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aria-controls="toc-Microbiology-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Microbiology subsection </button> <ul id="toc-Microbiology-sublist" class="vector-toc-list"> <li id="toc-Discovery" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Discovery"> <div class="vector-toc-text"> 3.1 Discovery </div> </a> <ul id="toc-Discovery-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Life_properties" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Life_properties"> <div class="vector-toc-text"> 3.2 Life properties </div> </a> <ul id="toc-Life_properties-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Structure"> <div class="vector-toc-text"> 3.3 Structure </div> </a> <ul id="toc-Structure-sublist" class="vector-toc-list"> <li id="toc-Giant_viruses" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Giant_viruses"> <div class="vector-toc-text"> 3.3.1 Giant viruses </div> </a> <ul id="toc-Giant_viruses-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genome" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genome"> <div class="vector-toc-text"> 3.4 Genome </div> </a> <ul id="toc-Genome-sublist" class="vector-toc-list"> <li id="toc-Genome_size" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Genome_size"> <div class="vector-toc-text"> 3.4.1 Genome size </div> </a> <ul id="toc-Genome_size-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genetic_mutation_and_recombination" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genetic_mutation_and_recombination"> <div class="vector-toc-text"> 3.5 Genetic mutation and recombination </div> </a> <ul id="toc-Genetic_mutation_and_recombination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Replication_cycle" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Replication_cycle"> <div class="vector-toc-text"> 3.6 Replication cycle </div> </a> <ul id="toc-Replication_cycle-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genome_replication" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genome_replication"> <div class="vector-toc-text"> 3.7 Genome replication </div> </a> <ul id="toc-Genome_replication-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cytopathic_effects_on_the_host_cell" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cytopathic_effects_on_the_host_cell"> <div class="vector-toc-text"> 3.8 Cytopathic effects on the host cell </div> </a> <ul id="toc-Cytopathic_effects_on_the_host_cell-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dormant_and_latent_infections" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dormant_and_latent_infections"> <div class="vector-toc-text"> 3.9 Dormant and latent infections </div> </a> <ul id="toc-Dormant_and_latent_infections-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Host_range" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Host_range"> <div class="vector-toc-text"> 3.10 Host range </div> </a> <ul id="toc-Host_range-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Novel_viruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Novel_viruses"> <div class="vector-toc-text"> 3.11 Novel viruses </div> </a> <ul id="toc-Novel_viruses-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Classification" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Classification"> <div class="vector-toc-text"> 4 Classification </div> </a> <button aria-controls="toc-Classification-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Classification subsection </button> <ul id="toc-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"> 4.1 ICTV classification </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"> 4.2 Baltimore classification </div> </a> <ul id="toc-Baltimore_classification-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Role_in_human_disease" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Role_in_human_disease"> <div class="vector-toc-text"> 5 Role in human disease </div> </a> <button aria-controls="toc-Role_in_human_disease-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Role in human disease subsection </button> <ul id="toc-Role_in_human_disease-sublist" class="vector-toc-list"> <li id="toc-Epidemiology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Epidemiology"> <div class="vector-toc-text"> 5.1 Epidemiology </div> </a> <ul id="toc-Epidemiology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Epidemics_and_pandemics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Epidemics_and_pandemics"> <div class="vector-toc-text"> 5.2 Epidemics and pandemics </div> </a> <ul id="toc-Epidemics_and_pandemics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cancer" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cancer"> <div class="vector-toc-text"> 5.3 Cancer </div> </a> <ul id="toc-Cancer-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Host_defence_mechanisms" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Host_defence_mechanisms"> <div class="vector-toc-text"> 5.4 Host defence mechanisms </div> </a> <ul id="toc-Host_defence_mechanisms-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Prevention_and_treatment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Prevention_and_treatment"> <div class="vector-toc-text"> 5.5 Prevention and treatment </div> </a> <ul id="toc-Prevention_and_treatment-sublist" class="vector-toc-list"> <li id="toc-Vaccines" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Vaccines"> <div class="vector-toc-text"> 5.5.1 Vaccines </div> </a> <ul id="toc-Vaccines-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Antiviral_drugs" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Antiviral_drugs"> <div class="vector-toc-text"> 5.5.2 Antiviral drugs </div> </a> <ul id="toc-Antiviral_drugs-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Infection_in_other_species" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Infection_in_other_species"> <div class="vector-toc-text"> 6 Infection in other species </div> </a> <button aria-controls="toc-Infection_in_other_species-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Infection in other species subsection </button> <ul id="toc-Infection_in_other_species-sublist" class="vector-toc-list"> <li id="toc-Animal_viruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Animal_viruses"> <div class="vector-toc-text"> 6.1 Animal viruses </div> </a> <ul id="toc-Animal_viruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Plant_viruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Plant_viruses"> <div class="vector-toc-text"> 6.2 Plant viruses </div> </a> <ul id="toc-Plant_viruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Bacterial_viruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bacterial_viruses"> <div class="vector-toc-text"> 6.3 Bacterial viruses </div> </a> <ul id="toc-Bacterial_viruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Archaeal_viruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Archaeal_viruses"> <div class="vector-toc-text"> 6.4 Archaeal viruses </div> </a> <ul id="toc-Archaeal_viruses-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Role_in_aquatic_ecosystems" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Role_in_aquatic_ecosystems"> <div class="vector-toc-text"> 7 Role in aquatic ecosystems </div> </a> <ul id="toc-Role_in_aquatic_ecosystems-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Role_in_evolution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Role_in_evolution"> <div class="vector-toc-text"> 8 Role in evolution </div> </a> <ul id="toc-Role_in_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Applications" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Applications"> <div class="vector-toc-text"> 9 Applications </div> </a> <button aria-controls="toc-Applications-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Applications subsection </button> <ul id="toc-Applications-sublist" class="vector-toc-list"> <li id="toc-Life_sciences_and_medicine" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Life_sciences_and_medicine"> <div class="vector-toc-text"> 9.1 Life sciences and medicine </div> </a> <ul id="toc-Life_sciences_and_medicine-sublist" class="vector-toc-list"> <li id="toc-Virotherapy" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Virotherapy"> <div class="vector-toc-text"> 9.1.1 Virotherapy </div> </a> <ul id="toc-Virotherapy-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Materials_science_and_nanotechnology" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Materials_science_and_nanotechnology"> <div class="vector-toc-text"> 9.2 Materials science and nanotechnology </div> </a> <ul id="toc-Materials_science_and_nanotechnology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Synthetic_viruses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Synthetic_viruses"> <div class="vector-toc-text"> 9.3 Synthetic viruses </div> </a> <ul id="toc-Synthetic_viruses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Weapons" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Weapons"> <div class="vector-toc-text"> 9.4 Weapons </div> </a> <ul id="toc-Weapons-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> 10 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 11 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 12 External links </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <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" > Toggle the table of contents </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">Virus</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 172 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-172" aria-hidden="true" > 172 languages </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Virus" title="Virus – Afrikaans" lang="af" hreflang="af" data-title="Virus" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target">Afrikaans</a></li><li class="interlanguage-link interwiki-als mw-list-item"><a href="https://als.wikipedia.org/wiki/Virus_(Medizin)" title="Virus (Medizin) – Alemannic" lang="gsw" hreflang="gsw" data-title="Virus (Medizin)" data-language-autonym="Alemannisch" data-language-local-name="Alemannic" class="interlanguage-link-target">Alemannisch</a></li><li class="interlanguage-link interwiki-am mw-list-item"><a href="https://am.wikipedia.org/wiki/%E1%89%AB%E1%8B%AD%E1%88%A8%E1%88%B5" title="ቫይረስ – Amharic" lang="am" hreflang="am" data-title="ቫይረስ" data-language-autonym="አማርኛ" data-language-local-name="Amharic" class="interlanguage-link-target">አማርኛ</a></li><li class="interlanguage-link interwiki-ang mw-list-item"><a href="https://ang.wikipedia.org/wiki/Cl%C4%93ofanwyrm" title="Clēofanwyrm – Old English" lang="ang" hreflang="ang" data-title="Clēofanwyrm" data-language-autonym="Ænglisc" data-language-local-name="Old English" class="interlanguage-link-target">Ænglisc</a></li><li class="interlanguage-link interwiki-ar badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ar.wikipedia.org/wiki/%D9%81%D9%8A%D8%B1%D9%88%D8%B3" title="فيروس – Arabic" lang="ar" hreflang="ar" data-title="فيروس" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-an mw-list-item"><a href="https://an.wikipedia.org/wiki/Virus" title="Virus – Aragonese" lang="an" hreflang="an" data-title="Virus" data-language-autonym="Aragonés" data-language-local-name="Aragonese" class="interlanguage-link-target">Aragonés</a></li><li class="interlanguage-link interwiki-arc mw-list-item"><a href="https://arc.wikipedia.org/wiki/%DC%92%DC%9D%DC%AA%DC%98%DC%A3" title="ܒܝܪܘܣ – Aramaic" lang="arc" hreflang="arc" data-title="ܒܝܪܘܣ" data-language-autonym="ܐܪܡܝܐ" data-language-local-name="Aramaic" class="interlanguage-link-target">ܐܪܡܝܐ</a></li><li class="interlanguage-link interwiki-hyw mw-list-item"><a href="https://hyw.wikipedia.org/wiki/%D4%BA%D5%A1%D5%B0%D6%80" title="Ժահր – Western Armenian" lang="hyw" hreflang="hyw" data-title="Ժահր" data-language-autonym="Արեւմտահայերէն" data-language-local-name="Western Armenian" class="interlanguage-link-target">Արեւմտահայերէն</a></li><li class="interlanguage-link interwiki-as mw-list-item"><a href="https://as.wikipedia.org/wiki/%E0%A6%AD%E0%A6%BE%E0%A6%87%E0%A7%B0%E0%A6%BE%E0%A6%9B" title="ভাইৰাছ – Assamese" lang="as" hreflang="as" data-title="ভাইৰাছ" data-language-autonym="অসমীয়া" data-language-local-name="Assamese" class="interlanguage-link-target">অসমীয়া</a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Virus" title="Virus – Asturian" lang="ast" hreflang="ast" data-title="Virus" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target">Asturianu</a></li><li class="interlanguage-link interwiki-gn mw-list-item"><a href="https://gn.wikipedia.org/wiki/V%C3%ADru" title="Víru – Guarani" lang="gn" hreflang="gn" data-title="Víru" data-language-autonym="Avañe'ẽ" data-language-local-name="Guarani" class="interlanguage-link-target">Avañe'ẽ</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Viruslar" title="Viruslar – Azerbaijani" lang="az" hreflang="az" data-title="Viruslar" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-azb mw-list-item"><a href="https://azb.wikipedia.org/wiki/%D9%88%DB%8C%D8%B1%D9%88%D8%B3" title="ویروس – South Azerbaijani" lang="azb" hreflang="azb" data-title="ویروس" data-language-autonym="تۆرکجه" data-language-local-name="South Azerbaijani" class="interlanguage-link-target">تۆرکجه</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" title="ভাইরাস – Bangla" lang="bn" hreflang="bn" data-title="ভাইরাস" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/P%C4%93%E2%81%BF-to%CC%8Dk" title="Pēⁿ-to̍k – Minnan" lang="nan" hreflang="nan" data-title="Pēⁿ-to̍k" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</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%D1%82%D0%B0%D1%80" title="Вирустар – Bashkir" lang="ba" hreflang="ba" data-title="Вирустар" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target">Башҡортса</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%D1%8B" title="Вірусы – Belarusian" lang="be" hreflang="be" data-title="Вірусы" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target">Беларуская</a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%92%D1%96%D1%80%D1%83%D1%81" title="Вірус – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Вірус" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target">Беларуская (тарашкевіца)</a></li><li class="interlanguage-link interwiki-bh mw-list-item"><a href="https://bh.wikipedia.org/wiki/%E0%A4%B5%E0%A4%BE%E0%A4%AF%E0%A4%B0%E0%A4%B8" title="वायरस – Bhojpuri" lang="bh" hreflang="bh" data-title="वायरस" data-language-autonym="भोजपुरी" data-language-local-name="Bhojpuri" class="interlanguage-link-target">भोजपुरी</a></li><li class="interlanguage-link interwiki-bcl mw-list-item"><a href="https://bcl.wikipedia.org/wiki/Birus" title="Birus – Central Bikol" lang="bcl" hreflang="bcl" data-title="Birus" data-language-autonym="Bikol Central" data-language-local-name="Central Bikol" class="interlanguage-link-target">Bikol Central</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%B8" title="Вируси – Bulgarian" lang="bg" hreflang="bg" data-title="Вируси" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-bo mw-list-item"><a href="https://bo.wikipedia.org/wiki/%E0%BD%93%E0%BD%91%E0%BC%8B%E0%BD%91%E0%BD%B4%E0%BD%82%E0%BC%8B" title="ནད་དུག་ – Tibetan" lang="bo" hreflang="bo" data-title="ནད་དུག་" data-language-autonym="བོད་ཡིག" data-language-local-name="Tibetan" class="interlanguage-link-target">བོད་ཡིག</a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Virus_(biologija)" title="Virus (biologija) – Bosnian" lang="bs" hreflang="bs" data-title="Virus (biologija)" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-br mw-list-item"><a href="https://br.wikipedia.org/wiki/Viruz" title="Viruz – Breton" lang="br" hreflang="br" data-title="Viruz" data-language-autonym="Brezhoneg" data-language-local-name="Breton" class="interlanguage-link-target">Brezhoneg</a></li><li class="interlanguage-link interwiki-bxr mw-list-item"><a href="https://bxr.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81" title="Вирус – Russia Buriat" lang="bxr" hreflang="bxr" data-title="Вирус" data-language-autonym="Буряад" data-language-local-name="Russia Buriat" class="interlanguage-link-target">Буряад</a></li><li class="interlanguage-link interwiki-ca badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ca.wikipedia.org/wiki/Virus" title="Virus – Catalan" lang="ca" hreflang="ca" data-title="Virus" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</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" title="Вирус – Chuvash" lang="cv" hreflang="cv" data-title="Вирус" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target">Чӑвашла</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Virus" title="Virus – Czech" lang="cs" hreflang="cs" data-title="Virus" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Firws" title="Firws – Welsh" lang="cy" hreflang="cy" data-title="Firws" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target">Cymraeg</a></li><li class="interlanguage-link interwiki-dag mw-list-item"><a href="https://dag.wikipedia.org/wiki/Vaaros" title="Vaaros – Dagbani" lang="dag" hreflang="dag" data-title="Vaaros" data-language-autonym="Dagbanli" data-language-local-name="Dagbani" class="interlanguage-link-target">Dagbanli</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Virus" title="Virus – Danish" lang="da" hreflang="da" data-title="Virus" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Viren" title="Viren – German" lang="de" hreflang="de" data-title="Viren" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-dv mw-list-item"><a href="https://dv.wikipedia.org/wiki/%DE%88%DE%A6%DE%87%DE%A8%DE%83%DE%A6%DE%90%DE%B0" title="ވައިރަސް – Divehi" lang="dv" hreflang="dv" data-title="ވައިރަސް" data-language-autonym="ދިވެހިބަސް" data-language-local-name="Divehi" class="interlanguage-link-target">ދިވެހިބަސް</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Viirused" title="Viirused – Estonian" lang="et" hreflang="et" data-title="Viirused" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%99%CF%8C%CF%82" title="Ιός – Greek" lang="el" hreflang="el" data-title="Ιός" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-myv mw-list-item"><a href="https://myv.wikipedia.org/wiki/%D0%9A%D0%B5%D0%B7%D0%BE%D0%B2%D0%BA%D1%88%D1%82" title="Кезовкшт – Erzya" lang="myv" hreflang="myv" data-title="Кезовкшт" data-language-autonym="Эрзянь" data-language-local-name="Erzya" class="interlanguage-link-target">Эрзянь</a></li><li class="interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://es.wikipedia.org/wiki/Virus" title="Virus – Spanish" lang="es" hreflang="es" data-title="Virus" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Viruso_(biologio)" title="Viruso (biologio) – Esperanto" lang="eo" hreflang="eo" data-title="Viruso (biologio)" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-ext mw-list-item"><a href="https://ext.wikipedia.org/wiki/Virus" title="Virus – Extremaduran" lang="ext" hreflang="ext" data-title="Virus" data-language-autonym="Estremeñu" data-language-local-name="Extremaduran" class="interlanguage-link-target">Estremeñu</a></li><li class="interlanguage-link interwiki-eu badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://eu.wikipedia.org/wiki/Birus" title="Birus – Basque" lang="eu" hreflang="eu" data-title="Birus" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%88%DB%8C%D8%B1%D9%88%D8%B3" title="ویروس – Persian" lang="fa" hreflang="fa" data-title="ویروس" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-hif mw-list-item"><a href="https://hif.wikipedia.org/wiki/Virus" title="Virus – Fiji Hindi" lang="hif" hreflang="hif" data-title="Virus" data-language-autonym="Fiji Hindi" data-language-local-name="Fiji Hindi" class="interlanguage-link-target">Fiji Hindi</a></li><li class="interlanguage-link interwiki-fo mw-list-item"><a href="https://fo.wikipedia.org/wiki/Virus" title="Virus – Faroese" lang="fo" hreflang="fo" data-title="Virus" data-language-autonym="Føroyskt" data-language-local-name="Faroese" class="interlanguage-link-target">Føroyskt</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Virus" title="Virus – French" lang="fr" hreflang="fr" data-title="Virus" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-fy mw-list-item"><a href="https://fy.wikipedia.org/wiki/Firus_(biology)" title="Firus (biology) – Western Frisian" lang="fy" hreflang="fy" data-title="Firus (biology)" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target">Frysk</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/V%C3%ADreas" title="Víreas – Irish" lang="ga" hreflang="ga" data-title="Víreas" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gv mw-list-item"><a href="https://gv.wikipedia.org/wiki/Veerys" title="Veerys – Manx" lang="gv" hreflang="gv" data-title="Veerys" data-language-autonym="Gaelg" data-language-local-name="Manx" class="interlanguage-link-target">Gaelg</a></li><li class="interlanguage-link interwiki-gd mw-list-item"><a href="https://gd.wikipedia.org/wiki/B%C3%ACoras" title="Bìoras – Scottish Gaelic" lang="gd" hreflang="gd" data-title="Bìoras" data-language-autonym="Gàidhlig" data-language-local-name="Scottish Gaelic" class="interlanguage-link-target">Gàidhlig</a></li><li class="interlanguage-link interwiki-gl badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://gl.wikipedia.org/wiki/Virus" title="Virus – Galician" lang="gl" hreflang="gl" data-title="Virus" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-inh mw-list-item"><a href="https://inh.