> is a powerful research method in virology.<sup id="cite_ref-pmid24899418_79-0" class="reference"><a href="#cite_note-pmid24899418-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> In this procedure complementary DNA (cDNA) copies of virus genomes called "infectious clones" are used to produce genetically modified viruses that can be then tested for changes in say, virulence or transmissibility.<sup id="cite_ref-pmid33528730_80-0" class="reference"><a href="#cite_note-pmid33528730-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Virus_classification">Virus classification</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=23" title="Edit section: Virus classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">For how viruses are classified with relation to other living things, see <a href="/wiki/Tree_of_life_(biology)" title="Tree of life (biology)">Tree of life (biology)</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/Virus_classification" title="Virus classification">Virus classification</a></div> <p>A major branch of virology is <a href="/wiki/Virus_classification" title="Virus classification">virus classification</a>. It is artificial in that it is not based on evolutionary <a href="/wiki/Phylogenetics" title="Phylogenetics">phylogenetics</a> but it is based shared or distinguishing properties of viruses.<sup id="cite_ref-pmid32740831_81-0" class="reference"><a href="#cite_note-pmid32740831-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid25015480_82-0" class="reference"><a href="#cite_note-pmid25015480-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> It seeks to describe the diversity of viruses by naming and grouping them on the basis of similarities.<sup id="cite_ref-pmid30039318_83-0" class="reference"><a href="#cite_note-pmid30039318-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> In 1962, <a href="/wiki/Andr%C3%A9_Lwoff" class="mw-redirect" title="André Lwoff">André Lwoff</a>, <a href="/wiki/Robert_Horne_(virologist)" title="Robert Horne (virologist)">Robert Horne</a>, and Paul Tournier were the first to develop a means of virus classification, based on the <a href="/wiki/Linnaean_taxonomy" title="Linnaean taxonomy">Linnaean</a> hierarchical system.<sup id="cite_ref-pmid14467544_84-0" class="reference"><a href="#cite_note-pmid14467544-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> This system based classification on <a href="/wiki/Phylum" title="Phylum">phylum</a>, <a href="/wiki/Class_(biology)" title="Class (biology)">class</a>, <a href="/wiki/Order_(biology)" title="Order (biology)">order</a>, <a href="/wiki/Family_(biology)" title="Family (biology)">family</a>, <a href="/wiki/Genus" title="Genus">genus</a>, and <a href="/wiki/Species" title="Species">species</a>. Viruses were grouped according to their shared properties (not those of their hosts) and the type of nucleic acid forming their genomes.<sup id="cite_ref-pmid13931895_85-0" class="reference"><a href="#cite_note-pmid13931895-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> In 1966, the <a href="/wiki/International_Committee_on_Taxonomy_of_Viruses" title="International Committee on Taxonomy of Viruses">International Committee on Taxonomy of Viruses</a> (ICTV) was formed. The system proposed by Lwoff, Horne and Tournier was initially not accepted by the ICTV because the small genome size of viruses and their high rate of mutation made it difficult to determine their ancestry beyond order. As such, the <a href="/wiki/Baltimore_classification" title="Baltimore classification">Baltimore classification</a> system has come to be used to supplement the more traditional hierarchy.<sup id="cite_ref-pmid16105179_86-0" class="reference"><a href="#cite_note-pmid16105179-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> Starting in 2018, the ICTV began to acknowledge deeper evolutionary relationships between viruses that have been discovered over time and adopted a 15-rank classification system ranging from realm to species.<sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> Additionally, some species within the same genus are grouped into a <i>genogroup</i>.<sup id="cite_ref-pmid34226482_88-0" class="reference"><a href="#cite_note-pmid34226482-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid22286874_89-0" class="reference"><a href="#cite_note-pmid22286874-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="ICTV_classification">ICTV classification</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=24" title="Edit section: ICTV classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ICTV developed the current classification system and wrote guidelines that put a greater weight on certain virus properties to maintain family uniformity. A unified taxonomy (a universal system for classifying viruses) has been established. Only a small part of the total diversity of viruses has been studied.<sup id="cite_ref-90" class="reference"><a href="#cite_note-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> As of 2021, 6 realms, 10 kingdoms, 17 phyla, 2 subphyla, 39 classes, 65 orders, 8 suborders, <a href="/wiki/List_of_virus_families_and_subfamilies" title="List of virus families and subfamilies">233 families, 168 subfamilies</a>, <a href="/wiki/List_of_virus_genera" title="List of virus genera">2,606 genera, 84 subgenera</a>, and <a href="/wiki/List_of_virus_species" title="List of virus species">10,434 species</a> of viruses have been defined by the ICTV.<sup id="cite_ref-ictv2021_91-0" class="reference"><a href="#cite_note-ictv2021-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> </p><p>The general taxonomic structure of taxon ranges and the suffixes used in taxonomic names are shown hereafter. As of 2021, the ranks of subrealm, subkingdom, and subclass are unused, whereas all other ranks are in use.<sup id="cite_ref-ictv2021_91-1" class="reference"><a href="#cite_note-ictv2021-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> </p> <dl><dd><a href="/wiki/Realm_(virology)" title="Realm (virology)">Realm</a> (<i>-viria</i>) <dl><dd>Subrealm (<i>-vira</i>) <dl><dd><a href="/wiki/Kingdom_(biology)" title="Kingdom (biology)">Kingdom</a> (<i>-virae</i>) <dl><dd>Subkingdom (<i>-virites</i>) <dl><dd><a href="/wiki/Phylum_(biology)" class="mw-redirect" title="Phylum (biology)">Phylum</a> (<i>-viricota</i>) <dl><dd>Subphylum (<i>-viricotina</i>) <dl><dd><a href="/wiki/Class_(biology)" title="Class (biology)">Class</a> (<i>-viricetes</i>) <dl><dd>Subclass (<i>-viricetidae</i>) <dl><dd><a href="/wiki/Order_(biology)" title="Order (biology)">Order</a> (<i>-virales</i>) <dl><dd>Suborder (<i>-virineae</i>) <dl><dd><a href="/wiki/Family_(biology)" title="Family (biology)">Family</a> (<i>-viridae</i>) <dl><dd>Subfamily (<i>-virinae</i>) <dl><dd><a href="/wiki/Genus" title="Genus">Genus</a> (<i>-virus</i>) <dl><dd>Subgenus (<i>-virus</i>) <dl><dd><a href="/wiki/Species" title="Species">Species</a></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl></dd></dl> <div class="mw-heading mw-heading3"><h3 id="Baltimore_classification">Baltimore classification</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=25" title="Edit section: Baltimore classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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/Baltimore_classification" title="Baltimore classification">Baltimore classification</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:VirusBaltimoreClassification.svg" class="mw-file-description"><img alt="A diagram showing how the Baltimore Classification is based on a virus's DNA or RNA and method of mRNA synthesis" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/VirusBaltimoreClassification.svg/330px-VirusBaltimoreClassification.svg.png" decoding="async" width="330" height="204" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/VirusBaltimoreClassification.svg/495px-VirusBaltimoreClassification.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a5/VirusBaltimoreClassification.svg/660px-VirusBaltimoreClassification.svg.png 2x" data-file-width="626" data-file-height="387" /></a><figcaption>The Baltimore Classification of viruses is based on the method of viral <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> synthesis.</figcaption></figure> <p>The Nobel Prize-winning biologist <a href="/wiki/David_Baltimore" title="David Baltimore">David Baltimore</a> devised the <a href="/wiki/Baltimore_classification" title="Baltimore classification">Baltimore classification</a> system.