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D0%B0%D1%88" title="Вирусаш – Ingush" lang="inh" hreflang="inh" data-title="Вирусаш" data-language-autonym="ГӀалгӀай" data-language-local-name="Ingush" class="interlanguage-link-target">ГӀалгӀай</a></li><li class="interlanguage-link interwiki-ki mw-list-item"><a href="https://ki.wikipedia.org/wiki/Virus" title="Virus – Kikuyu" lang="ki" hreflang="ki" data-title="Virus" data-language-autonym="Gĩkũyũ" data-language-local-name="Kikuyu" class="interlanguage-link-target">Gĩkũyũ</a></li><li class="interlanguage-link interwiki-gu mw-list-item"><a href="https://gu.wikipedia.org/wiki/%E0%AA%B5%E0%AA%BF%E0%AA%B7%E0%AA%BE%E0%AA%A3%E0%AB%81" title="વિષાણુ – Gujarati" lang="gu" hreflang="gu" data-title="વિષાણુ" data-language-autonym="ગુજરાતી" data-language-local-name="Gujarati" class="interlanguage-link-target">ગુજરાતી</a></li><li class="interlanguage-link interwiki-hak mw-list-item"><a href="https://hak.wikipedia.org/wiki/Phiang-thu%CC%8Dk" title="Phiang-thu̍k – Hakka Chinese" lang="hak" hreflang="hak" data-title="Phiang-thu̍k" data-language-autonym="客家語 / Hak-kâ-ngî" data-language-local-name="Hakka Chinese" class="interlanguage-link-target">客家語 / Hak-kâ-ngî</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" title="바이러스 – Korean" lang="ko" hreflang="ko" data-title="바이러스" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://hy.wikipedia.org/wiki/%D5%8E%D5%AB%D6%80%D5%B8%D6%82%D5%BD%D5%B6%D5%A5%D6%80" title="Վիրուսներ – Armenian" lang="hy" hreflang="hy" data-title="Վիրուսներ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</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" title="विषाणु – Hindi" lang="hi" hreflang="hi" data-title="विषाणु" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Virusi" title="Virusi – Croatian" lang="hr" hreflang="hr" data-title="Virusi" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-gor mw-list-item"><a href="https://gor.wikipedia.org/wiki/Virus" title="Virus – Gorontalo" lang="gor" hreflang="gor" data-title="Virus" data-language-autonym="Bahasa Hulontalo" data-language-local-name="Gorontalo" class="interlanguage-link-target">Bahasa Hulontalo</a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Viruso" title="Viruso – Ido" lang="io" hreflang="io" data-title="Viruso" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target">Ido</a></li><li class="interlanguage-link interwiki-ilo mw-list-item"><a href="https://ilo.wikipedia.org/wiki/Birus" title="Birus – Iloko" lang="ilo" hreflang="ilo" data-title="Birus" data-language-autonym="Ilokano" data-language-local-name="Iloko" class="interlanguage-link-target">Ilokano</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Virus" title="Virus – Indonesian" lang="id" hreflang="id" data-title="Virus" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-ia mw-list-item"><a href="https://ia.wikipedia.org/wiki/Virus" title="Virus – Interlingua" lang="ia" hreflang="ia" data-title="Virus" data-language-autonym="Interlingua" data-language-local-name="Interlingua" class="interlanguage-link-target">Interlingua</a></li><li class="interlanguage-link interwiki-ie mw-list-item"><a href="https://ie.wikipedia.org/wiki/Virus" title="Virus – Interlingue" lang="ie" hreflang="ie" data-title="Virus" data-language-autonym="Interlingue" data-language-local-name="Interlingue" class="interlanguage-link-target">Interlingue</a></li><li class="interlanguage-link interwiki-zu mw-list-item"><a href="https://zu.wikipedia.org/wiki/IGciwanyane" title="IGciwanyane – Zulu" lang="zu" hreflang="zu" data-title="IGciwanyane" data-language-autonym="IsiZulu" data-language-local-name="Zulu" class="interlanguage-link-target">IsiZulu</a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Veira" title="Veira – Icelandic" lang="is" hreflang="is" data-title="Veira" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target">Íslenska</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Virus_(biologia)" title="Virus (biologia) – Italian" lang="it" hreflang="it" data-title="Virus (biologia)" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A0%D7%92%D7%99%D7%A3" title="נגיף – Hebrew" lang="he" hreflang="he" data-title="נגיף" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Virus" title="Virus – Javanese" lang="jv" hreflang="jv" data-title="Virus" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target">Jawa</a></li><li class="interlanguage-link interwiki-kbp mw-list-item"><a href="https://kbp.wikipedia.org/wiki/K%CA%8Ad%C9%94%C5%8B_s%C9%94t%CA%8A" title="Kʊdɔŋ sɔtʊ – Kabiye" lang="kbp" hreflang="kbp" data-title="Kʊdɔŋ sɔtʊ" data-language-autonym="Kabɩyɛ" data-language-local-name="Kabiye" class="interlanguage-link-target">Kabɩyɛ</a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%B5%E0%B3%88%E0%B2%B0%E0%B2%BE%E0%B2%A3%E0%B3%81" title="ವೈರಾಣು – Kannada" lang="kn" hreflang="kn" data-title="ವೈರಾಣು" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target">ಕನ್ನಡ</a></li><li class="interlanguage-link interwiki-pam mw-list-item"><a href="https://pam.wikipedia.org/wiki/Virus" title="Virus – Pampanga" lang="pam" hreflang="pam" data-title="Virus" data-language-autonym="Kapampangan" data-language-local-name="Pampanga" class="interlanguage-link-target">Kapampangan</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%94%E1%83%91%E1%83%98" title="ვირუსები – Georgian" lang="ka" hreflang="ka" data-title="ვირუსები" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target">ქართული</a></li><li class="interlanguage-link interwiki-csb mw-list-item"><a href="https://csb.wikipedia.org/wiki/Wirus" title="Wirus – Kashubian" lang="csb" hreflang="csb" data-title="Wirus" data-language-autonym="Kaszëbsczi" data-language-local-name="Kashubian" class="interlanguage-link-target">Kaszëbsczi</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%D1%82%D0%B0%D1%80" title="Вирустар – Kazakh" lang="kk" hreflang="kk" data-title="Вирустар" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-kw mw-list-item"><a href="https://kw.wikipedia.org/wiki/Virus" title="Virus – Cornish" lang="kw" hreflang="kw" data-title="Virus" data-language-autonym="Kernowek" data-language-local-name="Cornish" class="interlanguage-link-target">Kernowek</a></li><li class="interlanguage-link interwiki-rn mw-list-item"><a href="https://rn.wikipedia.org/wiki/Virus" title="Virus – Rundi" lang="rn" hreflang="rn" data-title="Virus" data-language-autonym="Ikirundi" data-language-local-name="Rundi" class="interlanguage-link-target">Ikirundi</a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/Virusi" title="Virusi – Swahili" lang="sw" hreflang="sw" data-title="Virusi" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target">Kiswahili</a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Viris" title="Viris – Haitian Creole" lang="ht" hreflang="ht" data-title="Viris" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target">Kreyòl ayisyen</a></li><li class="interlanguage-link interwiki-gcr mw-list-item"><a href="https://gcr.wikipedia.org/wiki/Viris" title="Viris – Guianan Creole" lang="gcr" hreflang="gcr" data-title="Viris" data-language-autonym="Kriyòl gwiyannen" data-language-local-name="Guianan Creole" class="interlanguage-link-target">Kriyòl gwiyannen</a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/V%C3%AErus" title="Vîrus – Kurdish" lang="ku" hreflang="ku" data-title="Vîrus" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target">Kurdî</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" title="Вирус – Kyrgyz" lang="ky" hreflang="ky" data-title="Вирус" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target">Кыргызча</a></li><li class="interlanguage-link interwiki-lo mw-list-item"><a href="https://lo.wikipedia.org/wiki/%E2%80%8B%E0%BB%84%E0%BA%A7%E2%80%8B%E0%BA%A3%E0%BA%B1%E2%80%8B%E0%BA%AA" title="ໄວຣັສ – Lao" lang="lo" hreflang="lo" data-title="ໄວຣັສ" data-language-autonym="ລາວ" data-language-local-name="Lao" class="interlanguage-link-target">ລາວ</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Virus_biologicum" title="Virus biologicum – Latin" lang="la" hreflang="la" data-title="Virus biologicum" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/V%C4%ABruss" title="Vīruss – Latvian" lang="lv" hreflang="lv" data-title="Vīruss" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-lb mw-list-item"><a href="https://lb.wikipedia.org/wiki/Virus" title="Virus – Luxembourgish" lang="lb" hreflang="lb" data-title="Virus" data-language-autonym="Lëtzebuergesch" data-language-local-name="Luxembourgish" class="interlanguage-link-target">Lëtzebuergesch</a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Virusas" title="Virusas – Lithuanian" lang="lt" hreflang="lt" data-title="Virusas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-lij mw-list-item"><a href="https://lij.wikipedia.org/wiki/Virus" title="Virus – Ligurian" lang="lij" hreflang="lij" data-title="Virus" data-language-autonym="Ligure" data-language-local-name="Ligurian" class="interlanguage-link-target">Ligure</a></li><li class="interlanguage-link interwiki-li mw-list-item"><a href="https://li.wikipedia.org/wiki/Virusse" title="Virusse – Limburgish" lang="li" hreflang="li" data-title="Virusse" data-language-autonym="Limburgs" data-language-local-name="Limburgish" class="interlanguage-link-target">Limburgs</a></li><li class="interlanguage-link interwiki-ln mw-list-item"><a href="https://ln.wikipedia.org/wiki/Vil%C3%BAsu" title="Vilúsu – Lingala" lang="ln" hreflang="ln" data-title="Vilúsu" data-language-autonym="Lingála" data-language-local-name="Lingala" class="interlanguage-link-target">Lingála</a></li><li class="interlanguage-link interwiki-lfn mw-list-item"><a href="https://lfn.wikipedia.org/wiki/Virus" title="Virus – Lingua Franca Nova" lang="lfn" hreflang="lfn" data-title="Virus" data-language-autonym="Lingua Franca Nova" data-language-local-name="Lingua Franca Nova" class="interlanguage-link-target">Lingua Franca Nova</a></li><li class="interlanguage-link interwiki-lg mw-list-item"><a href="https://lg.wikipedia.org/wiki/Vayiraasi" title="Vayiraasi – Ganda" lang="lg" hreflang="lg" data-title="Vayiraasi" data-language-autonym="Luganda" data-language-local-name="Ganda" class="interlanguage-link-target">Luganda</a></li><li class="interlanguage-link interwiki-lmo mw-list-item"><a href="https://lmo.wikipedia.org/wiki/Virus_(biolojia)" title="Virus (biolojia) – Lombard" lang="lmo" hreflang="lmo" data-title="Virus (biolojia)" data-language-autonym="Lombard" data-language-local-name="Lombard" class="interlanguage-link-target">Lombard</a></li><li class="interlanguage-link interwiki-hu badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://hu.wikipedia.org/wiki/V%C3%ADrus" title="Vírus – Hungarian" lang="hu" hreflang="hu" data-title="Vírus" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://mk.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81" title="Вирус – Macedonian" lang="mk" hreflang="mk" data-title="Вирус" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-mg mw-list-item"><a href="https://mg.wikipedia.org/wiki/Virosy" title="Virosy – Malagasy" lang="mg" hreflang="mg" data-title="Virosy" data-language-autonym="Malagasy" data-language-local-name="Malagasy" class="interlanguage-link-target">Malagasy</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%B4%B8%E0%B5%8D" title="വൈറസ് – Malayalam" lang="ml" hreflang="ml" data-title="വൈറസ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%B5%E0%A4%BF%E0%A4%B7%E0%A4%BE%E0%A4%A3%E0%A5%82" title="विषाणू – Marathi" lang="mr" hreflang="mr" data-title="विषाणू" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target">मराठी</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%94%E1%83%A4%E1%83%98" title="ვირუსეფი – Mingrelian" lang="xmf" hreflang="xmf" data-title="ვირუსეფი" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target">მარგალური</a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D9%81%D9%8A%D8%B1%D9%88%D8%B3" title="فيروس – Egyptian Arabic" lang="arz" hreflang="arz" data-title="فيروس" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target">مصرى</a></li><li class="interlanguage-link interwiki-mnw mw-list-item"><a href="https://mnw.wikipedia.org/wiki/%E1%80%9D%E1%80%B3%E1%80%9B%E1%80%AF%E1%80%9E%E1%80%BA" title="ဝဳရုသ် – Mon" lang="mnw" hreflang="mnw" data-title="ဝဳရုသ်" data-language-autonym="ဘာသာမန်" data-language-local-name="Mon" class="interlanguage-link-target">ဘာသာမန်</a></li><li class="interlanguage-link interwiki-mzn mw-list-item"><a href="https://mzn.wikipedia.org/wiki/%D9%88%DB%8C%D8%B1%D9%88%D8%B3" title="ویروس – Mazanderani" lang="mzn" hreflang="mzn" data-title="ویروس" data-language-autonym="مازِرونی" data-language-local-name="Mazanderani" class="interlanguage-link-target">مازِرونی</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Virus" title="Virus – Malay" lang="ms" hreflang="ms" data-title="Virus" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-min mw-list-item"><a href="https://min.wikipedia.org/wiki/Virus" title="Virus – Minangkabau" lang="min" hreflang="min" data-title="Virus" data-language-autonym="Minangkabau" data-language-local-name="Minangkabau" class="interlanguage-link-target">Minangkabau</a></li><li class="interlanguage-link interwiki-cdo mw-list-item"><a href="https://cdo.wikipedia.org/wiki/B%C3%AAng-d%C5%ADk" title="Bêng-dŭk – Mindong" lang="cdo" hreflang="cdo" data-title="Bêng-dŭk" data-language-autonym="閩東語 / Mìng-dĕ̤ng-ngṳ̄" data-language-local-name="Mindong" class="interlanguage-link-target">閩東語 / Mìng-dĕ̤ng-ngṳ̄</a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81" title="Вирус – Mongolian" lang="mn" hreflang="mn" data-title="Вирус" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target">Монгол</a></li><li class="interlanguage-link interwiki-my mw-list-item"><a href="https://my.wikipedia.org/wiki/%E1%80%97%E1%80%AD%E1%80%AF%E1%80%84%E1%80%BA%E1%80%B8%E1%80%9B%E1%80%95%E1%80%BA%E1%80%85%E1%80%BA" title="ဗိုင်းရပ်စ် – Burmese" lang="my" hreflang="my" data-title="ဗိုင်းရပ်စ်" data-language-autonym="မြန်မာဘာသာ" data-language-local-name="Burmese" class="interlanguage-link-target">မြန်မာဘာသာ</a></li><li class="interlanguage-link interwiki-nl badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://nl.wikipedia.org/wiki/Virus_(biologie)" title="Virus (biologie) – Dutch" lang="nl" hreflang="nl" data-title="Virus (biologie)" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ne mw-list-item"><a href="https://ne.wikipedia.org/wiki/%E0%A4%B5%E0%A4%BF%E0%A4%B7%E0%A4%BE%E0%A4%A3%E0%A5%81" title="विषाणु – Nepali" lang="ne" hreflang="ne" data-title="विषाणु" data-language-autonym="नेपाली" data-language-local-name="Nepali" class="interlanguage-link-target">नेपाली</a></li><li class="interlanguage-link interwiki-new mw-list-item"><a href="https://new.wikipedia.org/wiki/%E0%A4%AD%E0%A4%BE%E0%A4%87%E0%A4%B0%E0%A4%B8" title="भाइरस – Newari" lang="new" hreflang="new" data-title="भाइरस" data-language-autonym="नेपाल भाषा" data-language-local-name="Newari" class="interlanguage-link-target">नेपाल भाषा</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" title="ウイルス – Japanese" lang="ja" hreflang="ja" data-title="ウイルス" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-frr mw-list-item"><a href="https://frr.wikipedia.org/wiki/Wiiren" title="Wiiren – Northern Frisian" lang="frr" hreflang="frr" data-title="Wiiren" data-language-autonym="Nordfriisk" data-language-local-name="Northern Frisian" class="interlanguage-link-target">Nordfriisk</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Virus" title="Virus – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Virus" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Virus" title="Virus – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Virus" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Virus" title="Virus – Occitan" lang="oc" hreflang="oc" data-title="Virus" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target">Occitan</a></li><li class="interlanguage-link interwiki-or mw-list-item"><a href="https://or.wikipedia.org/wiki/%E0%AC%AD%E0%AD%82%E0%AC%A4%E0%AC%BE%E0%AC%A3%E0%AD%81" title="ଭୂତାଣୁ – Odia" lang="or" hreflang="or" data-title="ଭୂତାଣୁ" data-language-autonym="ଓଡ଼ିଆ" data-language-local-name="Odia" class="interlanguage-link-target">ଓଡ଼ିଆ</a></li><li class="interlanguage-link interwiki-om mw-list-item"><a href="https://om.wikipedia.org/wiki/Vaayirasii" title="Vaayirasii – Oromo" lang="om" hreflang="om" data-title="Vaayirasii" data-language-autonym="Oromoo" data-language-local-name="Oromo" class="interlanguage-link-target">Oromoo</a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Viruslar" title="Viruslar – Uzbek" lang="uz" hreflang="uz" data-title="Viruslar" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target">Oʻzbekcha / ўзбекча</a></li><li class="interlanguage-link interwiki-pa mw-list-item"><a href="https://pa.wikipedia.org/wiki/%E0%A8%B5%E0%A8%BF%E0%A8%B8%E0%A8%BC%E0%A8%BE%E0%A8%A3%E0%A9%82" title="ਵਿਸ਼ਾਣੂ – Punjabi" lang="pa" hreflang="pa" data-title="ਵਿਸ਼ਾਣੂ" data-language-autonym="ਪੰਜਾਬੀ" data-language-local-name="Punjabi" class="interlanguage-link-target">ਪੰਜਾਬੀ</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%D8%B3" title="وائرس – Western Punjabi" lang="pnb" hreflang="pnb" data-title="وائرس" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target">پنجابی</a></li><li class="interlanguage-link interwiki-pap mw-list-item"><a href="https://pap.wikipedia.org/wiki/Virus" title="Virus – Papiamento" lang="pap" hreflang="pap" data-title="Virus" data-language-autonym="Papiamentu" data-language-local-name="Papiamento" class="interlanguage-link-target">Papiamentu</a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D9%88%DB%8C%D8%B1%D9%88%D8%B3" title="ویروس – Pashto" lang="ps" hreflang="ps" data-title="ویروس" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target">پښتو</a></li><li class="interlanguage-link interwiki-tpi mw-list-item"><a href="https://tpi.wikipedia.org/wiki/Virus" title="Virus – Tok Pisin" lang="tpi" hreflang="tpi" data-title="Virus" data-language-autonym="Tok Pisin" data-language-local-name="Tok Pisin" class="interlanguage-link-target">Tok Pisin</a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Viren" title="Viren – Low German" lang="nds" hreflang="nds" data-title="Viren" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target">Plattdüütsch</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Wirusy" title="Wirusy – Polish" lang="pl" hreflang="pl" data-title="Wirusy" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/V%C3%ADrus" title="Vírus – Portuguese" lang="pt" hreflang="pt" data-title="Vírus" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-kaa mw-list-item"><a href="https://kaa.wikipedia.org/wiki/Viruslar" title="Viruslar – Kara-Kalpak" lang="kaa" hreflang="kaa" data-title="Viruslar" data-language-autonym="Qaraqalpaqsha" data-language-local-name="Kara-Kalpak" class="interlanguage-link-target">Qaraqalpaqsha</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Virus" title="Virus – Romanian" lang="ro" hreflang="ro" data-title="Virus" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-qu mw-list-item"><a href="https://qu.wikipedia.org/wiki/A%C3%B1aw" title="Añaw – Quechua" lang="qu" hreflang="qu" data-title="Añaw" data-language-autonym="Runa Simi" data-language-local-name="Quechua" class="interlanguage-link-target">Runa Simi</a></li><li class="interlanguage-link interwiki-rue mw-list-item"><a href="https://rue.wikipedia.org/wiki/%D0%92%D1%96%D1%80%D1%83%D1%81" title="Вірус – Rusyn" lang="rue" hreflang="rue" data-title="Вірус" data-language-autonym="Русиньскый" data-language-local-name="Rusyn" class="interlanguage-link-target">Русиньскый</a></li><li class="interlanguage-link interwiki-ru badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ru.