<sup id="cite_ref-pmid34259570_92-0" class="reference"><a href="#cite_note-pmid34259570-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> </p><p>The Baltimore classification of viruses is based on the mechanism of <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> production. Viruses must generate mRNAs from their genomes to produce proteins and replicate themselves, but different mechanisms are used to achieve this in each virus family. Viral genomes may be single-stranded (ss) or double-stranded (ds), RNA or DNA, and may or may not use <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a> (RT). In addition, ssRNA viruses may be either <a href="/wiki/Sense_(molecular_biology)" title="Sense (molecular biology)">sense</a> (+) or antisense (−). This classification places viruses into seven groups: </p> <ul><li>I: <a href="/wiki/DsDNA_virus" class="mw-redirect" title="DsDNA virus">dsDNA viruses</a> (e.g. <a href="/wiki/Adenovirus" class="mw-redirect" title="Adenovirus">Adenoviruses</a>, <a href="/wiki/Herpesvirus" class="mw-redirect" title="Herpesvirus">Herpesviruses</a>, <a href="/wiki/Poxvirus" class="mw-redirect" title="Poxvirus">Poxviruses</a>)</li> <li>II: <a href="/wiki/SsDNA_virus" class="mw-redirect" title="SsDNA virus">ssDNA viruses</a> (+ strand or "sense") DNA (e.g. <a href="/wiki/Parvovirus" class="mw-redirect" title="Parvovirus">Parvoviruses</a>)</li> <li>III: <a href="/wiki/DsRNA_virus" class="mw-redirect" title="DsRNA virus">dsRNA viruses</a> (e.g. <a href="/wiki/Reovirus" class="mw-redirect" title="Reovirus">Reoviruses</a>)</li> <li>IV:<a href="/wiki/Positive-sense_ssRNA_virus" class="mw-redirect" title="Positive-sense ssRNA virus">(+)ssRNA viruses</a> (+ strand or sense) RNA (e.g. <a href="/wiki/Coronavirus" title="Coronavirus">Coronaviruses</a>, <a href="/wiki/Picornavirus" title="Picornavirus">Picornaviruses</a>, <a href="/wiki/Togavirus" class="mw-redirect" title="Togavirus">Togaviruses</a>)</li> <li>V: <a href="/wiki/Negative-sense_ssRNA_virus" class="mw-redirect" title="Negative-sense ssRNA virus">(−)ssRNA viruses</a> (− strand or antisense) RNA (e.g. <a href="/wiki/Orthomyxovirus" class="mw-redirect" title="Orthomyxovirus">Orthomyxoviruses</a>, <a href="/wiki/Rhabdovirus" class="mw-redirect" title="Rhabdovirus">Rhabdoviruses</a>)</li> <li>VI: <a href="/wiki/SsRNA-RT_virus" class="mw-redirect" title="SsRNA-RT virus">ssRNA-RT viruses</a> (+ strand or sense) RNA with DNA intermediate in life-cycle (e.g. <a href="/wiki/Retrovirus" title="Retrovirus">Retroviruses</a>)</li> <li>VII: <a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT viruses</a> DNA with RNA intermediate in life-cycle (e.g. <a href="/wiki/Hepadnavirus" class="mw-redirect" title="Hepadnavirus">Hepadnaviruses</a>)</li></ul> <style data-mw-deduplicate="TemplateStyles:r1239009302">.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{clear:left;float:left;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 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class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-pmid22408703-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-pmid22408703_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="CITEREFDoljaKoonin2011" class="citation journal cs1">Dolja VV, Koonin EV (November 2011). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3293486">"Common origins and host-dependent diversity of plant and animal viromes"</a>. <i>Current Opinion in Virology</i>. <b>1</b> (5): 322–31. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" 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Mahy B, Collier LA (eds.). <i>Topley and Wilson's Microbiology and Microbial Infections</i>. Virology. Vol. 1 (Ninth ed.). <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-340-66316-2" title="Special:BookSources/0-340-66316-2"><bdi>0-340-66316-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Topley+and+Wilson%27s+Microbiology+and+Microbial+Infections&rft.series=Virology&rft.edition=Ninth&rft.date=1998&rft.isbn=0-340-66316-2&rft.aulast=Collier&rft.aufirst=L&rft.au=Balows%2C+A&rft.au=Sussman%2C+M&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVirology" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDimmockEastonLeppard2007" class="citation book cs1">Dimmock NJ, Easton AJ, Leppard K (2007). <i>Introduction to Modern Virology</i> (Sixth ed.). Blackwell Publishing. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4051-3645-7" title="Special:BookSources/978-1-4051-3645-7"><bdi>978-1-4051-3645-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Introduction+to+Modern+Virology&rft.edition=Sixth&rft.pub=Blackwell+Publishing&rft.date=2007&rft.isbn=978-1-4051-3645-7&rft.aulast=Dimmock&rft.aufirst=NJ&rft.au=Easton%2C+AJ&rft.au=Leppard%2C+K&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVirology" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFShors2017" class="citation book cs1">Shors T (2017). <i>Understanding Viruses</i>. Jones and Bartlett Publishers. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-284-02592-7" title="Special:BookSources/978-1-284-02592-7"><bdi>978-1-284-02592-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Understanding+Viruses&rft.pub=Jones+and+Bartlett+Publishers&rft.date=2017&rft.isbn=978-1-284-02592-7&rft.aulast=Shors&rft.aufirst=T&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVirology" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Virology&action=edit&section=28" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Commons-logo.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></a></span> Media related to <a href="https://commons.wikimedia.org/wiki/Category:Virology" class="extiw" title="commons:Category:Virology">Virology</a> at Wikimedia Commons</li> <li><span class="official-website"><span class="url"><a rel="nofollow" class="external text" href="http://www.ictvonline.org/">Official website</a></span></span> of the International Committee on Taxonomy of Viruses</li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist 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.hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Biotechnology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output 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abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Biotechnology" title="Template:Biotechnology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Biotechnology" title="Template talk:Biotechnology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Biotechnology" title="Special:EditPage/Template:Biotechnology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Biotechnology" style="font-size:114%;margin:0 4em"><a href="/wiki/Biotechnology" title="Biotechnology">Biotechnology</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">History</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/History_of_biotechnology" title="History of biotechnology">History of biotechnology</a></li> <li><a href="/wiki/Timeline_of_biotechnology" title="Timeline of biotechnology">Timeline of biotechnology</a></li> <li><a href="/wiki/Competitions_and_prizes_in_biotechnology" title="Competitions and prizes in biotechnology">Competitions and prizes in biotechnology</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="8" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:DNA_replication_split.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/DNA_replication_split.svg/50px-DNA_replication_split.svg.png" decoding="async" width="50" height="101" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/DNA_replication_split.svg/75px-DNA_replication_split.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/70/DNA_replication_split.svg/100px-DNA_replication_split.svg.png 2x" data-file-width="315" data-file-height="635" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Branches</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/Colors_of_biotechnology" class="mw-redirect" title="Colors of biotechnology">Colors of biotechnology</a></li> <li><a href="/wiki/Industrial_biotechnology" class="mw-redirect" title="Industrial biotechnology">Industrial biotechnology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Biological concepts</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/Allele" title="Allele">Allele</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a></li> <li><a href="/wiki/DNA" title="DNA">DNA</a>/<a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Fermentation" title="Fermentation">Fermentation</a></li> <li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Protein" title="Protein">Protein</a></li> <li><a href="/wiki/Selective_breeding" title="Selective breeding">Selective breeding</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">General concepts</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/Biotechnology_industrial_park" class="mw-redirect" title="Biotechnology industrial park">Biotechnology industrial park</a></li> <li><a href="/wiki/Biotechnology_products" class="mw-redirect" title="Biotechnology products">Biotechnology products</a></li> <li><a href="/wiki/Biotechnology_law" class="mw-redirect" title="Biotechnology law">Biotechnology law</a></li> <li><a href="/wiki/Green_Revolution" title="Green Revolution">Green Revolution</a></li> <li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li> <li><a