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81%D1%8B" title="Вирусы – Russian" lang="ru" hreflang="ru" data-title="Вирусы" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</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" title="Вирус – Yakut" lang="sah" hreflang="sah" data-title="Вирус" data-language-autonym="Саха тыла" data-language-local-name="Yakut" class="interlanguage-link-target">Саха тыла</a></li><li class="interlanguage-link interwiki-sat mw-list-item"><a href="https://sat.wikipedia.org/wiki/%E1%B1%B5%E1%B1%B7%E1%B1%9F%E1%B1%AD%E1%B1%A8%E1%B1%9F%E1%B1%A5" title="ᱵᱷᱟᱭᱨᱟᱥ – Santali" lang="sat" hreflang="sat" data-title="ᱵᱷᱟᱭᱨᱟᱥ" data-language-autonym="ᱥᱟᱱᱛᱟᱲᱤ" data-language-local-name="Santali" class="interlanguage-link-target">ᱥᱟᱱᱛᱟᱲᱤ</a></li><li class="interlanguage-link interwiki-sc mw-list-item"><a href="https://sc.wikipedia.org/wiki/Virus_(biologia)" title="Virus (biologia) – Sardinian" lang="sc" hreflang="sc" data-title="Virus (biologia)" data-language-autonym="Sardu" data-language-local-name="Sardinian" class="interlanguage-link-target">Sardu</a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/Virus" title="Virus – Scots" lang="sco" hreflang="sco" data-title="Virus" data-language-autonym="Scots" data-language-local-name="Scots" class="interlanguage-link-target">Scots</a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Virusi" title="Virusi – Albanian" lang="sq" hreflang="sq" data-title="Virusi" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target">Shqip</a></li><li class="interlanguage-link interwiki-scn mw-list-item"><a href="https://scn.wikipedia.org/wiki/Virus_(biolugg%C3%ACa)" title="Virus (bioluggìa) – Sicilian" lang="scn" hreflang="scn" data-title="Virus (bioluggìa)" data-language-autonym="Sicilianu" data-language-local-name="Sicilian" class="interlanguage-link-target">Sicilianu</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" title="වෛරස – Sinhala" lang="si" hreflang="si" data-title="වෛරස" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target">සිංහල</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Virus" title="Virus – Simple English" lang="en-simple" hreflang="en-simple" data-title="Virus" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sd mw-list-item"><a href="https://sd.wikipedia.org/wiki/%D9%88%D8%A7%D8%A6%D8%B1%D8%B3" title="وائرس – Sindhi" lang="sd" hreflang="sd" data-title="وائرس" data-language-autonym="سنڌي" data-language-local-name="Sindhi" class="interlanguage-link-target">سنڌي</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/V%C3%ADrus" title="Vírus – Slovak" lang="sk" hreflang="sk" data-title="Vírus" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Virusi" title="Virusi – Slovenian" lang="sl" hreflang="sl" data-title="Virusi" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-so mw-list-item"><a href="https://so.wikipedia.org/wiki/Fayrus" title="Fayrus – Somali" lang="so" hreflang="so" data-title="Fayrus" data-language-autonym="Soomaaliga" data-language-local-name="Somali" class="interlanguage-link-target">Soomaaliga</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%DA%A4%D8%A7%DB%8C%D8%B1%DB%86%D8%B3" title="ڤایرۆس – Central Kurdish" lang="ckb" hreflang="ckb" data-title="ڤایرۆس" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81" title="Вирус – Serbian" lang="sr" hreflang="sr" data-title="Вирус" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Virus_(biologija)" title="Virus (biologija) – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Virus (biologija)" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/Virus" title="Virus – Sundanese" lang="su" hreflang="su" data-title="Virus" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target">Sunda</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Virukset" title="Virukset – Finnish" lang="fi" hreflang="fi" data-title="Virukset" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Virus" title="Virus – Swedish" lang="sv" hreflang="sv" data-title="Virus" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Birus" title="Birus – Tagalog" lang="tl" hreflang="tl" data-title="Birus" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target">Tagalog</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%A4%E0%AF%80%E0%AE%A8%E0%AF%81%E0%AE%A3%E0%AF%8D%E0%AE%AE%E0%AE%BF" title="தீநுண்மி – Tamil" lang="ta" hreflang="ta" data-title="தீநுண்மி" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</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%BB%D0%B0%D1%80" title="Вируслар – Tatar" lang="tt" hreflang="tt" data-title="Вируслар" data-language-autonym="Татарча / tatarça" data-language-local-name="Tatar" class="interlanguage-link-target">Татарча / tatarça</a></li><li class="interlanguage-link interwiki-te badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://te.wikipedia.org/wiki/%E0%B0%B5%E0%B1%88%E0%B0%B0%E0%B0%B8%E0%B1%8D" title="వైరస్ – Telugu" lang="te" hreflang="te" data-title="వైరస్" data-language-autonym="తెలుగు" data-language-local-name="Telugu" class="interlanguage-link-target">తెలుగు</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%84%E0%B8%A7%E0%B8%A3%E0%B8%B1%E0%B8%AA" title="ไวรัส – Thai" lang="th" hreflang="th" data-title="ไวรัส" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target">ไทย</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%D2%B3%D0%BE" title="Вирусҳо – Tajik" lang="tg" hreflang="tg" data-title="Вирусҳо" data-language-autonym="Тоҷикӣ" data-language-local-name="Tajik" class="interlanguage-link-target">Тоҷикӣ</a></li><li class="interlanguage-link interwiki-tcy mw-list-item"><a href="https://tcy.wikipedia.org/wiki/%E0%B2%B5%E0%B3%88%E0%B2%B0%E0%B2%B8%E0%B3%8D" title="ವೈರಸ್ – Tulu" lang="tcy" hreflang="tcy" data-title="ವೈರಸ್" data-language-autonym="ತುಳು" data-language-local-name="Tulu" class="interlanguage-link-target">ತುಳು</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Vir%C3%BCs" title="Virüs – Turkish" lang="tr" hreflang="tr" data-title="Virüs" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-tyv mw-list-item"><a href="https://tyv.wikipedia.org/wiki/%D0%92%D0%B8%D1%80%D1%83%D1%81" title="Вирус – Tuvinian" lang="tyv" hreflang="tyv" data-title="Вирус" data-language-autonym="Тыва дыл" data-language-local-name="Tuvinian" class="interlanguage-link-target">Тыва дыл</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" title="Вірус – Ukrainian" lang="uk" hreflang="uk" data-title="Вірус" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D9%88%D8%A7%D8%A6%D8%B1%D8%B3" title="وائرس – Urdu" lang="ur" hreflang="ur" data-title="وائرس" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target">اردو</a></li><li class="interlanguage-link interwiki-ug mw-list-item"><a href="https://ug.wikipedia.org/wiki/%DB%8B%D9%89%D8%B1%DB%87%D8%B3" title="ۋىرۇس – Uyghur" lang="ug" hreflang="ug" data-title="ۋىرۇس" data-language-autonym="ئۇيغۇرچە / Uyghurche" data-language-local-name="Uyghur" class="interlanguage-link-target">ئۇيغۇرچە / Uyghurche</a></li><li class="interlanguage-link interwiki-za mw-list-item"><a href="https://za.wikipedia.org/wiki/Binghdoeg" title="Binghdoeg – Zhuang" lang="za" hreflang="za" data-title="Binghdoeg" data-language-autonym="Vahcuengh" data-language-local-name="Zhuang" class="interlanguage-link-target">Vahcuengh</a></li><li class="interlanguage-link interwiki-vec mw-list-item"><a href="https://vec.wikipedia.org/wiki/Virus_(bio%C5%82ozia)" title="Virus (biołozia) – Venetian" lang="vec" hreflang="vec" data-title="Virus (biołozia)" data-language-autonym="Vèneto" data-language-local-name="Venetian" class="interlanguage-link-target">Vèneto</a></li><li class="interlanguage-link interwiki-vi badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://vi.wikipedia.org/wiki/Virus" title="Virus – Vietnamese" lang="vi" hreflang="vi" data-title="Virus" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target">Tiếng Việt</a></li><li class="interlanguage-link interwiki-wa mw-list-item"><a href="https://wa.wikipedia.org/wiki/Vir%C3%BBsse" title="Virûsse – Walloon" lang="wa" hreflang="wa" data-title="Virûsse" data-language-autonym="Walon" data-language-local-name="Walloon" class="interlanguage-link-target">Walon</a></li><li class="interlanguage-link interwiki-zh-classical mw-list-item"><a href="https://zh-classical.wikipedia.org/wiki/%E7%97%85%E6%AF%92" title="病毒 – Literary Chinese" lang="lzh" hreflang="lzh" data-title="病毒" data-language-autonym="文言" data-language-local-name="Literary Chinese" class="interlanguage-link-target">文言</a></li><li class="interlanguage-link interwiki-war mw-list-item"><a href="https://war.wikipedia.org/wiki/Birus" title="Birus – Waray" lang="war" hreflang="war" data-title="Birus" data-language-autonym="Winaray" data-language-local-name="Waray" class="interlanguage-link-target">Winaray</a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E7%97%85%E6%AF%92" title="病毒 – Wu" lang="wuu" hreflang="wuu" data-title="病毒" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target">吴语</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%95%D7%A1" title="ווירוס – Yiddish" lang="yi" hreflang="yi" data-title="ווירוס" data-language-autonym="ייִדיש" data-language-local-name="Yiddish" class="interlanguage-link-target">ייִדיש</a></li><li class="interlanguage-link interwiki-yo mw-list-item"><a href="https://yo.wikipedia.org/wiki/%C3%88r%C3%A0n" title="Èràn – Yoruba" lang="yo" hreflang="yo" data-title="Èràn" data-language-autonym="Yorùbá" data-language-local-name="Yoruba" class="interlanguage-link-target">Yorùbá</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" title="病毒 – Cantonese" lang="yue" hreflang="yue" data-title="病毒" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target">粵語</a></li><li class="interlanguage-link interwiki-bat-smg mw-list-item"><a href="https://bat-smg.wikipedia.org/wiki/Virosos" title="Virosos – Samogitian" lang="sgs" hreflang="sgs" data-title="Virosos" data-language-autonym="Žemaitėška" data-language-local-name="Samogitian" class="interlanguage-link-target">Žemaitėška</a></li><li class="interlanguage-link interwiki-zh badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://zh.wikipedia.org/wiki/%E7%97%85%E6%AF%92" title="病毒 – Chinese" lang="zh" hreflang="zh" data-title="病毒" 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Click here for more information."><img alt="Featured article" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/20px-Cscr-featured.svg.png" decoding="async" width="20" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/30px-Cscr-featured.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/40px-Cscr-featured.svg.png 2x" data-file-width="466" data-file-height="443" /></a></div></div> <div id="mw-indicator-pp-default" class="mw-indicator"><div class="mw-parser-output"><a href="/wiki/Wikipedia:Protection_policy#semi" title="This article is semi-protected."><img alt="Page semi-protected" src="//upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/20px-Semi-protection-shackle.svg.png" decoding="async" width="20" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/30px-Semi-protection-shackle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/1/1b/Semi-protection-shackle.svg/40px-Semi-protection-shackle.svg.png 2x" data-file-width="512" data-file-height="512" /></a></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Infectious agent that replicates in cells</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about the type of pathogen. For the type of malware, see <a href="/wiki/Computer_virus" title="Computer virus">Computer virus</a>. For other uses, see <a href="/wiki/Virus_(disambiguation)" class="mw-disambig" title="Virus (disambiguation)">Virus (disambiguation)</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">For a more accessible and less technical introduction to this topic, see <a href="/wiki/Introduction_to_viruses" title="Introduction to viruses">Introduction to viruses</a>.</div> <table class="infobox biota" style="text-align: left; width: 200px; font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(250,250,190)">Virus </th></tr> <tr> <td colspan="2" style="text-align: center"><a href="/wiki/File:Coronavirus._SARS-CoV-2.png" class="mw-file-description"><img alt=""Severe acute respiratory syndrome coronavirus 2"" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/94/Coronavirus._SARS-CoV-2.png/220px-Coronavirus._SARS-CoV-2.png" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/94/Coronavirus._SARS-CoV-2.png/330px-Coronavirus._SARS-CoV-2.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/94/Coronavirus._SARS-CoV-2.png/440px-Coronavirus._SARS-CoV-2.png 2x" data-file-width="2048" data-file-height="2048" /></a> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%"><a href="/wiki/SARS-CoV-2" title="SARS-CoV-2">SARS-CoV-2</a>, a member of the subfamily <a href="/wiki/Coronavirinae" class="mw-redirect" title="Coronavirinae">Coronavirinae</a> </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(250,250,190)"><a href="/wiki/Virus_classification" title="Virus classification">Virus classification</a> <a href="/wiki/Template:Taxonomy/Virus" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a> </th></tr> <tr> <td>(unranked): </td> <td>Virus </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(250,250,190)"><a href="/wiki/Realm_(virology)" title="Realm (virology)">Realms</a> </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul><li><a href="/wiki/Adnaviria" title="Adnaviria">Adnaviria</a></li> <li><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></li> <li><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></li> <li><a href="/wiki/Riboviria" title="Riboviria">Riboviria</a></li> <li><a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a></li> <li><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></li></ul> </td></tr> </tbody></table><style data-mw-deduplicate="TemplateStyles:r1238732961">@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .infobox.biota img{background:transparent}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .infobox.biota img{background:white}}.mw-parser-output .infobox.biota .taxobox-edit-taxonomy img{background:transparent!important}body.skin-vector .mw-parser-output table.biota.infobox{margin-top:0.5em}body.skin--responsive .mw-parser-output table.biota.infobox tr.taxonrow td{padding:2px 10px}</style> A virus is a submicroscopic <a href="/wiki/Infectious_agent" class="mw-redirect" title="Infectious agent">infectious agent</a> that replicates only inside the living <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a> of an <a href="/wiki/Organism" title="Organism">organism</a>.<a href="#cite_note-WU_2020-1">[1]</a> Viruses infect all <a href="/wiki/Life_forms" class="mw-redirect" title="Life forms">life forms</a>, from animals and plants to <a href="/wiki/Microorganism" title="Microorganism">microorganisms</a>, including <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> and <a href="/wiki/Archaea" title="Archaea">archaea</a>.<a href="#cite_note-Koonin_2006-2">[2]</a><a href="#cite_note-Zimmer_2021-3">[3]</a> Viruses are found in almost every <a href="/wiki/Ecosystem" title="Ecosystem">ecosystem</a> on Earth and are the most numerous type of biological entity.<a href="#cite_note-Lawrence_2009-4">[4]</a><a href="#cite_note-5">[5]</a> Since <a href="/wiki/Dmitri_Ivanovsky" title="Dmitri Ivanovsky">Dmitri Ivanovsky</a>'s 1892 article describing a non-bacterial <a href="/wiki/Pathogen" title="Pathogen">pathogen</a> infecting tobacco plants and the discovery of the <a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">tobacco mosaic virus</a> by <a href="/wiki/Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a> in 1898,<a href="#cite_note-Dimmock_2007-6">[6]</a>: 4  more than 11,000 of the millions of <a href="/wiki/List_of_virus_species" title="List of virus species">virus species</a> have been described in detail.<a href="#cite_note-ictv_2022-7">[7]</a><a href="#cite_note-Breitbart_2005-8">[8]</a> The study of viruses is known as <a href="/wiki/Virology" title="Virology">virology</a>, a subspeciality of <a href="/wiki/Microbiology" title="Microbiology">microbiology</a>. When infected, a <a href="/wiki/Host_cell" class="mw-redirect" title="Host cell">host cell</a> is often forced to rapidly produce thousands of copies of the original virus. When not inside an infected cell or in the process of infecting a cell, viruses exist in the form of independent viral particles, or virions, consisting of (i) <a href="/wiki/Genetic_material" class="mw-redirect" title="Genetic material">genetic material</a>, i.e., long <a href="/wiki/Molecule" title="Molecule">molecules</a> of <a href="/wiki/DNA" title="DNA">DNA</a> or <a href="/wiki/RNA" title="RNA">RNA</a> that encode the structure of the proteins by which the virus acts; (ii) a <a href="/wiki/Protein" title="Protein">protein</a> coat, the <a href="/wiki/Capsid" title="Capsid">capsid</a>, which surrounds and protects the genetic material; and in some cases (iii) an outside <a href="/wiki/Viral_envelope" title="Viral envelope">envelope</a> of <a href="/wiki/Lipid" title="Lipid">lipids</a>. The shapes of these virus particles range from simple <a href="/wiki/Helix" title="Helix">helical</a> and <a href="/wiki/Icosahedron" title="Icosahedron">icosahedral</a> forms to more complex structures. Most virus species have virions too small to be seen with an <a href="/wiki/Optical_microscope" title="Optical microscope">optical microscope</a> and are one-hundredth the size of most bacteria. The origins of viruses in the <a href="/wiki/Evolutionary_history_of_life" class="mw-redirect" title="Evolutionary history of life">evolutionary history of life</a> are still unclear. Some viruses may have evolved from <a href="/wiki/Plasmid" title="Plasmid">plasmids</a>, which are pieces of DNA that can move between cells. Other viruses may have evolved from bacteria. In evolution, viruses are an important means of <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>, which increases <a href="/wiki/Genetic_diversity" title="Genetic diversity">genetic diversity</a> in a way analogous to <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual reproduction</a>.<a href="#cite_note-Canchaya_2003-9">[9]</a> Viruses are considered by some <a href="/wiki/Biologist" title="Biologist">biologists</a> to be a life form, because they carry genetic material, reproduce, and evolve through <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>, although they lack some key characteristics, such as cell structure, that are generally considered necessary criteria for defining life. Because they possess some but not all such qualities, viruses have been described as "organisms at the edge of life"<a href="#cite_note-Rybicki_1990-10">[10]</a> and as <a href="/wiki/Viral_replication" title="Viral replication">replicators</a>.<a href="#cite_note-Koonin_2016-11">[11]</a> <a href="/wiki/Pathogen_transmission" title="Pathogen transmission">Viruses spread</a> in many ways. One transmission pathway is through disease-bearing organisms known as <a href="/wiki/Vector_(epidemiology)" class="mw-redirect" title="Vector (epidemiology)">vectors</a>: for example, viruses are often transmitted from plant to plant by insects that feed on <a href="/wiki/Plant_sap" class="mw-redirect" title="Plant sap">plant sap</a>, such as <a href="/wiki/Aphid" title="Aphid">aphids</a>; and viruses in animals can be carried by <a href="/wiki/Hematophagy" title="Hematophagy">blood-sucking</a> insects. Many viruses spread <a href="/wiki/Airborne_transmission" title="Airborne transmission">in the air</a> by coughing and sneezing, including <a href="/wiki/Influenza" title="Influenza">influenza viruses</a>, <a href="/wiki/SARS-CoV-2" title="SARS-CoV-2">SARS-CoV-2</a>, <a href="/wiki/Chickenpox" title="Chickenpox">chickenpox</a>, <a href="/wiki/Smallpox" title="Smallpox">smallpox</a>, and <a href="/wiki/Measles" title="Measles">measles</a>. <a href="/wiki/Norovirus" title="Norovirus">Norovirus</a> and <a href="/wiki/Rotavirus" title="Rotavirus">rotavirus</a>, common causes of viral <a href="/wiki/Gastroenteritis" title="Gastroenteritis">gastroenteritis</a>, are transmitted by the <a href="/wiki/Fecal%E2%80%93oral_route" title="Fecal–oral route">faecal–oral route</a>, passed by hand-to-mouth contact or in food or water. The <a href="/wiki/Infectious_dose" class="mw-redirect" title="Infectious dose">infectious dose</a> of norovirus required to produce infection in humans is fewer than 100 particles.<a href="#cite_note-Robilotti_2015-12">[12]</a> <a href="/wiki/HIV" title="HIV">HIV</a> is one of several viruses <a href="/wiki/Sexually_transmitted_infection" title="Sexually transmitted infection">transmitted through sexual contact</a> and by exposure to infected blood. The variety of host cells that a virus can infect is called its <a href="/wiki/Host_specificity" class="mw-redirect" title="Host specificity">host range</a>: this is narrow for viruses specialized to infect only a few species, or broad for viruses capable of infecting many.<a href="#cite_note-Shors_2017-13">[13]</a>: 123–124  Viral infections in animals provoke an <a href="/wiki/Immune_response" title="Immune response">immune response</a> that usually eliminates the infecting virus. Immune responses can also be produced by <a href="/wiki/Vaccine" title="Vaccine">vaccines</a>, which confer an <a href="/wiki/Immunity_(medical)" class="mw-redirect" title="Immunity (medical)">artificially acquired immunity</a> to the specific viral infection. Some viruses, including those that cause <a href="/wiki/HIV/AIDS" title="HIV/AIDS">HIV/AIDS</a>, <a href="/wiki/Human_papillomavirus_infection" title="Human papillomavirus infection">HPV infection</a>, and <a href="/wiki/Viral_hepatitis" title="Viral hepatitis">viral hepatitis</a>, evade these immune responses and result in <a href="/wiki/Chronic_(medical)" class="mw-redirect" title="Chronic (medical)">chronic</a> infections. Several classes of <a href="/wiki/Antiviral_drug" title="Antiviral drug">antiviral drugs</a> have been developed. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Etymology">Etymology</h2></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Plural_form_of_words_ending_in_-us" title="Plural form of words ending in -us">Plural form of words ending in -us</a></div> The English word "virus" comes from the <a href="/wiki/Latin" title="Latin">Latin</a> vīrus, which refers to <a href="/wiki/Poison" title="Poison">poison</a> and other noxious liquids. Vīrus comes from the same <a href="/wiki/Indo-European_languages" title="Indo-European languages">Indo-European</a> root as <a href="/wiki/Sanskrit" title="Sanskrit">Sanskrit</a> viṣa, <a href="/wiki/Avestan" title="Avestan">Avestan</a> vīša, and <a href="/wiki/Ancient_Greek" title="Ancient Greek">Ancient Greek</a> ἰός (iós), which all mean "poison". The first attested use of "virus" in English appeared in 1398 in <a href="/wiki/John_Trevisa" title="John Trevisa">John Trevisa</a>'s translation of <a href="/wiki/Bartholomeus_Anglicus" class="mw-redirect" title="Bartholomeus Anglicus">Bartholomeus Anglicus</a>'s De Proprietatibus Rerum.<a href="#cite_note-14">[14]</a><a href="#cite_note-Etymology_Dictionary-15">[15]</a> Virulent, from Latin virulentus ('poisonous'), dates to <abbr title="circa">c.</abbr> 1400.<a href="#cite_note-16">[16]</a><a href="#cite_note-17">[17]</a> A meaning of 'agent that causes infectious disease' is first recorded in 1728,<a href="#cite_note-Etymology_Dictionary-15">[15]</a> long before the discovery of viruses by <a href="/wiki/Dmitri_Ivanovsky" title="Dmitri Ivanovsky">Dmitri Ivanovsky</a> in 1892. The English <a href="/wiki/Plural_form_of_words_ending_in_-us#Virus" title="Plural form of words ending in -us">plural</a> is viruses (sometimes also vira),<a href="#cite_note-18">[18]</a> whereas the Latin word is a <a href="/wiki/Mass_noun" title="Mass noun">mass noun</a>, which has no <a href="/wiki/Classical_Latin" title="Classical Latin">classically</a> attested plural (vīra is used in <a href="/wiki/Neo-Latin" title="Neo-Latin">Neo-Latin</a><a href="#cite_note-19">[19]</a>). The adjective viral dates to 1948.<a href="#cite_note-20">[20]</a> The term virion (plural virions), which dates from 1959,<a href="#cite_note-Etymology_Dictionary2-21">[21]</a> is also used to refer to a single viral particle that is released from the cell and is capable of infecting other cells of the same type.<a href="#cite_note-22">[22]</a> <div class="mw-heading mw-heading2"><h2 id="Origins">Origins</h2></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Viral_evolution" title="Viral evolution">Viral evolution</a></div> Viruses are found wherever there is life and have probably existed since living cells <a href="/wiki/Abiogenesis" title="Abiogenesis">first evolved</a>.<a href="#cite_note-Iyer_2006-23">[23]</a> The origin of viruses is unclear because they do not form fossils, so <a href="/wiki/Molecular_biology" title="Molecular biology">molecular techniques</a> are used to infer how they arose.<a href="#cite_note-Sanjuán_2010-24">[24]</a> In addition, viral genetic material occasionally integrates into the <a href="/wiki/Germline" title="Germline">germline</a> of the host organisms, by which they can be passed on <a href="/wiki/Vertical_transfer" class="mw-redirect" title="Vertical transfer">vertically</a> to the offspring of the host for many generations. This provides an invaluable source of information for <a href="/wiki/Paleovirology" title="Paleovirology">paleovirologists</a> to trace back ancient viruses that existed as far back as millions of years ago. There are three main hypotheses that aim to explain the origins of viruses:<a href="#cite_note-Krupovic_2019-25">[25]</a> <dl><dt>Regressive hypothesis</dt> <dd>Viruses may have once been small cells that <a href="/wiki/Parasitism" title="Parasitism">parasitised</a> larger cells. Over time, genes not required by their parasitism were lost. The bacteria <a href="/wiki/Rickettsia" title="Rickettsia">rickettsia</a> and <a href="/wiki/Chlamydia_(genus)" title="Chlamydia (genus)">chlamydia</a> are living cells that, like viruses, can reproduce only inside host cells. They lend support to this hypothesis, as their dependence on parasitism is likely to have caused the loss of genes that enabled them to survive outside a cell. This is also called the "degeneracy hypothesis",<a href="#cite_note-Dimmock_2007-6">[6]</a>: 16 <a href="#cite_note-Collier_1998-26">[26]</a>: 11  or "reduction hypothesis".<a href="#cite_note-Mahy_2009-27">[27]</a>: 24 </dd> <dt>Cellular origin hypothesis</dt> <dd>Some viruses may have evolved from bits of DNA or RNA that "escaped" from the genes of a larger organism. The escaped DNA could have come from <a href="/wiki/Plasmid" title="Plasmid">plasmids</a> (pieces of naked DNA that can move between cells) or <a href="/wiki/Transposons" class="mw-redirect" title="Transposons">transposons</a> (molecules of DNA that replicate and move around to different positions within the genes of the cell).<a href="#cite_note-Shors_2017-13">[13]</a>: 810  Once called "jumping genes", transposons are examples of <a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">mobile genetic elements</a> and could be the origin of some viruses. They were discovered in maize by <a href="/wiki/Barbara_McClintock" title="Barbara McClintock">Barbara McClintock</a> in 1950.<a href="#cite_note-28">[28]</a> This is sometimes called the "vagrancy hypothesis",<a href="#cite_note-Dimmock_2007-6">[6]</a>: 16 <a href="#cite_note-Collier_1998-26">[26]</a>: 11–12  or the "escape hypothesis".<a href="#cite_note-Mahy_2009-27">[27]</a>: 24 </dd> <dt>Co-evolution hypothesis</dt> <dd>This is also called the "virus-first hypothesis"<a href="#cite_note-Mahy_2009-27">[27]</a>: 24  and proposes that viruses may have evolved from complex molecules of protein and <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acid</a> at the same time that cells first appeared on Earth and would have been dependent on cellular life for billions of years. <a href="/wiki/Viroids" class="mw-redirect" title="Viroids">Viroids</a> are molecules of RNA that are not classified as viruses because they lack a protein coat. They have characteristics that are common to several viruses and are often called <a href="/wiki/Virus_classification#Subviral_agents" title="Virus classification">subviral agents</a>.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 55  Viroids are important pathogens of plants.<a href="#cite_note-Shors_2017-13">[13]</a>: 791  They do not code for proteins but interact with the host cell and use the host machinery for their replication.<a href="#cite_note-Tsagris_2008-29">[29]</a> The <a href="/wiki/Hepatitis_delta_virus" class="mw-redirect" title="Hepatitis delta virus">hepatitis delta virus</a> of humans has an RNA <a href="/wiki/Genome" title="Genome">genome</a> similar to viroids but has a protein coat derived from hepatitis B virus and cannot produce one of its own. It is, therefore, a defective virus. Although hepatitis delta virus genome may replicate independently once inside a host cell, it requires the help of hepatitis B virus to provide a protein coat so that it can be transmitted to new cells.<a href="#cite_note-Shors_2017-13">[13]</a>: 460  In similar manner, the <a href="/wiki/Sputnik_virophage" title="Sputnik virophage">sputnik virophage</a> is dependent on <a href="/wiki/Mimivirus" title="Mimivirus">mimivirus</a>, which infects the protozoan <a href="/wiki/Acanthamoeba" title="Acanthamoeba">Acanthamoeba</a> castellanii.<a href="#cite_note-La_Scola_2008-30">[30]</a> These viruses, which are dependent on the presence of other virus species in the host cell, are called "<a href="/wiki/Satellite_(biology)" title="Satellite (biology)">satellites</a>" and may represent evolutionary intermediates of viroids and viruses.<a href="#cite_note-Collier_1998-26">[26]</a>: 777 <a href="#cite_note-Dimmock_2007-6">[6]</a>: 55–57 </dd></dl> In the past, there were problems with all of these hypotheses: the regressive hypothesis did not explain why even the smallest of cellular parasites do not resemble viruses in any way. The escape hypothesis did not explain the complex capsids and other structures on virus particles. The virus-first hypothesis contravened the definition of viruses in that they require host cells.<a href="#cite_note-Mahy_2009-27">[27]</a>: 24  Viruses are now recognised as ancient and as having origins that pre-date the divergence of life into the <a href="/wiki/Three-domain_system" title="Three-domain system">three domains</a>.<a href="#cite_note-Mahy_2009-27">[27]</a>: 28  This discovery has led modern virologists to reconsider and re-evaluate these three classical hypotheses.<a href="#cite_note-Mahy_2009-27">[27]</a>: 28  The evidence for an <a href="/wiki/RNA_world" title="RNA world">ancestral world of RNA</a> cells<a href="#cite_note-Mahy_2009-27">[27]</a>: 26  and computer analysis of viral and host DNA sequences give a better understanding of the evolutionary relationships between different viruses and may help identify the ancestors of modern viruses. To date, such analyses have not proved which of these hypotheses is correct.<a href="#cite_note-Mahy_2009-27">[27]</a>: 26  It seems unlikely that all currently known viruses have a common ancestor, and viruses have probably arisen numerous times in the past by one or more mechanisms.<a href="#cite_note-31">[31]</a> <div class="mw-heading mw-heading2"><h2 id="Microbiology">Microbiology</h2></div> <div class="mw-heading mw-heading3"><h3 id="Discovery">Discovery</h3></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/History_of_virology" title="History of virology">History of virology</a></div> The first evidence of the existence of viruses came from experiments with filters that had pores small enough to retain bacteria. In 1892, <a href="/wiki/Dmitri_Ivanovsky" title="Dmitri Ivanovsky">Dmitri Ivanovsky</a> used one of these filters to show that sap from a diseased tobacco plant remained infectious to healthy tobacco plants despite having been filtered. Martinus Beijerinck called the filtered, infectious substance a "virus" and this discovery is considered to be the beginning of virology. The subsequent discovery and partial characterization of <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophages</a> by <a href="/wiki/Frederick_Twort" title="Frederick Twort">Frederick Twort</a> and <a href="/wiki/F%C3%A9lix_d%27Herelle" class="mw-redirect" title="Félix d'Herelle">Félix d'Herelle</a> further catalyzed the field, and by the early 20th century many viruses had been discovered. In 1926, <a href="/wiki/Thomas_Milton_Rivers" title="Thomas Milton Rivers">Thomas Milton Rivers</a> defined viruses as obligate parasites. Viruses were demonstrated to be particles, rather than a fluid, by <a href="/wiki/Wendell_Meredith_Stanley" title="Wendell Meredith Stanley">Wendell Meredith Stanley</a>, and the invention of the <a href="/wiki/Electron_microscope" title="Electron microscope">electron microscope</a> in 1931 allowed their complex structures to be visualised.<a href="#cite_note-pmid36291572-32">[32]</a> <div class="mw-heading mw-heading3"><h3 id="Life_properties">Life properties</h3></div> Scientific opinions differ on whether viruses are a form of life or organic structures that interact with living organisms.<a href="#cite_note-Koonin_2016-11">[11]</a> They have been described as "organisms at the edge of life",<a href="#cite_note-Rybicki_1990-10">[10]</a> since they resemble organisms in that they possess <a href="/wiki/Genes" class="mw-redirect" title="Genes">genes</a>, evolve by <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>,<a href="#cite_note-Holmes_2007-33">[33]</a> and reproduce by creating multiple copies of themselves through self-assembly. Although they have genes, they do not have a cellular structure, which is often seen as the basic unit of life. Viruses do not have their own <a href="/wiki/Metabolism" title="Metabolism">metabolism</a> and require a host cell to make new products. They therefore cannot naturally reproduce outside a host cell<a href="#cite_note-Wimmer_2009-34">[34]</a>—although some bacteria such as <a href="/wiki/Rickettsia" title="Rickettsia">rickettsia</a> and <a href="/wiki/Chlamydia_(genus)" title="Chlamydia (genus)">chlamydia</a> are considered living organisms despite the same limitation.<a href="#cite_note-Horn_2008-35">[35]</a><a href="#cite_note-Ammerman_2008-36">[36]</a> Accepted forms of life use <a href="/wiki/Cell_division" title="Cell division">cell division</a> to reproduce, whereas viruses spontaneously assemble within cells. They differ from <a href="/wiki/Self-organisation" class="mw-redirect" title="Self-organisation">autonomous growth</a> of <a href="/wiki/Crystallization" title="Crystallization">crystals</a> as they inherit genetic mutations while being subject to natural selection. Virus self-assembly within host cells has implications for the study of the <a href="/wiki/Origin_of_life" class="mw-redirect" title="Origin of life">origin of life</a>, as it lends further credence to the hypothesis that life could have started as <a href="/wiki/Molecular_self-assembly" title="Molecular self-assembly">self-assembling organic molecules</a>.<a href="#cite_note-Koonin_2006-2">[2]</a> The <a href="/wiki/Virocell" class="mw-redirect" title="Virocell">virocell</a> model first proposed by <a href="/wiki/Patrick_Forterre" title="Patrick Forterre">Patrick Forterre</a> considers the infected cell to be the "living form" of viruses and that virus particles (virions) are analogous to <a href="/wiki/Spore" title="Spore">spores</a>.<a href="#cite_note-37">[37]</a> Although the living versus non-living debate continues, the virocell model has gained some acceptance.<a href="#cite_note-pmid34522049-38">[38]</a> <div class="mw-heading mw-heading3"><h3 id="Structure">Structure</h3></div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:204px;max-width:204px"><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:Virus_size.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Virus_size.png/200px-Virus_size.png" decoding="async" width="200" height="117" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Virus_size.png/300px-Virus_size.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/70/Virus_size.png/400px-Virus_size.png 2x" data-file-width="922" data-file-height="538" /></a></div><div class="thumbcaption">Virions of some of the most common human viruses with their relative size. The nucleic acids are not to scale.</div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:Hexon.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Hexon.svg/200px-Hexon.svg.png" decoding="async" width="200" height="132" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Hexon.svg/300px-Hexon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Hexon.svg/400px-Hexon.svg.png 2x" data-file-width="439" data-file-height="290" /></a></div><div class="thumbcaption">Diagram of how a virus <a href="/wiki/Capsid" title="Capsid">capsid</a> can be constructed using multiple copies of just two protein molecules</div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:TMV_structure_simple.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/TMV_structure_simple.png/200px-TMV_structure_simple.png" decoding="async" width="200" height="142" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/TMV_structure_simple.png/300px-TMV_structure_simple.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8e/TMV_structure_simple.png/400px-TMV_structure_simple.png 2x" data-file-width="1474" data-file-height="1050" /></a></div><div class="thumbcaption">Structure of <a href="/wiki/Tobacco_mosaic_virus" title="Tobacco mosaic virus">tobacco mosaic virus</a>: <a href="/wiki/RNA" title="RNA">RNA</a> coiled in a helix of repeating protein sub-units</div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:Icosahedral_Adenoviruses.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Icosahedral_Adenoviruses.jpg/200px-Icosahedral_Adenoviruses.jpg" decoding="async" width="200" height="80" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Icosahedral_Adenoviruses.jpg/300px-Icosahedral_Adenoviruses.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Icosahedral_Adenoviruses.jpg/400px-Icosahedral_Adenoviruses.jpg 2x" data-file-width="889" data-file-height="356" /></a></div><div class="thumbcaption">Structure of icosahedral <a href="/wiki/Adenovirus" class="mw-redirect" title="Adenovirus">adenovirus</a>. Electron micrograph with an illustration to show shape</div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:Varicella_(Chickenpox)_Virus_PHIL_1878_lores.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Varicella_%28Chickenpox%29_Virus_PHIL_1878_lores.jpg/200px-Varicella_%28Chickenpox%29_Virus_PHIL_1878_lores.jpg" decoding="async" width="200" height="199" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Varicella_%28Chickenpox%29_Virus_PHIL_1878_lores.jpg/300px-Varicella_%28Chickenpox%29_Virus_PHIL_1878_lores.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/1/16/Varicella_%28Chickenpox%29_Virus_PHIL_1878_lores.jpg 2x" data-file-width="367" data-file-height="366" /></a></div><div class="thumbcaption">Structure of <a href="/wiki/Chickenpox" title="Chickenpox">chickenpox</a> virus. They have a lipid envelope.</div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:CowpeaMosaicVirus3D.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/CowpeaMosaicVirus3D.png/200px-CowpeaMosaicVirus3D.png" decoding="async" width="200" height="203" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/CowpeaMosaicVirus3D.png/300px-CowpeaMosaicVirus3D.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c0/CowpeaMosaicVirus3D.png/400px-CowpeaMosaicVirus3D.png 2x" data-file-width="777" data-file-height="789" /></a></div><div class="thumbcaption">Structure of an icosahedral <a href="/wiki/Cowpea_mosaic_virus" title="Cowpea mosaic virus">cowpea mosaic virus</a></div></div></div><div class="trow"><div class="tsingle" style="width:202px;max-width:202px"><div class="thumbimage"><a href="/wiki/File:Ms2capsid_surface.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7e/Ms2capsid_surface.png/200px-Ms2capsid_surface.png" decoding="async" width="200" height="200" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7e/Ms2capsid_surface.png/300px-Ms2capsid_surface.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7e/Ms2capsid_surface.png/400px-Ms2capsid_surface.png 2x" data-file-width="1600" data-file-height="1600" /></a></div><div class="thumbcaption">Bacteriophage <a href="/wiki/Escherichia_virus_MS2" class="mw-redirect" title="Escherichia virus MS2">Escherichia virus MS2</a> capsid. This spherical virus also has icosahedral symmetry.</div></div></div></div></div> Viruses display a wide diversity of sizes and shapes, called '<a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphologies</a>'. In general, viruses are much smaller than bacteria and more than a thousand bacteriophage viruses would fit inside an <a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a> bacterium's cell.<a href="#cite_note-Krasner_2014-39">[39]</a>: 98  Many viruses that have been studied are spherical and have a diameter between 20 and 300 <a href="/wiki/Nanometres" class="mw-redirect" title="Nanometres">nanometres</a>. Some <a href="/wiki/Filovirus" class="mw-redirect" title="Filovirus">filoviruses</a>, which are filaments, have a total length of up to 1400 nm; their diameters are only about 80 nm.<a href="#cite_note-Collier_1998-26">[26]</a>: 33–55  Most viruses cannot be seen with an <a href="/wiki/Optical_microscope" title="Optical microscope">optical microscope</a>, so scanning and transmission <a href="/wiki/Electron_microscope" title="Electron microscope">electron microscopes</a> are used to visualise them.<a href="#cite_note-Collier_1998-26">[26]</a>: 33–37  To increase the contrast between viruses and the background, electron-dense "stains" are used. These are solutions of <a href="/wiki/Salt_(chemistry)" title="Salt (chemistry)">salts</a> of heavy metals, such as <a href="/wiki/Tungsten" title="Tungsten">tungsten</a>, that scatter the electrons from regions covered with the stain. When virions are coated with stain (positive staining), fine detail is obscured. <a href="/wiki/Negative_staining" class="mw-redirect" title="Negative staining">Negative staining</a> overcomes this problem by staining the background only.<a href="#cite_note-Kiselev_1990-40">[40]</a> A complete virus particle, known as a virion, consists of nucleic acid surrounded by a protective coat of protein called a <a href="/wiki/Capsid" title="Capsid">capsid</a>. These are formed from protein subunits called <a href="/wiki/Capsomere" title="Capsomere">capsomeres</a>.<a href="#cite_note-Collier_1998-26">[26]</a>: 40  Viruses can have a <a href="/wiki/Lipid" title="Lipid">lipid</a> "envelope" derived from the host <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a>. The capsid is made from proteins encoded by the viral <a href="/wiki/Genome" title="Genome">genome</a> and its shape serves as the basis for morphological distinction.<a href="#cite_note-Caspar_1962-41">[41]</a><a href="#cite_note-Crick_1956-42">[42]</a> Virally-coded protein subunits will self-assemble to form a capsid, in general requiring the presence of the virus genome. Complex viruses code for proteins that assist in the construction of their capsid. Proteins associated with nucleic acid are known as <a href="/wiki/Nucleoprotein" title="Nucleoprotein">nucleoproteins</a>, and the association of viral capsid proteins with viral nucleic acid is called a nucleocapsid. The capsid and entire virus structure can be mechanically (physically) probed through <a href="/wiki/Atomic_force_microscopy" title="Atomic force microscopy">atomic force microscopy</a>.<a href="#cite_note-Falvo_1997-43">[43]</a><a href="#cite_note-Kuznetsov_2001-44">[44]</a> In general, there are five main morphological virus types: <dl><dt>Helical</dt> <dd>These viruses are composed of a single type of capsomere stacked around a central axis to form a <a href="/wiki/Helix" title="Helix">helical</a> structure, which may have a central cavity, or tube. This arrangement results in virions which can be short and highly rigid rods, or long and very flexible filaments. The genetic material (typically single-stranded RNA, but single-stranded DNA in some cases) is bound into the protein helix by interactions between the negatively charged nucleic acid and positive charges on the protein. Overall, the length of a helical capsid is related to the length of the nucleic acid contained within it, and the diameter is dependent on the size and arrangement of capsomeres. The well-studied tobacco mosaic virus<a href="#cite_note-Collier_1998-26">[26]</a>: 37 and inovirus<a href="#cite_note-45">[45]</a> are examples of helical viruses.</dd></dl> <dl><dt>Icosahedral</dt> <dd>Most animal viruses are icosahedral or near-spherical with chiral <a href="/wiki/Icosahedral_symmetry" title="Icosahedral symmetry">icosahedral symmetry</a>. A <a href="/wiki/Regular_icosahedron" title="Regular icosahedron">regular icosahedron</a> is the optimum way of forming a closed shell from identical subunits. The minimum number of capsomeres required for each triangular face is 3, which gives 60 for the icosahedron. Many viruses, such as rotavirus, have more than 60 capsomers and appear spherical but they retain this symmetry. To achieve this, the capsomeres at the apices are surrounded by five other capsomeres and are called pentons. Capsomeres on the triangular faces are surrounded by six others and are called <a href="/wiki/Hexon_protein" title="Hexon protein">hexons</a>.<a href="#cite_note-Collier_1998-26">[26]</a>: 40, 42  Hexons are in essence flat and pentons, which form the 12 vertices, are curved. The same protein may act as the subunit of both the pentamers and hexamers or they may be composed of different proteins.<a href="#cite_note-Wilson_2016-46">[46]</a></dd></dl> <dl><dt>Prolate</dt> <dd>This is an icosahedron elongated along the fivefold axis and is a common arrangement of the heads of bacteriophages. This structure is composed of a cylinder with a cap at either end.<a href="#cite_note-47">[47]</a></dd></dl> <dl><dt>Enveloped</dt> <dd>Some species of virus <a href="/wiki/Viral_envelope" title="Viral envelope">envelop</a> themselves in a modified form of one of the <a href="/wiki/Cell_membranes" class="mw-redirect" title="Cell membranes">cell membranes</a>, either the outer membrane surrounding an infected host cell or internal membranes such as a nuclear membrane or <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a>, thus gaining an outer lipid bilayer known as a <a href="/wiki/Viral_envelope" title="Viral envelope">viral envelope</a>. This membrane is studded with proteins coded for by the viral genome and host genome; the lipid membrane itself and any carbohydrates present originate entirely from the host. <a href="/wiki/Influenza_virus" class="mw-redirect" title="Influenza virus">Influenza virus</a>, <a href="/wiki/HIV" title="HIV">HIV</a> (which causes <a href="/wiki/AIDS" class="mw-redirect" title="AIDS">AIDS</a>), and <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> (which causes <a href="/wiki/COVID-19" title="COVID-19">COVID-19</a>)<a href="#cite_note-Dhama_2020-48">[48]</a> use this strategy. Most enveloped viruses are dependent on the envelope for their infectivity.<a href="#cite_note-Collier_1998-26">[26]</a>: 42–43 </dd></dl> <dl><dt>Complex</dt> <dd>These viruses possess a capsid that is neither purely helical nor purely icosahedral, and that may possess extra structures such as protein tails or a complex outer wall. Some bacteriophages, such as <a href="/wiki/Enterobacteria_phage_T4" class="mw-redirect" title="Enterobacteria phage T4">Enterobacteria phage T4</a>, have a complex structure consisting of an icosahedral head bound to a helical tail, which may have a <a href="/wiki/Hexagon" title="Hexagon">hexagonal</a> base plate with protruding protein tail fibres. This tail structure acts like a molecular syringe, attaching to the bacterial host and then injecting the viral genome into the cell.<a href="#cite_note-49">[49]</a></dd></dl> The <a href="/wiki/Poxviridae" title="Poxviridae">poxviruses</a> are large, complex viruses that have an unusual morphology. The viral genome is associated with proteins within a central disc structure known as a <a href="/wiki/Nucleoid" title="Nucleoid">nucleoid</a>. The nucleoid is surrounded by a membrane and two lateral bodies of unknown function. The virus has an outer envelope with a thick layer of protein studded over its surface. The whole virion is slightly <a href="/wiki/Pleomorphism_(microbiology)" title="Pleomorphism (microbiology)">pleomorphic</a>, ranging from ovoid to brick-shaped.<a href="#cite_note-Long_1970-50">[50]</a> <div class="mw-heading mw-heading4"><h4 id="Giant_viruses">Giant viruses</h4></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/Giant_virus" title="Giant virus">Giant virus</a></div> <a href="/wiki/Mimivirus" title="Mimivirus">Mimivirus</a> is one of the largest characterised viruses, with a capsid diameter of 400 nm. Protein filaments measuring 100 nm project from the surface. The capsid appears hexagonal under an electron microscope, therefore the capsid is probably icosahedral.<a href="#cite_note-Suzan-Monti_2006-51">[51]</a> In 2011, researchers discovered the largest then known virus in samples of water collected from the ocean floor off the coast of Las Cruces, Chile. Provisionally named <a href="/wiki/Megavirus" title="Megavirus">Megavirus</a> chilensis, it can be seen with a basic optical microscope.<a href="#cite_note-Arslan_2011-52">[52]</a> In 2013, the <a href="/wiki/Pandoravirus" title="Pandoravirus">Pandoravirus</a> genus was discovered in Chile and Australia, and has genomes about twice as large as Megavirus and Mimivirus.<a href="#cite_note-Philippe_2013-53">[53]</a> All giant viruses have dsDNA genomes and they are classified into several families: <a href="/wiki/Mimiviridae" title="Mimiviridae">Mimiviridae</a>, Pithoviridae, <a href="/wiki/Pandoraviridae" title="Pandoraviridae">Pandoraviridae</a>, <a href="/wiki/Phycodnaviridae" title="Phycodnaviridae">Phycodnaviridae</a>, and the <a href="/wiki/Mollivirus" class="mw-redirect" title="Mollivirus">Mollivirus</a> genus.<a href="#cite_note-Brandes_2019-54">[54]</a> Some viruses that infect <a href="/wiki/Archaea" title="Archaea">Archaea</a> have complex structures unrelated to any other form of virus, with a wide variety of unusual shapes, ranging from spindle-shaped structures to viruses that resemble hooked rods, teardrops or even bottles. Other archaeal viruses resemble the tailed bacteriophages, and can have multiple tail structures.<a href="#cite_note-Prangishvili-55">[55]</a> <div class="mw-heading mw-heading3"><h3 id="Genome">Genome</h3></div> <table class="wikitable floatright" width="275px"> <caption>Genomic diversity among viruses </caption> <tbody><tr> <th>Property</th> <th>Parameters </th></tr> <tr> <td>Nucleic acid</td> <td> <ul><li>DNA</li> <li>RNA</li> <li>Both DNA and RNA (at different stages in the life cycle)</li></ul> </td></tr> <tr> <td>Shape</td> <td> <ul><li>Linear</li> <li>Circular</li> <li>Segmented</li></ul> </td></tr> <tr> <td>Strandedness</td> <td> <ul><li>Single-stranded (ss)</li> <li>Double-stranded (ds)</li> <li>Double-stranded with regions of single-strandedness</li></ul> </td></tr> <tr> <td><a href="/wiki/Sense_(molecular_biology)" title="Sense (molecular biology)">Sense</a></td> <td> <ul><li>Positive sense (+)</li> <li>Negative sense (−)</li> <li>Ambisense (+/−)</li></ul> </td></tr></tbody></table> An enormous variety of genomic structures can be seen among <a href="/wiki/Viral_species" class="mw-redirect" title="Viral species">viral species</a>; as a group, they contain more structural genomic diversity than plants, animals, archaea, or bacteria. There are millions of different types of viruses,<a href="#cite_note-Breitbart_2005-8">[8]</a> although fewer than 7,000 types have been described in detail.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 49 As of January 2021, the <a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a> Virus genome database has more than 193,000 complete genome sequences,<a href="#cite_note-NCBI_VGDB-56">[56]</a> but there are doubtlessly many more to be discovered.<a href="#cite_note-57">[57]</a><a href="#cite_note-58">[58]</a> A virus has either a <a href="/wiki/DNA" title="DNA">DNA</a> or an <a href="/wiki/RNA" title="RNA">RNA</a> genome and is called a <a href="/wiki/DNA_virus" title="DNA virus">DNA virus</a> or an <a href="/wiki/RNA_virus" title="RNA virus">RNA virus</a>, respectively. Most viruses have RNA genomes. Plant viruses tend to have single-stranded RNA genomes and bacteriophages tend to have double-stranded DNA genomes.<a href="#cite_note-Collier_1998-26">[26]</a>: 96–99  Viral genomes are circular, as in the <a href="/wiki/Polyomavirus" class="mw-redirect" title="Polyomavirus">polyomaviruses</a>, or linear, as in the <a href="/wiki/Adenoviridae" title="Adenoviridae">adenoviruses</a>. The type of nucleic acid is irrelevant to the shape of the genome. Among RNA viruses and certain DNA viruses, the genome is often divided into separate parts, in which case it is called segmented. For RNA viruses, each segment often codes for only one protein and they are usually found together in one capsid. All segments are not required to be in the same virion for the virus to be infectious, as demonstrated by <a href="/wiki/Brome_mosaic_virus" title="Brome mosaic virus">brome mosaic virus</a> and several other plant viruses.<a href="#cite_note-Collier_1998-26">[26]</a>: 33–35  A viral genome, irrespective of nucleic acid type, is almost always either single-stranded (ss) or double-stranded (ds). Single-stranded genomes consist of an unpaired nucleic acid, analogous to one-half of a ladder split down the middle. Double-stranded genomes consist of two complementary paired nucleic acids, analogous to a ladder. The virus particles of some virus families, such as those belonging to the <a href="/wiki/Hepadnaviridae" title="Hepadnaviridae">Hepadnaviridae</a>, contain a genome that is partially double-stranded and partially single-stranded.<a href="#cite_note-Collier_1998-26">[26]</a>: 96–99  For most viruses with RNA genomes and some with single-stranded DNA (ssDNA) genomes, the single strands are said to be either <a href="/wiki/Positive-sense" class="mw-redirect" title="Positive-sense">positive-sense</a> (called the 'plus-strand') or <a href="/wiki/Negative-sense" class="mw-redirect" title="Negative-sense">negative-sense</a> (called the 'minus-strand'), depending on if they are complementary to the viral <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> (mRNA). Positive-sense viral RNA is in the same sense as viral mRNA and thus at least a part of it can be immediately <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">translated</a> by the host cell. Negative-sense viral RNA is complementary to mRNA and thus must be converted to positive-sense RNA by an <a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA-dependent RNA polymerase</a> before translation. DNA nomenclature for viruses with genomic ssDNA is similar to RNA nomenclature, in that positive-strand viral ssDNA is identical in sequence to the viral mRNA and is thus a coding strand, while negative-sense viral ssDNA is complementary to the viral mRNA and is thus a template strand.<a href="#cite_note-Collier_1998-26">[26]</a>: 96–99  Several types of ssDNA and ssRNA viruses have genomes that are <a href="/wiki/Ambisense" class="mw-redirect" title="Ambisense">ambisense</a> in that transcription can occur off both strands in a double-stranded replicative intermediate. Examples include <a href="/wiki/Geminiviridae" title="Geminiviridae">geminiviruses</a>, which are ssDNA plant viruses and <a href="/wiki/Arenavirus" title="Arenavirus">arenaviruses</a>, which are ssRNA viruses of animals.<a href="#cite_note-isbn0-470-02387-2-59">[59]</a> <div class="mw-heading mw-heading4"><h4 id="Genome_size">Genome size</h4></div> Genome size varies greatly between species. The smallest—the ssDNA circoviruses, family <a href="/wiki/Circoviridae" title="Circoviridae">Circoviridae</a>—code for only two proteins and have a genome size of only two kilobases;<a href="#cite_note-60">[60]</a> the largest—the <a href="/wiki/Pandoravirus" title="Pandoravirus">pandoraviruses</a>—have genome sizes of around two megabases which code for about 2500 proteins.<a href="#cite_note-Philippe_2013-53">[53]</a> Virus genes rarely have <a href="/wiki/Intron" title="Intron">introns</a> and often are arranged in the genome so that they <a href="/wiki/Overlapping_gene" title="Overlapping gene">overlap</a>.<a href="#cite_note-Brandes_2016-61">[61]</a> In general, RNA viruses have smaller genome sizes than DNA viruses because of a higher error-rate when replicating, and have a maximum upper size limit.<a href="#cite_note-Sanjuán_2010-24">[24]</a> Beyond this, errors when replicating render the virus useless or uncompetitive. To compensate, RNA viruses often have segmented genomes—the genome is split into smaller molecules—thus reducing the chance that an error in a single-component genome will incapacitate the entire genome. In contrast, DNA viruses generally have larger genomes because of the high fidelity of their replication enzymes.<a href="#cite_note-62">[62]</a> Single-strand DNA viruses are an exception to this rule, as mutation rates for these genomes can approach the extreme of the ssRNA virus case.<a href="#cite_note-Duffy_2009-63">[63]</a> <div class="mw-heading mw-heading3"><h3 id="Genetic_mutation_and_recombination">Genetic mutation and recombination</h3></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Viruses-10-00497-g003.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Viruses-10-00497-g003.png/220px-Viruses-10-00497-g003.png" decoding="async" width="220" height="154" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Viruses-10-00497-g003.png/330px-Viruses-10-00497-g003.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6d/Viruses-10-00497-g003.png/440px-Viruses-10-00497-g003.png 2x" data-file-width="3517" data-file-height="2462" /></a><figcaption>Evolution mechanisms of Influenza A virus. (A) Antigenic Drift: Gradual accumulation of mutations in the genome of IAVs leads to emergence of new virus variants. (B) Antigenic Shift: The reassortment of genetic segments between two or more invading IAVs in a host cell can lead to emergence of an antigenically novel subtype.</figcaption></figure> Viruses undergo genetic change by several mechanisms. These include a process called <a href="/wiki/Antigenic_drift" title="Antigenic drift">antigenic drift</a> where individual bases in the DNA or RNA <a href="/wiki/Mutate" class="mw-redirect" title="Mutate">mutate</a> to other bases. Most of these <a href="/wiki/Point_mutations" class="mw-redirect" title="Point mutations">point mutations</a> are "silent"—they do not change the protein that the gene encodes—but others can confer evolutionary advantages such as resistance to <a href="/wiki/Antiviral_drugs" class="mw-redirect" title="Antiviral drugs">antiviral drugs</a>.<a href="#cite_note-Sandbulte_2011-64">[64]</a><a href="#cite_note-Moss_2010-65">[65]</a> <a href="/wiki/Antigenic_shift" title="Antigenic shift">Antigenic shift</a> occurs when there is a major change in the genome of the virus. This can be a result of <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a> or <a href="/wiki/Reassortment" title="Reassortment">reassortment</a>. The <a href="/wiki/Influenza_A_virus" title="Influenza A virus">Influenza A virus</a> is highly prone to reassortment; occasionally this has resulted in novel <a href="/wiki/Strain_(biology)" title="Strain (biology)">strains</a> which have caused <a href="/wiki/Pandemics" class="mw-redirect" title="Pandemics">pandemics</a>.<a href="#cite_note-Hampson_2006-66">[66]</a> RNA viruses often exist as <a href="/wiki/Quasispecies" class="mw-redirect" title="Quasispecies">quasispecies</a> or swarms of viruses of the same species but with slightly different genome nucleoside sequences. Such quasispecies are a prime target for natural selection.<a href="#cite_note-Metzner_2006-67">[67]</a> Segmented genomes confer evolutionary advantages; different strains of a virus with a segmented genome can shuffle and combine genes and produce progeny viruses (or offspring) that have unique characteristics. This is called reassortment or 'viral sex'.<a href="#cite_note-68">[68]</a> <a href="/wiki/Genetic_recombination" title="Genetic recombination">Genetic recombination</a> is a process by which a strand of DNA (or RNA) is broken and then joined to the end of a different DNA (or RNA) molecule. This can occur when viruses infect cells simultaneously and studies of <a href="/wiki/Viral_evolution" title="Viral evolution">viral evolution</a> have shown that recombination has been rampant in the species studied.<a href="#cite_note-Worobey_1999-69">[69]</a> Recombination is common to both RNA and DNA viruses.<a href="#cite_note-Lukashev_2005-70">[70]</a><a href="#cite_note-Umene_1999-71">[71]</a> <a href="/wiki/Coronavirus" title="Coronavirus">Coronaviruses</a> have a single-strand positive-sense <a href="/wiki/RNA" title="RNA">RNA</a> genome. Replication of the <a href="/wiki/Genome" title="Genome">genome</a> is catalyzed by an <a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA-dependent RNA polymerase</a>. The mechanism of <a href="/wiki/Homologous_recombination" title="Homologous recombination">recombination</a> used by coronaviruses likely involves template switching by the polymerase during genome replication.<a href="#cite_note-Su_2016-72">[72]</a> This process appears to be an adaptation for coping with genome damage.<a href="#cite_note-Barr_2010-73">[73]</a> <div class="mw-heading mw-heading3"><h3 id="Replication_cycle">Replication cycle</h3></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:HepC_replication.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/da/HepC_replication.png/220px-HepC_replication.png" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/da/HepC_replication.png/330px-HepC_replication.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/da/HepC_replication.png/440px-HepC_replication.png 2x" data-file-width="960" data-file-height="720" /></a><figcaption>A typical virus replication cycle</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Phage_injecting_its_genome_into_bacteria.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/Phage_injecting_its_genome_into_bacteria.svg/220px-Phage_injecting_its_genome_into_bacteria.svg.png" decoding="async" width="220" height="191" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/Phage_injecting_its_genome_into_bacteria.svg/330px-Phage_injecting_its_genome_into_bacteria.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/29/Phage_injecting_its_genome_into_bacteria.svg/440px-Phage_injecting_its_genome_into_bacteria.svg.png 2x" data-file-width="2316" data-file-height="2013" /></a><figcaption>Some bacteriophages inject their <a href="/wiki/Genome" title="Genome">genomes</a> into bacterial cells (not to scale)</figcaption></figure> Viral populations do not grow through cell division, because they are acellular. Instead, they use the machinery and metabolism of a host cell to produce multiple copies of themselves, and they assemble in the cell.<a href="#cite_note-Freed_2015-74">[74]</a> When infected, the host cell is forced to rapidly produce thousands of copies of the original virus.