href="/wiki/Pharmaceutical_company" class="mw-redirect" title="Pharmaceutical company">Pharmaceutical company</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Basic techniques<br /> and tools</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%">Biology field</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/Bioreactor" title="Bioreactor">Bioreactor</a></li> <li><a href="/wiki/Cell_culture" title="Cell culture">Cell culture</a></li> <li><a href="/wiki/Cultured_meat" title="Cultured meat">Cultured meat</a></li> <li><a href="/wiki/Flow_cytometry" title="Flow cytometry">Flow cytometry</a></li> <li><a href="/wiki/Hybridoma_technology" title="Hybridoma technology">Hybridoma technology</a></li> <li><a href="/wiki/High-performance_liquid_chromatography" title="High-performance liquid chromatography">HPLC</a></li> <li><a href="/wiki/Nuclear_magnetic_resonance" title="Nuclear magnetic resonance">NMR</a></li> <li><a href="/wiki/Spectroscopy" title="Spectroscopy">Spectroscopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Chemical field</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/Centrifugation" title="Centrifugation">Centrifugation</a></li> <li><a href="/wiki/Continuous_stirred-tank_reactor" title="Continuous stirred-tank reactor">CSTR</a></li> <li><a href="/wiki/Crystallization" title="Crystallization">Crystallization</a></li> <li><a href="/wiki/Chromatography" title="Chromatography">Chromatography</a></li> <li><a href="/wiki/Kidney_dialysis" title="Kidney dialysis">Kidney dialysis</a></li> <li><a href="/wiki/Electrophoresis" title="Electrophoresis">Electrophoresis</a></li> <li><a href="/wiki/Extraction_(chemistry)" title="Extraction (chemistry)">Extraction</a></li> <li><a href="/wiki/Fed-batch_culture" title="Fed-batch culture">Fed Batch</a></li> <li><a href="/wiki/Filtration" title="Filtration">Filtration</a></li> <li><a href="/wiki/Plug_flow_reactor_model" title="Plug flow reactor model">PFR</a></li> <li><a href="/wiki/Sedimentation" title="Sedimentation">Sedimentation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Applications</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/Animal_cell_culture" class="mw-redirect" title="Animal cell culture">Animal cell culture</a></li> <li><a href="/wiki/Biofabrication" title="Biofabrication">Biofabrication</a></li> <li><a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</a></li> <li><a href="/wiki/Biosynthesis" title="Biosynthesis">Biosynthesis</a></li> <li><a href="/wiki/Bionic_architecture" title="Bionic architecture">Bionic architecture</a></li> <li><a href="/wiki/Cell_immunity" class="mw-redirect" title="Cell immunity">Cell immunity</a></li> <li><a href="/wiki/Cloning" title="Cloning">Cloning</a> <ul><li><a href="/wiki/Reproductive_cloning" class="mw-redirect" title="Reproductive cloning">Reproductive cloning</a></li> <li><a href="/wiki/Therapeutic_cloning" class="mw-redirect" title="Therapeutic cloning">Therapeutic cloning</a></li></ul></li> <li><a href="/wiki/Embryology" title="Embryology">Embryology</a></li> <li><a href="/wiki/Environmental_biotechnology" title="Environmental biotechnology">Environmental biotechnology</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a> <ul><li><a href="/wiki/Genetically_modified_organism" title="Genetically modified organism">Genetically modified organism</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular genetics</a></li></ul></li> <li><a href="/wiki/Gene_therapy" title="Gene therapy">Gene therapy</a></li> <li><a href="/wiki/Microbial_biodegradation" title="Microbial biodegradation">Microbial biodegradation</a></li> <li><a href="/wiki/Omics" title="Omics">Omics</a></li> <li><a href="/wiki/Pharmacogenomics" title="Pharmacogenomics">Pharmacogenomics</a></li> <li><a href="/wiki/Stem_cells" class="mw-redirect" title="Stem cells">Stem cells</a></li> <li><a href="/wiki/Telomere" title="Telomere">Telomere</a></li> <li><a href="/wiki/Tissue_culture" title="Tissue culture">Tissue culture</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Interdisciplinary <br />fields</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/Biobased_economy" class="mw-redirect" title="Biobased economy">Bioeconomy</a></li> <li><a href="/wiki/Bioelectronics" title="Bioelectronics">Bioelectronics</a></li> <li><a href="/wiki/Biological_engineering" title="Biological engineering">Bioengineering</a></li> <li><a href="/wiki/Biology" title="Biology">Biology</a></li> <li><a href="/wiki/Biopharmaceutical" title="Biopharmaceutical">Biopharmacology</a></li> <li><a