<a href="#cite_note-Yin_2018-75">[75]</a> Their life cycle differs greatly between species, but there are six basic stages in their life cycle:<a href="#cite_note-Collier_1998-26">[26]</a>: 75–91  Attachment is a specific binding between viral capsid proteins and specific receptors on the host cellular surface. This specificity determines the host range and type of host cell of a virus. For example, HIV infects a limited range of human <a href="/wiki/Leucocytes" class="mw-redirect" title="Leucocytes">leucocytes</a>. This is because its surface protein, <a href="/wiki/Gp120" class="mw-redirect" title="Gp120">gp120</a>, specifically interacts with the <a href="/wiki/CD4" title="CD4">CD4</a> molecule—a <a href="/wiki/Chemokine_receptor" title="Chemokine receptor">chemokine receptor</a>—which is most commonly found on the surface of <a href="/wiki/CD4%2B" class="mw-redirect" title="CD4+">CD4+</a> <a href="/wiki/T-Cells" class="mw-redirect" title="T-Cells">T-Cells</a>. This mechanism has evolved to favour those viruses that infect only cells in which they are capable of replication. Attachment to the receptor can induce the viral envelope protein to undergo changes that result in the <a href="/wiki/Lipid_bilayer_fusion" title="Lipid bilayer fusion">fusion</a> of viral and cellular membranes, or changes of non-enveloped virus surface proteins that allow the virus to enter.<a href="#cite_note-Más_2013-76">[76]</a> Penetration or <a href="/wiki/Viral_entry" title="Viral entry">viral entry</a> follows attachment: Virions enter the host cell through receptor-mediated <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a> or <a href="/wiki/Lipid_bilayer_fusion" title="Lipid bilayer fusion">membrane fusion</a>. The infection of plant and fungal cells is different from that of animal cells. Plants have a rigid cell wall made of <a href="/wiki/Cellulose" title="Cellulose">cellulose</a>, and fungi one of chitin, so most viruses can get inside these cells only after trauma to the cell wall.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 70  Nearly all plant viruses (such as tobacco mosaic virus) can also move directly from cell to cell, in the form of single-stranded nucleoprotein complexes, through pores called <a href="/wiki/Plasmodesma" title="Plasmodesma">plasmodesmata</a>.<a href="#cite_note-77">[77]</a> Bacteria, like plants, have strong cell walls that a virus must breach to infect the cell. Given that bacterial cell walls are much thinner than plant cell walls due to their much smaller size, some viruses have evolved mechanisms that inject their genome into the bacterial cell across the cell wall, while the viral capsid remains outside.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 71  Uncoating is a process in which the viral capsid is removed: This may be by degradation by viral enzymes or host enzymes or by simple dissociation; the end-result is the releasing of the viral genomic nucleic acid.<a href="#cite_note-Blaas_2016-78">[78]</a> <a href="/wiki/Viral_replication" title="Viral replication">Replication</a> of viruses involves primarily multiplication of the genome. Replication involves the synthesis of viral messenger RNA (mRNA) from "early" genes (with exceptions for positive-sense RNA viruses), viral <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">protein synthesis</a>, possible assembly of viral proteins, then viral genome replication mediated by early or regulatory protein expression. This may be followed, for complex viruses with larger genomes, by one or more further rounds of mRNA synthesis: "late" gene expression is, in general, of structural or virion proteins.<a href="#cite_note-Isomura_2013-79">[79]</a> Assembly – Following the structure-mediated self-assembly of the virus particles, some modification of the proteins often occurs. In viruses such as HIV, this modification (sometimes called maturation) occurs after the virus has been released from the host cell.<a href="#cite_note-Barman_2001-80">[80]</a> Release – Viruses can be <a href="/wiki/Viral_shedding" title="Viral shedding">released</a> from the host cell by <a href="/wiki/Lysis" title="Lysis">lysis</a>, a process that kills the cell by bursting its membrane and cell wall if present: this is a feature of many bacterial and some animal viruses. Some viruses undergo a <a href="/wiki/Lysogenic_cycle" title="Lysogenic cycle">lysogenic cycle</a> where the viral genome is incorporated by <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a> into a specific place in the host's chromosome. The viral genome is then known as a "<a href="/wiki/Provirus" title="Provirus">provirus</a>" or, in the case of bacteriophages a "<a href="/wiki/Prophage" title="Prophage">prophage</a>".<a href="#cite_note-Shors_2017-13">[13]</a>: 836  Whenever the host divides, the viral genome is also replicated. The viral genome is mostly silent within the host. At some point, the provirus or prophage may give rise to the active virus, which may lyse the host cells.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 243–259  Enveloped viruses (e.g., HIV) typically are released from the host cell by <a href="/wiki/Viral_shedding" title="Viral shedding">budding</a>. During this process, the virus acquires its envelope, which is a modified piece of the host's plasma or other, internal membrane.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 185–187  <div class="mw-heading mw-heading3"><h3 id="Genome_replication">Genome replication</h3></div> The genetic material within virus particles, and the method by which the material is replicated, varies considerably between different types of viruses. <dl><dt>DNA viruses</dt> <dd>The genome replication of most <a href="/wiki/DNA_virus" title="DNA virus">DNA viruses</a> takes place in the cell's <a href="/wiki/Cell_nucleus" title="Cell nucleus">nucleus</a>. If the cell has the appropriate receptor on its surface, these viruses enter the cell either by direct fusion with the cell membrane (e.g., herpesviruses) or—more usually—by receptor-mediated endocytosis. Most DNA viruses are entirely dependent on the host cell's DNA and RNA synthesising machinery and RNA processing machinery. Viruses with larger genomes may encode much of this machinery themselves. In eukaryotes, the viral genome must cross the cell's nuclear membrane to access this machinery, while in bacteria it need only enter the cell.<a href="#cite_note-Shors_2017-13">[13]</a>: 118 <a href="#cite_note-Collier_1998-26">[26]</a>: 78 </dd></dl> <dl><dt>RNA viruses</dt> <dd>Replication of <a href="/wiki/RNA_virus" title="RNA virus">RNA viruses</a> usually takes place in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>. RNA viruses can be placed into four different groups depending on their modes of replication. The <a href="/wiki/Sense_(molecular_biology)" title="Sense (molecular biology)">polarity</a> (whether or not it can be used directly by ribosomes to make proteins) of single-stranded RNA viruses largely determines the replicative mechanism; the other major criterion is whether the genetic material is single-stranded or double-stranded. All RNA viruses use their own <a href="/wiki/RNA_replicase" class="mw-redirect" title="RNA replicase">RNA replicase</a> enzymes to create copies of their genomes.<a href="#cite_note-Collier_1998-26">[26]</a>: 79 </dd></dl> <dl><dt>Reverse transcribing viruses</dt> <dd><a href="/wiki/Reverse_transcribing_viruses" class="mw-redirect" title="Reverse transcribing viruses">Reverse transcribing viruses</a> have ssRNA (<a href="/wiki/Retroviridae" class="mw-redirect" title="Retroviridae">Retroviridae</a>, <a href="/wiki/Metaviridae" title="Metaviridae">Metaviridae</a>, <a href="/wiki/Pseudoviridae" title="Pseudoviridae">Pseudoviridae</a>) or dsDNA (<a href="/wiki/Caulimoviridae" title="Caulimoviridae">Caulimoviridae</a>, and <a href="/wiki/Hepadnaviridae" title="Hepadnaviridae">Hepadnaviridae</a>) in their particles. Reverse transcribing viruses with RNA genomes (<a href="/wiki/Retrovirus" title="Retrovirus">retroviruses</a>) use a DNA intermediate to replicate, whereas those with DNA genomes (<a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">pararetroviruses</a>) use an RNA intermediate during genome replication. Both types use a <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a>, or RNA-dependent DNA polymerase enzyme, to carry out the nucleic acid conversion. Retroviruses integrate the DNA produced by <a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">reverse transcription</a> into the host genome as a provirus as a part of the replication process; pararetroviruses do not, although integrated genome copies of especially plant pararetroviruses can give rise to infectious virus.<a href="#cite_note-Staginnus_2006-81">[81]</a> They are susceptible to <a href="/wiki/Antiviral_drug" title="Antiviral drug">antiviral drugs</a> that inhibit the reverse transcriptase enzyme, e.g. <a href="/wiki/Zidovudine" title="Zidovudine">zidovudine</a> and <a href="/wiki/Lamivudine" title="Lamivudine">lamivudine</a>. An example of the first type is HIV, which is a retrovirus. Examples of the second type are the <a href="/wiki/Hepadnaviridae" title="Hepadnaviridae">Hepadnaviridae</a>, which includes Hepatitis B virus.<a href="#cite_note-Collier_1998-26">[26]</a>: 88–89 </dd></dl> <div class="mw-heading mw-heading3"><h3 id="Cytopathic_effects_on_the_host_cell">Cytopathic effects on the host cell</h3></div> The range of structural and biochemical effects that viruses have on the host cell is extensive.<a href="#cite_note-Collier_1998-26">[26]</a>: 115–146  These are called '<a href="/wiki/Cytopathic_effect" title="Cytopathic effect">cytopathic effects</a>'.<a href="#cite_note-Collier_1998-26">[26]</a>: 115  Most virus infections eventually result in the death of the host cell. The causes of death include cell lysis, alterations to the cell's surface membrane and <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>.<a href="#cite_note-Roulston_1999-82">[82]</a> Often cell death is caused by cessation of its normal activities because of suppression by virus-specific proteins, not all of which are components of the virus particle.<a href="#cite_note-Alwine_2008-83">[83]</a> The distinction between cytopathic and harmless is gradual. Some viruses, such as <a href="/wiki/Epstein%E2%80%93Barr_virus" title="Epstein–Barr virus">Epstein–Barr virus</a>, can cause cells to proliferate without causing malignancy,<a href="#cite_note-Barozzi_2007-84">[84]</a> while others, such as <a href="/wiki/Papillomavirus" class="mw-redirect" title="Papillomavirus">papillomaviruses</a>, are established causes of cancer.<a href="#cite_note-Subramanya_2008-85">[85]</a> <div class="mw-heading mw-heading3"><h3 id="Dormant_and_latent_infections">Dormant and latent infections</h3></div> Some viruses cause no apparent changes to the infected cell. Cells in which the virus is <a href="/wiki/Virus_latency" title="Virus latency">latent</a> and inactive show few signs of infection and often function normally.<a href="#cite_note-Sinclair_2008-86">[86]</a> This causes persistent infections and the virus is often dormant for many months or years. This is often the case with <a href="/wiki/Herpes_simplex" class="mw-redirect" title="Herpes simplex">herpes viruses</a>.<a href="#cite_note-Jordan_1984-87">[87]</a><a href="#cite_note-Sissons_2002-88">[88]</a> <div class="mw-heading mw-heading3"><h3 id="Host_range">Host range</h3></div> Viruses are by far the most abundant biological entities on Earth and they outnumber all the others put together.<a href="#cite_note-Crawford_2011-89">[89]</a> They infect all types of cellular life including animals, plants, <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteria</a> and <a href="/wiki/Mycovirus" title="Mycovirus">fungi</a>.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 49  Different types of viruses can infect only a limited range of hosts and many are species-specific. Some, such as <a href="/wiki/Smallpox_virus" class="mw-redirect" title="Smallpox virus">smallpox virus</a> for example, can infect only one species—in this case humans,<a href="#cite_note-Shors_2017-13">[13]</a>: 643  and are said to have a narrow <a href="/wiki/Host_specificity" class="mw-redirect" title="Host specificity">host range</a>. Other viruses, such as rabies virus, can infect different species of mammals and are said to have a broad range.<a href="#cite_note-Shors_2017-13">[13]</a>: 631  The viruses that infect plants are harmless to animals, and most viruses that infect other animals are harmless to humans.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 272  The host range of some bacteriophages is limited to a single <a href="/wiki/Strain_(biology)" title="Strain (biology)">strain</a> of bacteria and they can be used to trace the source of outbreaks of infections by a method called <a href="/wiki/Phage_typing" title="Phage typing">phage typing</a>.<a href="#cite_note-Baggesen_2010-90">[90]</a> The complete set of viruses in an organism or habitat is called the <a href="/wiki/Virome" title="Virome">virome</a>; for example, all human viruses constitute the <a href="/wiki/Human_virome" title="Human virome">human virome</a>.<a href="#cite_note-Parker_2016-91">[91]</a> <div class="mw-heading mw-heading3"><h3 id="Novel_viruses">Novel viruses</h3></div> A novel virus is one that has not previously been recorded. It can be a virus that is isolated from its <a href="/wiki/Natural_reservoir" title="Natural reservoir">natural reservoir</a> or isolated as the result of <a href="/wiki/Zoonosis" title="Zoonosis">spread to an animal or human host</a> where the virus had not been identified before. It can be an <a href="/wiki/Emergent_virus" title="Emergent virus">emergent virus</a>, one that represents a new virus, but it can also be an extant virus that has not been <a href="/wiki/Virus_classification" title="Virus classification">previously identified</a>.<a href="#cite_note-Woolhouse_2012-92">[92]</a> The <a href="/wiki/SARS-CoV-2" title="SARS-CoV-2">SARS-CoV-2</a> coronavirus that caused the COVID-19 pandemic is an example of a novel virus.<a href="#cite_note-Ashour_2020-93">[93]</a> <div class="mw-heading mw-heading2"><h2 id="Classification">Classification</h2></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> Classification seeks to describe the diversity of viruses by naming and grouping them on the basis of similarities. 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.<a href="#cite_note-Lwoff_1962-94">[94]</a> 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.<a href="#cite_note-Lwoff_1962b-95">[95]</a> 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.<a href="#cite_note-Fauquet_2005-96">[96]</a> 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.<a href="#cite_note-97">[97]</a> Additionally, some species within the same genus are grouped into a genogroup.<a href="#cite_note-Khan_2021-98">[98]</a><a href="#cite_note-Eberle_2012-99">[99]</a> <div class="mw-heading mw-heading3"><h3 id="ICTV_classification">ICTV classification</h3></div> 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.<a href="#cite_note-100">[100]</a> Only a small part of the total diversity of viruses has been studied.<a href="#cite_note-101">[101]</a> As of 2022, 6 realms, 10 kingdoms, 17 phyla, 2 subphyla, 40 classes, 72 orders, 8 suborders, <a href="/wiki/List_of_virus_families_and_subfamilies" title="List of virus families and subfamilies">264 families, 182 subfamilies</a>, <a href="/wiki/List_of_virus_genera" title="List of virus genera">2,818 genera, 84 subgenera</a>, and <a href="/wiki/List_of_virus_species" title="List of virus species">11,273 species</a> of viruses have been defined by the ICTV.<a href="#cite_note-ictv_2022-7">[7]</a> The general taxonomic structure of taxon ranges and the suffixes used in taxonomic names are shown hereafter. As of 2022, the ranks of subrealm, subkingdom, and subclass are unused, whereas all other ranks are in use.<a href="#cite_note-ictv_2022-7">[7]</a> <dl><dd><a href="/wiki/Realm_(virology)" title="Realm (virology)">Realm</a> (-viria) <dl><dd>Subrealm (-vira) <dl><dd><a href="/wiki/Kingdom_(biology)" title="Kingdom (biology)">Kingdom</a> (-virae) <dl><dd>Subkingdom (-virites) <dl><dd><a href="/wiki/Phylum_(biology)" class="mw-redirect" title="Phylum (biology)">Phylum</a> (-viricota) <dl><dd>Subphylum (-viricotina) <dl><dd><a href="/wiki/Class_(biology)" title="Class (biology)">Class</a> (-viricetes) <dl><dd>Subclass (-viricetidae) <dl><dd><a href="/wiki/Order_(biology)" title="Order (biology)">Order</a> (-virales) <dl><dd>Suborder (-virineae) <dl><dd><a href="/wiki/Family_(biology)" title="Family (biology)">Family</a> (-viridae) <dl><dd>Subfamily (-virinae) <dl><dd><a href="/wiki/Genus" title="Genus">Genus</a> (-virus) <dl><dd>Subgenus (-virus) <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></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> 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.<a href="#cite_note-Temin_1972-102">[102]</a><a href="#cite_note-Baltimore_1974-103">[103]</a> The ICTV classification system is used in conjunction with the Baltimore classification system in modern virus classification.<a href="#cite_note-van_Regenmortel_2004-104">[104]</a><a href="#cite_note-Mayo_1999-105">[105]</a><a href="#cite_note-de_Villiers_2004-106">[106]</a> 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: <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> <div class="mw-heading mw-heading2"><h2 id="Role_in_human_disease">Role in human disease</h2></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Viral_disease" title="Viral disease">Viral disease</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Viral_infections_and_involved_species.svg" class="mw-file-description"><img alt="A photograph of the upper body of a man labelled with the names of viruses that infect the different parts" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c2/Viral_infections_and_involved_species.svg/220px-Viral_infections_and_involved_species.svg.png" decoding="async" width="220" height="210" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c2/Viral_infections_and_involved_species.svg/330px-Viral_infections_and_involved_species.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c2/Viral_infections_and_involved_species.svg/440px-Viral_infections_and_involved_species.svg.png 2x" data-file-width="1421" data-file-height="1358" /></a><figcaption>Overview of the main types of viral infection and the most notable species involved<a href="#cite_note-Fisher_2007-107">[107]</a></figcaption></figure> Examples of common human diseases caused by viruses include the <a href="/wiki/Common_cold" title="Common cold">common cold</a>, <a href="/wiki/Influenza" title="Influenza">influenza</a>, <a href="/wiki/Chickenpox" title="Chickenpox">chickenpox</a>, and <a href="/wiki/Cold_sores" class="mw-redirect" title="Cold sores">cold sores</a>. Many serious diseases such as <a href="/wiki/Rabies" title="Rabies">rabies</a>, <a href="/wiki/Ebola_virus_disease" class="mw-redirect" title="Ebola virus disease">Ebola virus disease</a>, <a href="/wiki/HIV/AIDS" title="HIV/AIDS">AIDS (HIV)</a>, <a href="/wiki/Avian_influenza" title="Avian influenza">avian influenza</a>, and <a href="/wiki/SARS" title="SARS">SARS</a> are caused by viruses. The relative ability of viruses to cause disease is described in terms of <a href="/wiki/Virulence" title="Virulence">virulence</a>. Other diseases are under investigation to discover if they have a virus as the causative agent, such as the possible connection between <a href="/wiki/Human_herpesvirus_6" title="Human herpesvirus 6">human herpesvirus 6</a> (HHV6) and neurological diseases such as <a href="/wiki/Multiple_sclerosis" title="Multiple sclerosis">multiple sclerosis</a> and <a href="/wiki/Chronic_fatigue_syndrome" class="mw-redirect" title="Chronic fatigue syndrome">chronic fatigue syndrome</a>.<a href="#cite_note-Komaroff_2006-108">[108]</a> There is controversy over whether the <a href="/wiki/Bornavirus" class="mw-redirect" title="Bornavirus">bornavirus</a>, previously thought to cause <a href="/wiki/Neurology" title="Neurology">neurological</a> diseases in horses, could be responsible for <a href="/wiki/Psychiatry" title="Psychiatry">psychiatric</a> illnesses in humans.<a href="#cite_note-Chen_1999-109">[109]</a> Viruses have different mechanisms by which they produce disease in an organism, which depends largely on the viral species. Mechanisms at the cellular level primarily include cell lysis, the breaking open and subsequent death of the cell. In <a href="/wiki/Multicellular_organism" title="Multicellular organism">multicellular organisms</a>, if enough cells die, the whole organism will start to suffer the effects. Although viruses cause disruption of healthy <a href="/wiki/Homeostasis" title="Homeostasis">homeostasis</a>, resulting in disease, they may exist relatively harmlessly within an organism. An example would include the ability of the <a href="/wiki/Herpes_simplex_virus" title="Herpes simplex virus">herpes simplex virus</a>, which causes cold sores, to remain in a dormant state within the human body. This is called latency<a href="#cite_note-Margolis_2007-110">[110]</a> and is a characteristic of the herpes viruses, including Epstein–Barr virus, which causes glandular fever, and <a href="/wiki/Varicella_zoster_virus" title="Varicella zoster virus">varicella zoster virus</a>, which causes chickenpox and <a href="/wiki/Shingles" title="Shingles">shingles</a>. Most people have been infected with at least one of these types of herpes virus.