href="/wiki/Biomedical_engineering" title="Biomedical engineering">Biomedical engineering</a></li> <li><a href="/wiki/Biomedicine" title="Biomedicine">Biomedicine</a></li> <li><a href="/wiki/Biomimetics" title="Biomimetics">Biomimetics</a></li> <li><a href="/wiki/Biochemistry" title="Biochemistry">Biochemicals</a></li> <li><a href="/wiki/Biorobotics" title="Biorobotics">Biorobotics</a></li> <li><a href="/wiki/Chemical_engineering" title="Chemical engineering">Chemical engineering</a></li> <li><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a></li> <li><a href="/wiki/Mining" title="Mining">Mining</a></li> <li><a href="/wiki/Molecular_biology" title="Molecular biology">Molecular biology</a></li> <li><a href="/wiki/Nanobiotechnology" title="Nanobiotechnology">Nanobiotechnology</a></li> <li><a class="mw-selflink selflink">Virology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</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/Index_of_biotechnology_articles" title="Index of biotechnology articles">Index of biotechnology articles</a></li> <li><a href="/wiki/List_of_biotechnology_articles" class="mw-redirect" title="List of biotechnology articles">List of biotechnology articles</a></li> <li><a href="/wiki/List_of_largest_biomedical_companies_by_market_capitalization" title="List of largest biomedical companies by market capitalization">List of largest biomedical companies by market capitalization</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Biotechnology" title="Category:Biotechnology">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span> <b><a href="https://commons.wikimedia.org/wiki/Category:Biotechnology" class="extiw" title="commons:Category:Biotechnology">Commons</a></b></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="Infectious_disease_and_microbiology" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse 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:Infectious_disease" title="Template:Infectious disease"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Infectious_disease" title="Template talk:Infectious disease"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Infectious_disease" title="Special:EditPage/Template:Infectious disease"><abbr title="Edit this template">e</abbr></a></li></ul></div><div 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title="Mycosis">fungi</a></li></ul></li> <li><a href="/wiki/Parasitology" title="Parasitology">Parasitology</a> <ul><li><a href="/wiki/Protozoa" title="Protozoa">protozoa</a></li> <li><a href="/wiki/Helminths" class="mw-redirect" title="Helminths">helminths</a></li></ul></li> <li><a href="/wiki/Medical_entomology" title="Medical entomology">Entomology</a> <ul><li><a href="/wiki/Ectoparasite" class="mw-redirect" title="Ectoparasite">ectoparasites</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="Branches_of_biology" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse 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:Branches_of_biology" title="Template:Branches of biology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Branches_of_biology" title="Template talk:Branches of biology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Branches_of_biology" title="Special:EditPage/Template:Branches of biology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Branches_of_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Outline_of_biology#Branches" title="Outline of biology">Branches of biology</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Aerobiology" title="Aerobiology">Aerobiology</a></li> <li><a href="/wiki/Agronomy" title="Agronomy">Agronomy</a></li> <li><a href="/wiki/Agrostology" title="Agrostology">Agrostology</a></li> <li><a href="/wiki/Anatomy" title="Anatomy">Anatomy</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Bacteriology" title="Bacteriology">Bacteriology</a></li> <li><a href="/wiki/Biochemistry" title="Biochemistry">Biochemistry</a></li> <li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Biogeology" title="Biogeology">Biogeology</a></li> <li><a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</a></li> <li><a href="/wiki/Biological_engineering" title="Biological engineering">Biological engineering</a></li> <li><a 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href="/wiki/Conservation_biology" title="Conservation biology">Conservation biology</a></li> <li><a