<a href="#cite_note-111">[111]</a> These latent viruses might sometimes be beneficial, as the presence of the virus can increase immunity against bacterial pathogens, such as <a href="/wiki/Yersinia_pestis" title="Yersinia pestis">Yersinia pestis</a>.<a href="#cite_note-112">[112]</a> Some viruses can cause lifelong or <a href="/wiki/Chronic_(medical)" class="mw-redirect" title="Chronic (medical)">chronic</a> infections, where the viruses continue to replicate in the body despite the host's defence mechanisms.<a href="#cite_note-Bertoletti_2007-113">[113]</a> This is common in hepatitis B virus and hepatitis C virus infections. People chronically infected are known as carriers, as they serve as reservoirs of infectious virus.<a href="#cite_note-Rodrigues_2001-114">[114]</a> In populations with a high proportion of carriers, the disease is said to be <a href="/wiki/Endemic_(epidemiology)" title="Endemic (epidemiology)">endemic</a>.<a href="#cite_note-Nguyen_2007-115">[115]</a> <div class="mw-heading mw-heading3"><h3 id="Epidemiology">Epidemiology</h3></div> Viral <a href="/wiki/Epidemiology" title="Epidemiology">epidemiology</a> is the branch of medical science that deals with the transmission and control of virus infections in humans. Transmission of viruses can be vertical, which means from mother to child, or horizontal, which means from person to person. Examples of <a href="/wiki/Vertical_transmission" title="Vertical transmission">vertical transmission</a> include hepatitis B virus and HIV, where the baby is born already infected with the virus.<a href="#cite_note-Fowler_2007-116">[116]</a> Another, more rare, example is the <a href="/wiki/Varicella_zoster_virus" title="Varicella zoster virus">varicella zoster virus</a>, which, although causing relatively mild infections in children and adults, can be fatal to the foetus and newborn baby.<a href="#cite_note-Sauerbrei_2000-117">[117]</a> <a href="/wiki/Horizontal_transmission" title="Horizontal transmission">Horizontal transmission</a> is the most common mechanism of spread of viruses in populations.<a href="#cite_note-Antonovics_2007-118">[118]</a> Horizontal transmission can occur when body fluids are exchanged during sexual activity, by exchange of saliva or when contaminated food or water is ingested. It can also occur when <a href="/wiki/Aerosol" title="Aerosol">aerosols</a> containing viruses are inhaled or by insect <a href="/wiki/Vector_(epidemiology)" class="mw-redirect" title="Vector (epidemiology)">vectors</a> such as when infected mosquitoes penetrate the skin of a host.<a href="#cite_note-Antonovics_2007-118">[118]</a> Most types of viruses are restricted to just one or two of these mechanisms and they are referred to as "respiratory viruses" or "enteric viruses" and so forth. The rate or speed of transmission of viral infections depends on factors that include population density, the number of susceptible individuals, (i.e., those not immune),<a href="#cite_note-Garnett_2005-119">[119]</a> the quality of healthcare and the weather.<a href="#cite_note-Platonov_2006-120">[120]</a> Epidemiology is used to break the chain of infection in populations during outbreaks of <a href="/wiki/Viral_disease" title="Viral disease">viral diseases</a>.<a href="#cite_note-Shors_2017-13">[13]</a>: 264  Control measures are used that are based on knowledge of how the virus is transmitted. It is important to find the source, or sources, of the outbreak and to identify the virus. Once the virus has been identified, the chain of transmission can sometimes be broken by vaccines. When vaccines are not available, sanitation and disinfection can be effective. Often, infected people are isolated from the rest of the community, and those that have been exposed to the virus are placed in <a href="/wiki/Quarantine" title="Quarantine">quarantine</a>.<a href="#cite_note-Shors_2017-13">[13]</a>: 894  To control the <a href="/wiki/2001_United_Kingdom_foot-and-mouth_outbreak" title="2001 United Kingdom foot-and-mouth outbreak">outbreak</a> of <a href="/wiki/Foot-and-mouth_disease" title="Foot-and-mouth disease">foot-and-mouth disease</a> in cattle in Britain in 2001, thousands of cattle were slaughtered.<a href="#cite_note-Jewell_2009-121">[121]</a> Most viral infections of humans and other animals have <a href="/wiki/Incubation_period" title="Incubation period">incubation periods</a> during which the infection causes no signs or symptoms.<a href="#cite_note-Shors_2017-13">[13]</a>: 170  Incubation periods for viral diseases range from a few days to weeks, but are known for most infections.<a href="#cite_note-Shors_2017-13">[13]</a>: 170–172  Somewhat overlapping, but mainly following the incubation period, there is a period of communicability—a time when an infected individual or animal is contagious and can infect another person or animal.<a href="#cite_note-Shors_2017-13">[13]</a>: 170–172  This, too, is known for many viral infections, and knowledge of the length of both periods is important in the control of outbreaks.<a href="#cite_note-Shors_2017-13">[13]</a>: 272  When outbreaks cause an unusually high proportion of cases in a population, community, or region, they are called epidemics. If outbreaks spread worldwide, they are called <a href="/wiki/Pandemic" title="Pandemic">pandemics</a>.<a href="#cite_note-Shors_2017-13">[13]</a>: 891  <div class="mw-heading mw-heading3"><h3 id="Epidemics_and_pandemics">Epidemics and pandemics</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/1918_flu_pandemic" class="mw-redirect" title="1918 flu pandemic">1918 flu pandemic</a>, <a href="/wiki/AIDS" class="mw-redirect" title="AIDS">AIDS</a>, <a href="/wiki/Ebola_virus_disease" class="mw-redirect" title="Ebola virus disease">Ebola virus disease</a>, and <a href="/wiki/COVID-19_pandemic" title="COVID-19 pandemic">COVID-19 pandemic</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/List_of_epidemics" class="mw-redirect" title="List of epidemics">List of epidemics</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Reconstructed_Spanish_Flu_Virus.jpg" class="mw-file-description"><img alt="An electron micrograph of the virus that caused Spanish influenza" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Reconstructed_Spanish_Flu_Virus.jpg/220px-Reconstructed_Spanish_Flu_Virus.jpg" decoding="async" width="220" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Reconstructed_Spanish_Flu_Virus.jpg/330px-Reconstructed_Spanish_Flu_Virus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ee/Reconstructed_Spanish_Flu_Virus.jpg/440px-Reconstructed_Spanish_Flu_Virus.jpg 2x" data-file-width="2126" data-file-height="1312" /></a><figcaption><a href="/wiki/Transmission_electron_microscopy" title="Transmission electron microscopy">Transmission electron microscope</a> image of a recreated 1918 influenza virus</figcaption></figure> A <a href="/wiki/Pandemic" title="Pandemic">pandemic</a> is a worldwide <a href="/wiki/Epidemic" title="Epidemic">epidemic</a>. The <a href="/wiki/1918_flu_pandemic" class="mw-redirect" title="1918 flu pandemic">1918 flu pandemic</a>, which lasted until 1919, was a <a href="/wiki/Pandemic_Severity_Index" class="mw-redirect" title="Pandemic Severity Index">category 5</a> influenza pandemic caused by an unusually severe and deadly influenza A virus. The victims were often healthy young adults, in contrast to most influenza outbreaks, which predominantly affect juvenile, elderly, or otherwise-weakened patients.<a href="#cite_note-Collier_1998-26">[26]</a>: 409–415  Older estimates say it killed 40–50 million people,<a href="#cite_note-Patterson1-122">[122]</a> while more recent research suggests that it may have killed as many as 100 million people, or 5% of the world's population in 1918.<a href="#cite_note-Johnson_2002-123">[123]</a> Although viral pandemics are rare events, HIV—which evolved from viruses found in monkeys and chimpanzees—has been pandemic since at least the 1980s.<a href="#cite_note-Eisinger_2018-124">[124]</a> During the 20th century there were four pandemics caused by influenza virus and those that occurred in 1918, 1957 and 1968 were severe.<a href="#cite_note-Qin_2018-125">[125]</a> Most researchers believe that HIV originated in <a href="/wiki/Sub-Saharan_Africa" title="Sub-Saharan Africa">sub-Saharan Africa</a> during the 20th century;<a href="#cite_note-Gao_1999-126">[126]</a> it is now a pandemic, with an estimated 37.9 million people now living with the disease worldwide.<a href="#cite_note-UNAIDS_DATA_2019-127">[127]</a> There were about 770,000 deaths from AIDS in 2018.<a href="#cite_note-UNAIDS_resources-128">[128]</a> The <a href="/wiki/Joint_United_Nations_Programme_on_HIV/AIDS" title="Joint United Nations Programme on HIV/AIDS">Joint United Nations Programme on HIV/AIDS</a> (UNAIDS) and the <a href="/wiki/World_Health_Organization" title="World Health Organization">World Health Organization</a> (WHO) estimate that AIDS has killed more than 25 million people since it was first recognised on 5 June 1981, making it one of the most destructive epidemics in recorded history.<a href="#cite_note-Mawar_2005-129">[129]</a> In 2007 there were 2.7 million new HIV infections and 2 million HIV-related deaths.<a href="#cite_note-130">[130]</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237032888/mw-parser-output/.tmulti"><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:154px;max-width:154px"><div class="trow"><div class="tsingle" style="width:152px;max-width:152px"><div class="thumbimage"><a href="/wiki/File:Ebola_virus_em.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Ebola_virus_em.png/150px-Ebola_virus_em.png" decoding="async" width="150" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Ebola_virus_em.png/225px-Ebola_virus_em.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3f/Ebola_virus_em.png/300px-Ebola_virus_em.png 2x" data-file-width="2043" data-file-height="2887" /></a></div></div></div><div class="trow"><div class="tsingle" style="width:152px;max-width:152px"><div class="thumbimage"><a href="/wiki/File:Marburg_virus.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Marburg_virus.jpg/150px-Marburg_virus.jpg" decoding="async" width="150" height="102" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Marburg_virus.jpg/225px-Marburg_virus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Marburg_virus.jpg/300px-Marburg_virus.jpg 2x" data-file-width="700" data-file-height="474" /></a></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">Ebola (top) and Marburg viruses (bottom)</div></div></div></div> Several highly lethal viral pathogens are members of the <a href="/wiki/Filoviridae" title="Filoviridae">Filoviridae</a>. Filoviruses are filament-like viruses that cause <a href="/wiki/Viral_hemorrhagic_fever" title="Viral hemorrhagic fever">viral hemorrhagic fever</a>, and include <a href="/wiki/Ebolavirus" title="Ebolavirus">ebolaviruses</a> and <a href="/wiki/Marburgvirus" title="Marburgvirus">marburgviruses</a>. <a href="/wiki/Marburg_virus" title="Marburg virus">Marburg virus</a>, first discovered in 1967, attracted widespread press attention in April 2005 for an outbreak in <a href="/wiki/Angola" title="Angola">Angola</a>.<a href="#cite_note-Towner_2006-131">[131]</a> <a href="/wiki/Ebola_virus_disease" class="mw-redirect" title="Ebola virus disease">Ebola virus disease</a> has also caused <a href="/wiki/List_of_Ebola_outbreaks" title="List of Ebola outbreaks">intermittent outbreaks</a> with high mortality rates since 1976 when it was first identified. The worst and most recent one is the 2013–2016 <a href="/wiki/Ebola_virus_epidemic_in_West_Africa" class="mw-redirect" title="Ebola virus epidemic in West Africa">West Africa epidemic</a>.<a href="#cite_note-132">[132]</a> Except for smallpox, most pandemics are caused by newly evolved viruses. These <a href="/wiki/Emergent_virus" title="Emergent virus">"emergent"</a> viruses are usually mutants of less harmful viruses that have circulated previously either in humans or other animals.<a href="#cite_note-133">[133]</a> Severe acute respiratory syndrome (<a href="/wiki/SARS" title="SARS">SARS</a>) and <a href="/wiki/Middle_East_respiratory_syndrome" class="mw-redirect" title="Middle East respiratory syndrome">Middle East respiratory syndrome</a> (MERS) are caused by new types of <a href="/wiki/Coronavirus" title="Coronavirus">coronaviruses</a>. Other coronaviruses are known to cause mild infections in humans,<a href="#cite_note-Weiss_2011-134">[134]</a> so the virulence and rapid spread of SARS infections—that by July 2003 had caused around 8,000 cases and 800 deaths—was unexpected and most countries were not prepared.<a href="#cite_note-Wong_2017-135">[135]</a> A related coronavirus, <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 (SARS-Cov-2)</a>, thought to have originated in bats, emerged in <a href="/wiki/Wuhan" title="Wuhan">Wuhan</a>, China in November 2019 and spread rapidly around the world. Infections with the virus caused the <a href="/wiki/COVID-19_pandemic" title="COVID-19 pandemic">COVID-19 pandemic</a> that started in 2020.<a href="#cite_note-Ashour_2020-93">[93]</a><a href="#cite_note-Deng_2020-136">[136]</a><a href="#cite_note-Han_2020-137">[137]</a> Unprecedented restrictions in peacetime were placed on international travel,<a href="#cite_note-138">[138]</a> and <a href="/wiki/Curfews" class="mw-redirect" title="Curfews">curfews</a> were imposed in several major cities worldwide in response to the pandemic.<a href="#cite_note-139">[139]</a> <div class="mw-heading mw-heading3"><h3 id="Cancer">Cancer</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Oncovirus" title="Oncovirus">Oncovirus</a></div> Viruses are an established cause of cancer in humans and other species. Viral cancers occur only in a minority of infected persons (or animals). Cancer viruses come from a range of virus families, including both RNA and DNA viruses, and so there is no single type of "<a href="/wiki/Oncovirus" title="Oncovirus">oncovirus</a>" (an obsolete term originally used for acutely transforming retroviruses). The development of cancer is determined by a variety of factors such as host immunity<a href="#cite_note-Einstein_2009-140">[140]</a> and mutations in the host.<a href="#cite_note-Shuda_2008-141">[141]</a> Viruses accepted to cause human cancers include some genotypes of <a href="/wiki/Human_papillomavirus" class="mw-redirect" title="Human papillomavirus">human papillomavirus</a>, <a href="/wiki/Hepatitis_B_virus" title="Hepatitis B virus">hepatitis B virus</a>, <a href="/wiki/Hepatitis_C_virus" title="Hepatitis C virus">hepatitis C virus</a>, <a href="/wiki/Epstein%E2%80%93Barr_virus" title="Epstein–Barr virus">Epstein–Barr virus</a>, <a href="/wiki/Kaposi%27s_sarcoma-associated_herpesvirus" title="Kaposi's sarcoma-associated herpesvirus">Kaposi's sarcoma-associated herpesvirus</a> and <a href="/wiki/Human_T-lymphotropic_virus" class="mw-redirect" title="Human T-lymphotropic virus">human T-lymphotropic virus</a>. The most recently discovered human cancer virus is a polyomavirus (<a href="/wiki/Merkel_cell_polyomavirus" title="Merkel cell polyomavirus">Merkel cell polyomavirus</a>) that causes most cases of a rare form of skin cancer called <a href="/wiki/Merkel_cell_carcinoma" class="mw-redirect" title="Merkel cell carcinoma">Merkel cell carcinoma</a>.<a href="#cite_note-Pulitzer_2009-142">[142]</a> Hepatitis viruses can develop into a chronic viral infection that leads to <a href="/wiki/Hepatocellular_carcinoma" title="Hepatocellular carcinoma">liver cancer</a>.<a href="#cite_note-143">[143]</a><a href="#cite_note-144">[144]</a> Infection by human T-lymphotropic virus can lead to <a href="/wiki/Tropical_spastic_paraparesis" title="Tropical spastic paraparesis">tropical spastic paraparesis</a> and <a href="/wiki/Adult_T-cell_leukemia" class="mw-redirect" title="Adult T-cell leukemia">adult T-cell leukaemia</a>.<a href="#cite_note-145">[145]</a> Human papillomaviruses are an established cause of cancers of <a href="/wiki/Cervix" title="Cervix">cervix</a>, skin, <a href="/wiki/Anus" title="Anus">anus</a>, and <a href="/wiki/Penis" title="Penis">penis</a>.<a href="#cite_note-146">[146]</a> Within the <a href="/wiki/Herpesviridae" title="Herpesviridae">Herpesviridae</a>, <a href="/wiki/Kaposi%27s_sarcoma-associated_herpesvirus" title="Kaposi's sarcoma-associated herpesvirus">Kaposi's sarcoma-associated herpesvirus</a> causes <a href="/wiki/Kaposi%27s_sarcoma" title="Kaposi's sarcoma">Kaposi's sarcoma</a> and <a href="/wiki/Primary_effusion_lymphoma" title="Primary effusion lymphoma">body-cavity lymphoma</a>, and Epstein–Barr virus causes <a href="/wiki/Burkitt%27s_lymphoma" class="mw-redirect" title="Burkitt's lymphoma">Burkitt's lymphoma</a>, <a href="/wiki/Hodgkin%27s_lymphoma" class="mw-redirect" title="Hodgkin's lymphoma">Hodgkin's lymphoma</a>, <a href="/wiki/B_cell" title="B cell">B</a> <a href="/wiki/Lymphoproliferative_disorders" title="Lymphoproliferative disorders">lymphoproliferative disorder</a>, and <a href="/wiki/Nasopharyngeal_carcinoma" title="Nasopharyngeal carcinoma">nasopharyngeal carcinoma</a>.<a href="#cite_note-147">[147]</a> Merkel cell polyomavirus closely related to <a href="/wiki/SV40" title="SV40">SV40</a> and mouse polyomaviruses that have been used as animal models for cancer viruses for over 50 years.<a href="#cite_note-Zur_Hausen_2008-148">[148]</a> <div class="mw-heading mw-heading3"><h3 id="Host_defence_mechanisms">Host defence mechanisms</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Immune_system" title="Immune system">Immune system</a></div> The body's first line of defence against viruses is the <a href="/wiki/Innate_immune_system" title="Innate immune system">innate immune system</a>. This comprises cells and other mechanisms that defend the host from infection in a non-specific manner. This means that the cells of the innate system recognise, and respond to, pathogens in a generic way, but, unlike the <a href="/wiki/Adaptive_immune_system" title="Adaptive immune system">adaptive immune system</a>, it does not confer long-lasting or protective immunity to the host.<a href="#cite_note-Alberts_2002-149">[149]</a> <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a> is an important innate defence against viruses.<a href="#cite_note-Ding_2007-150">[150]</a> Many viruses have a replication strategy that involves double-stranded RNA (dsRNA). When such a virus infects a cell, it releases its RNA molecule or molecules, which immediately bind to a protein complex called a <a href="/wiki/Dicer" title="Dicer">dicer</a> that cuts the RNA into smaller pieces. A biochemical pathway—the <a href="/wiki/RNA-induced_silencing_complex" title="RNA-induced silencing complex">RISC complex</a>—is activated, which ensures cell survival by degrading the viral mRNA. Rotaviruses have evolved to avoid this defence mechanism by not uncoating fully inside the cell, and releasing newly produced mRNA through pores in the particle's inner capsid. Their genomic dsRNA remains protected inside the core of the virion.<a href="#cite_note-Patton_2005-151">[151]</a><a href="#cite_note-Jayaram_2004-152">[152]</a> When the <a href="/wiki/Adaptive_immune_system" title="Adaptive immune system">adaptive immune system</a> of a <a href="/wiki/Vertebrate" title="Vertebrate">vertebrate</a> encounters a virus, it produces specific <a href="/wiki/Antibodies" class="mw-redirect" title="Antibodies">antibodies</a> that bind to the virus and often render it non-infectious. This is called <a href="/wiki/Humoral_immunity" title="Humoral immunity">humoral immunity</a>. Two types of antibodies are important. The first, called <a href="/wiki/Immunoglobulin_M" title="Immunoglobulin M">IgM</a>, is highly effective at neutralising viruses but is produced by the cells of the immune system only for a few weeks. The second, called <a href="/wiki/Immunoglobulin_G" title="Immunoglobulin G">IgG</a>, is produced indefinitely. The presence of IgM in the blood of the host is used to test for acute infection, whereas IgG indicates an infection sometime in the past.<a href="#cite_note-Greer_1995-153">[153]</a> IgG antibody is measured when tests for <a href="/wiki/Immunity_(medical)" class="mw-redirect" title="Immunity (medical)">immunity</a> are carried out.<a href="#cite_note-Matter_1997-154">[154]</a> Antibodies can continue to be an effective defence mechanism even after viruses have managed to gain entry to the host cell. A protein that is in cells, called <a href="/wiki/TRIM21" title="TRIM21">TRIM21</a>, can attach to the antibodies on the surface of the virus particle. This primes the subsequent destruction of the virus by the enzymes of the cell's <a href="/wiki/Proteosome" class="mw-redirect" title="Proteosome">proteosome</a> system.