href="/wiki/Cryobiology" title="Cryobiology">Cryobiology</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Dendrology" title="Dendrology">Dendrology</a></li> <li><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Ecology" title="Ecology">Ecology</a></li> <li><a href="/wiki/Embryology" title="Embryology">Embryology</a></li> <li><a href="/wiki/Epidemiology" title="Epidemiology">Epidemiology</a></li> <li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Freshwater_biology" title="Freshwater biology">Freshwater biology</a></li> <li><a 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href="/wiki/Mathematical_and_theoretical_biology" title="Mathematical and theoretical biology">Mathematical biology</a></li> <li><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a></li> <li><a href="/wiki/Molecular_biology" title="Molecular biology">Molecular biology</a></li> <li><a href="/wiki/Mycology" title="Mycology">Mycology</a></li> <li><a href="/wiki/Neontology" title="Neontology">Neontology</a></li> <li><a href="/wiki/Neuroscience" title="Neuroscience">Neuroscience</a></li> <li><a href="/wiki/Nutritional_science" title="Nutritional science">Nutrition</a></li> <li><a href="/wiki/Ornithology" title="Ornithology">Ornithology</a></li> <li><a href="/wiki/Osteology" title="Osteology">Osteology</a></li> <li><a href="/wiki/Paleontology" title="Paleontology">Paleontology</a></li> <li><a href="/wiki/Parasitology" title="Parasitology">Parasitology</a></li> <li><a href="/wiki/Pathology" title="Pathology">Pathology</a></li> <li><a href="/wiki/Pharmacology" title="Pharmacology">Pharmacology</a></li> <li><a href="/wiki/Photobiology" title="Photobiology">Photobiology</a></li> <li><a href="/wiki/Phycology" title="Phycology">Phycology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a></li> <li><a href="/wiki/Physiology" title="Physiology">Physiology</a></li> <li><a href="/wiki/Pomology" title="Pomology">Pomology</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a></li> <li><a href="/wiki/Proteomics" title="Proteomics">Proteomics</a></li> <li><a href="/wiki/Protistology" title="Protistology">Protistology</a></li> <li><a href="/wiki/Quantum_biology" title="Quantum biology">Quantum biology</a></li> <li><a href="/wiki/Relational_biology" class="mw-redirect" title="Relational biology">Relational biology</a></li> <li><a href="/wiki/Reproductive_biology" title="Reproductive biology">Reproductive biology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li> <li><a href="/wiki/Structural_biology" title="Structural biology">Structural biology</a></li> <li><a href="/wiki/Synthetic_biology" title="Synthetic biology">Synthetic biology</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Systems_biology" title="Systems biology">Systems biology</a></li> <li><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Taxonomy</a></li> <li><a href="/wiki/Teratology" title="Teratology">Teratology</a></li> <li><a href="/wiki/Toxicology" title="Toxicology">Toxicology</a></li> <li><a class="mw-selflink selflink">Virology</a></li> <li><a href="/wiki/Virophysics" title="Virophysics">Virophysics</a></li> <li><a href="/wiki/Xenobiology" title="Xenobiology">Xenobiology</a></li> <li><a href="/wiki/Zoology" title="Zoology">Zoology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</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/History_of_biology" title="History of biology">History of biology</a></li> <li><a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a></li> <li><a href="/wiki/Timeline_of_biology_and_organic_chemistry" title="Timeline of biology and organic chemistry">Timeline of biology and organic chemistry</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"><link rel="mw-deduplicated-inline-style" 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0.25em"><ul><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Virologie"><a rel="nofollow" class="external text" href="https://d-nb.info/gnd/4063597-1">Germany</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Virology"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85143799">United States</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Virologie"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb11939856g">France</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Virologie"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb11939856g">BnF data</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="virologie"><a rel="nofollow" class="external text" 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