<a href="#cite_note-Mallery_2010-155">[155]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Rotavirus_with_antibody.jpg" class="mw-file-description"><img alt="Two spherical rotavirus particles, one is coated with antibody which looks like many small birds, regularly spaced on the surface of the virus" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Rotavirus_with_antibody.jpg/220px-Rotavirus_with_antibody.jpg" decoding="async" width="220" height="119" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Rotavirus_with_antibody.jpg/330px-Rotavirus_with_antibody.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/98/Rotavirus_with_antibody.jpg/440px-Rotavirus_with_antibody.jpg 2x" data-file-width="1072" data-file-height="579" /></a><figcaption>Two <a href="/wiki/Rotavirus" title="Rotavirus">rotaviruses</a>: the one on the right is coated with antibodies that prevent its attachment to cells and infecting them.</figcaption></figure> A second defence of vertebrates against viruses is called <a href="/wiki/Cell-mediated_immunity" title="Cell-mediated immunity">cell-mediated immunity</a> and involves immune cells known as <a href="/wiki/T_cells" class="mw-redirect" title="T cells">T cells</a>. The body's cells constantly display short fragments of their proteins on the cell's surface, and, if a T cell recognises a suspicious viral fragment there, the host cell is destroyed by 'killer T' cells and the virus-specific T-cells proliferate. Cells such as the <a href="/wiki/Macrophage" title="Macrophage">macrophage</a> are specialists at this <a href="/wiki/Antigen_presentation" title="Antigen presentation">antigen presentation</a>.<a href="#cite_note-Cascalho_2007-156">[156]</a> The production of <a href="/wiki/Interferon" title="Interferon">interferon</a> is an important host defence mechanism. This is a hormone produced by the body when viruses are present. Its role in immunity is complex; it eventually stops the viruses from reproducing by killing the infected cell and its close neighbours.<a href="#cite_note-Le_Page_2000-157">[157]</a> Not all virus infections produce a protective immune response in this way. HIV evades the immune system by constantly changing the amino acid sequence of the proteins on the surface of the virion. This is known as "escape mutation" as the viral epitopes escape recognition by the host immune response. These persistent viruses evade immune control by sequestration, blockade of <a href="/wiki/Antigen_presentation" title="Antigen presentation">antigen presentation</a>, <a href="/wiki/Cytokine" title="Cytokine">cytokine</a> resistance, evasion of <a href="/wiki/Natural_killer_cell" title="Natural killer cell">natural killer cell</a> activities, escape from <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, and <a href="/wiki/Antigenic_shift" title="Antigenic shift">antigenic shift</a>.<a href="#cite_note-Hilleman_2004-158">[158]</a> Other viruses, called '<a href="/wiki/Neurotropic_virus" title="Neurotropic virus">neurotropic viruses</a>', are disseminated by neural spread where the immune system may be unable to reach them due to <a href="/wiki/Immune_privilege" title="Immune privilege">immune privilege</a>.<a href="#cite_note-Libbey_2014-159">[159]</a> <div class="mw-heading mw-heading3"><h3 id="Prevention_and_treatment">Prevention and treatment</h3></div> Because viruses use vital metabolic pathways within host cells to replicate, they are difficult to eliminate without using drugs that cause toxic effects to host cells in general. The most effective medical approaches to viral diseases are <a href="/wiki/Vaccination" title="Vaccination">vaccinations</a> to provide immunity to infection, and <a href="/wiki/Antiviral_drugs" class="mw-redirect" title="Antiviral drugs">antiviral drugs</a> that selectively interfere with viral replication. <div class="mw-heading mw-heading4"><h4 id="Vaccines">Vaccines</h4></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Vaccination" title="Vaccination">Vaccination</a></div> Vaccination is a cheap and effective way of preventing infections by viruses. Vaccines were used to prevent viral infections long before the discovery of the actual viruses. Their use has resulted in a dramatic decline in morbidity (illness) and mortality (death) associated with viral infections such as <a href="/wiki/Polio" title="Polio">polio</a>, <a href="/wiki/Measles" title="Measles">measles</a>, <a href="/wiki/Mumps" title="Mumps">mumps</a> and <a href="/wiki/Rubella" title="Rubella">rubella</a>.<a href="#cite_note-Asaria_2006-160">[160]</a> <a href="/wiki/Smallpox" title="Smallpox">Smallpox</a> infections have been eradicated.<a href="#cite_note-Lane_2006-161">[161]</a> Vaccines are available to prevent over thirteen viral infections of humans,<a href="#cite_note-Arvin_2006-162">[162]</a> and more are used to prevent viral infections of animals.<a href="#cite_note-Pastoret_2007-163">[163]</a> Vaccines can consist of live-attenuated or killed viruses, viral proteins (<a href="/wiki/Antigens" class="mw-redirect" title="Antigens">antigens</a>), or <a href="/wiki/RNA_vaccine" class="mw-redirect" title="RNA vaccine">RNA</a>.<a href="#cite_note-Palese_2006-164">[164]</a><a href="#cite_note-Anand_2021-165">[165]</a> Live vaccines contain weakened forms of the virus, which do not cause the disease but, nonetheless, confer immunity. Such viruses are called attenuated. Live vaccines can be dangerous when given to people with a weak immunity (who are described as <a href="/wiki/Immunocompromised" class="mw-redirect" title="Immunocompromised">immunocompromised</a>), because in these people, the weakened virus can cause the original disease.<a href="#cite_note-Thomssen_1975-166">[166]</a> Biotechnology and genetic engineering techniques are used to produce subunit vaccines. These vaccines use only the capsid proteins of the virus. Hepatitis B vaccine is an example of this type of vaccine.<a href="#cite_note-McLean_1986-167">[167]</a> Subunit vaccines are safe for <a href="/wiki/Immunocompromised" class="mw-redirect" title="Immunocompromised">immunocompromised</a> patients because they cannot cause the disease.<a href="#cite_note-Casswall_2005-168">[168]</a> The <a href="/wiki/Yellow_fever_vaccine" title="Yellow fever vaccine">yellow fever virus vaccine</a>, a live-attenuated strain called 17D, is probably the safest and most effective vaccine ever generated.<a href="#cite_note-Barnett_2008-169">[169]</a> <div class="mw-heading mw-heading4"><h4 id="Antiviral_drugs">Antiviral drugs</h4></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Antiviral_drug" title="Antiviral drug">Antiviral drug</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Guanosine_aciclovir_comparison.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Guanosine_aciclovir_comparison.svg/170px-Guanosine_aciclovir_comparison.svg.png" decoding="async" width="170" height="205" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Guanosine_aciclovir_comparison.svg/255px-Guanosine_aciclovir_comparison.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Guanosine_aciclovir_comparison.svg/340px-Guanosine_aciclovir_comparison.svg.png 2x" data-file-width="512" data-file-height="618" /></a><figcaption>The structure of the DNA base <a href="/wiki/Guanosine" title="Guanosine">guanosine</a> and the antiviral drug <a href="/wiki/Acyclovir" class="mw-redirect" title="Acyclovir">acyclovir</a></figcaption></figure> Antiviral drugs are often <a href="/wiki/Nucleoside_analogues" class="mw-redirect" title="Nucleoside analogues">nucleoside analogues</a> (fake DNA building-blocks), which viruses mistakenly incorporate into their genomes during replication.<a href="#cite_note-De_Clercq_2016-170">[170]</a> The life-cycle of the virus is then halted because the newly synthesised DNA is inactive. This is because these analogues lack the <a href="/wiki/Hydroxyl" class="mw-redirect" title="Hydroxyl">hydroxyl</a> groups, which, along with <a href="/wiki/Phosphorus" title="Phosphorus">phosphorus</a> atoms, link together to form the strong "backbone" of the DNA molecule. This is called DNA <a href="/wiki/Chain_termination" title="Chain termination">chain termination</a>.<a href="#cite_note-Magden_2005-171">[171]</a> Examples of nucleoside analogues are <a href="/wiki/Aciclovir" title="Aciclovir">aciclovir</a> for Herpes simplex virus infections and <a href="/wiki/Lamivudine" title="Lamivudine">lamivudine</a> for HIV and hepatitis B virus infections. Aciclovir is one of the oldest and most frequently prescribed antiviral drugs.<a href="#cite_note-Mindel_1983-172">[172]</a> Other antiviral drugs in use target different stages of the viral life cycle. HIV is dependent on a proteolytic enzyme called the <a href="/wiki/HIV-1_protease" title="HIV-1 protease">HIV-1 protease</a> for it to become fully infectious. There is a large class of drugs called <a href="/wiki/Protease_inhibitors" class="mw-redirect" title="Protease inhibitors">protease inhibitors</a> that inactivate this enzyme.<a href="#cite_note-Palmisano_2011-173">[173]</a> There are around thirteen classes of antiviral drugs each targeting different viruses or stages of viral replication.<a href="#cite_note-De_Clercq_2016-170">[170]</a> Hepatitis C is caused by an RNA virus. In 80% of people infected, the disease is chronic, and without treatment, they are <a href="/wiki/Infection" title="Infection">infected</a> for the remainder of their lives. There are effective treatments that use <a href="/wiki/Direct-acting_antivirals" class="mw-redirect" title="Direct-acting antivirals">direct-acting antivirals</a>.<a href="#cite_note-Falade-Nwulia_2017-174">[174]</a> The treatment of chronic <a href="/wiki/Asymptomatic_carrier" title="Asymptomatic carrier">carriers</a> of the hepatitis B virus has also been developed by using similar strategies that include lamivudine and other anti-viral drugs.<a href="#cite_note-Nguyen_2020-175">[175]</a> <div class="mw-heading mw-heading2"><h2 id="Infection_in_other_species">Infection in other species</h2></div> Viruses infect all cellular life and, although viruses occur universally, each cellular species has its own specific range that often infects only that species.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 3  Some viruses, called <a href="/wiki/Satellite_(biology)" title="Satellite (biology)">satellites</a>, can replicate only within cells that have already been infected by another virus.<a href="#cite_note-La_Scola_2008-30">[30]</a> <div class="mw-heading mw-heading3"><h3 id="Animal_viruses">Animal viruses</h3></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/Animal_virus" title="Animal virus">Animal virus</a> and <a href="/wiki/Veterinary_virology" title="Veterinary virology">Veterinary virology</a></div> Viruses are important pathogens of livestock. Diseases such as foot-and-mouth disease and <a href="/wiki/Bluetongue" class="mw-redirect" title="Bluetongue">bluetongue</a> are caused by viruses.<a href="#cite_note-Goris_2008-176">[176]</a> Companion animals such as cats, dogs, and horses, if not vaccinated, are susceptible to serious viral infections. <a href="/wiki/Canine_parvovirus" title="Canine parvovirus">Canine parvovirus</a> is caused by a small DNA virus and infections are often fatal in pups.<a href="#cite_note-177">[177]</a> Like all <a href="/wiki/Invertebrates" class="mw-redirect" title="Invertebrates">invertebrates</a>, the honey bee is susceptible to many viral infections.<a href="#cite_note-178">[178]</a> Most viruses co-exist harmlessly in their host and cause no signs or symptoms of disease.<a href="#cite_note-Dimmock_2007-6">[6]</a>: 4  <div class="mw-heading mw-heading3"><h3 id="Plant_viruses">Plant viruses</h3></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/Plant_virus" title="Plant virus">Plant virus</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Pepper_mild_mottle_virus.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Pepper_mild_mottle_virus.png/220px-Pepper_mild_mottle_virus.png" decoding="async" width="220" height="191" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Pepper_mild_mottle_virus.png/330px-Pepper_mild_mottle_virus.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Pepper_mild_mottle_virus.png/440px-Pepper_mild_mottle_virus.png 2x" data-file-width="650" data-file-height="565" /></a><figcaption><a href="/wiki/Capsicum" title="Capsicum">Peppers</a> infected by mild mottle virus</figcaption></figure> There are many types of plant viruses, but often they cause only a loss of <a href="/wiki/Crop_yield" title="Crop yield">yield</a>, and it is not economically viable to try to control them. Plant viruses are often spread from plant to plant by organisms, known as <a href="/wiki/Vector_(epidemiology)" class="mw-redirect" title="Vector (epidemiology)">vectors</a>. These are usually insects, but some fungi, <a href="/wiki/Nematode" title="Nematode">nematode worms</a>, <a href="/wiki/Protozoa" title="Protozoa">single-celled organisms</a>, and parasitic plants are vectors.<a href="#cite_note-179">[179]</a> When control of plant virus infections is considered economical, for perennial fruits, for example, efforts are concentrated on killing the vectors and removing alternate hosts such as weeds.<a href="#cite_note-Shors_2017-13">[13]</a>: 802  Plant viruses cannot infect humans and other animals because they can reproduce only in living plant cells.<a href="#cite_note-Shors_2017-13">[13]</a>: 799–807  Originally from Peru, the potato has become a staple crop worldwide.<a href="#cite_note-Zaheer_2016-180">[180]</a> The <a href="/wiki/Potato_virus_Y" title="Potato virus Y">potato virus Y</a> causes disease in potatoes and related species including tomatoes and peppers. In the 1980s, this virus acquired economical importance when it proved difficult to control in seed potato crops. Transmitted by <a href="/wiki/Aphid" title="Aphid">aphids</a>, this virus can reduce crop yields by up to 80 per cent, causing significant losses to potato yields.<a href="#cite_note-Fuentes_2019-181">[181]</a> Plants have elaborate and effective defence mechanisms against viruses. One of the most effective is the presence of so-called resistance (R) genes. Each R gene confers resistance to a particular virus by triggering localised areas of cell death around the infected cell, which can often be seen with the unaided eye as large spots. This stops the infection from spreading.<a href="#cite_note-182">[182]</a> RNA interference is also an effective defence in plants.<a href="#cite_note-Shors_2017-13">[13]</a>: 809 > When they are infected, plants often produce natural disinfectants that kill viruses, such as <a href="/wiki/Salicylic_acid" title="Salicylic acid">salicylic acid</a>, <a href="/wiki/Nitric_oxide" title="Nitric oxide">nitric oxide</a>, and <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen molecules</a>.<a href="#cite_note-183">[183]</a> Plant virus particles or virus-like particles (VLPs) have applications in both <a href="/wiki/Biotechnology" title="Biotechnology">biotechnology</a> and <a href="/wiki/Nanotechnology" title="Nanotechnology">nanotechnology</a>. The capsids of most plant viruses are simple and robust structures and can be produced in large quantities either by the infection of plants or by expression in a variety of heterologous systems. Plant virus particles can be modified genetically and chemically to encapsulate foreign material and can be incorporated into supramolecular structures for use in biotechnology.<a href="#cite_note-LomonossoffGP-184">[184]</a> <div class="mw-heading mw-heading3"><h3 id="Bacterial_viruses">Bacterial viruses</h3></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/Bacteriophage" title="Bacteriophage">Bacteriophage</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Phage.jpg" class="mw-file-description"><img alt="An electron micrograph showing a portion of a bacterium covered with viruses" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Phage.jpg/170px-Phage.jpg" decoding="async" width="170" height="199" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Phage.jpg/255px-Phage.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/52/Phage.jpg/340px-Phage.jpg 2x" data-file-width="1480" data-file-height="1733" /></a><figcaption>Transmission electron micrograph of multiple bacteriophages attached to a bacterial cell wall</figcaption></figure> Bacteriophages are a common and diverse group of viruses and are the most abundant biological entity in aquatic environments—there are up to ten times more of these viruses in the oceans than there are bacteria,<a href="#cite_note-185">[185]</a> reaching levels of 250,000,000 bacteriophages per millilitre of seawater.<a href="#cite_note-186">[186]</a> These viruses infect specific bacteria by binding to <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">surface receptor molecules</a> and then entering the cell. Within a short amount of time, in some cases, just minutes, bacterial <a href="/wiki/Polymerase" title="Polymerase">polymerase</a> starts translating viral mRNA into protein. These proteins go on to become either new virions within the cell, helper proteins, which help assembly of new virions, or proteins involved in cell lysis. Viral enzymes aid in the breakdown of the cell membrane, and, in the case of the <a href="/wiki/T4_phage" class="mw-redirect" title="T4 phage">T4 phage</a>, in just over twenty minutes after injection over three hundred phages could be released.<a href="#cite_note-Shors_2017-13">[13]</a>: 834–835  The major way bacteria defend themselves from bacteriophages is by producing enzymes that destroy foreign DNA. These enzymes, called <a href="/wiki/Restriction_endonucleases" class="mw-redirect" title="Restriction endonucleases">restriction endonucleases</a>, cut up the viral DNA that bacteriophages inject into bacterial cells.<a href="#cite_note-187">[187]</a> Bacteria also contain a system that uses <a href="/wiki/CRISPR" title="CRISPR">CRISPR</a> sequences to retain fragments of the genomes of viruses that the bacteria have come into contact with previously, which allows them to block the virus's replication through a form of <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a>.<a href="#cite_note-188">[188]</a><a href="#cite_note-189">[189]</a> This genetic system provides bacteria with <a href="/wiki/Immunity_(medical)" class="mw-redirect" title="Immunity (medical)">acquired immunity</a> to infection.<a href="#cite_note-Mojica_2016-190">[190]</a> Some bacteriophages are called "<a href="/wiki/Temperateness_(virology)" title="Temperateness (virology)">temperate</a>" because they cause latent infections and do not immediately destroy their host cells. Instead, their DNA is incorporated with the host cell's as a <a href="/wiki/Prophage" title="Prophage">prophage</a>. These latent infections become productive when the prophage DNA is activated by stimuli such as changes in the environment.<a href="#cite_note-Weiss_2017-191">[191]</a> The intestines of animals, including humans, contain temperate bacteriophages, which are activated by various stimuli including changes in diet and antibiotics.<a href="#cite_note-Hu_2021-192">[192]</a> Although first observed in bacteriophages, many other viruses are known to form <a href="/wiki/Proviruses" class="mw-redirect" title="Proviruses">proviruses</a> including HIV.<a href="#cite_note-Weiss_2017-191">[191]</a><a href="#cite_note-Russ_2023-193">[193]</a> <div class="mw-heading mw-heading3"><h3 id="Archaeal_viruses">Archaeal viruses</h3></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/Archaeal_virus" title="Archaeal virus">Archaeal virus</a></div> Some viruses replicate within <a href="/wiki/Archaea" title="Archaea">archaea</a>: these are DNA viruses with unusual and sometimes unique shapes.<a href="#cite_note-Lawrence_2009-4">[4]</a><a href="#cite_note-Prangishvili-55">[55]</a> These viruses have been studied in most detail in the <a href="/wiki/Thermophile" title="Thermophile">thermophilic</a> archaea, particularly the orders <a href="/wiki/Sulfolobales" title="Sulfolobales">Sulfolobales</a> and <a href="/wiki/Thermoproteales" title="Thermoproteales">Thermoproteales</a>.<a href="#cite_note-194">[194]</a> Defences against these viruses involve RNA interference from <a href="/wiki/Repetitive_DNA" class="mw-redirect" title="Repetitive DNA">repetitive DNA</a> sequences within archaean genomes that are related to the genes of the viruses.<a href="#cite_note-195">[195]</a><a href="#cite_note-196">[196]</a> Most archaea have CRISPR–Cas systems as an adaptive defence against viruses. These enable archaea to retain sections of viral DNA, which are then used to target and eliminate subsequent infections by the virus using a process similar to RNA interference.<a href="#cite_note-van_der_Oost_2014-197">[197]</a> <div class="mw-heading mw-heading2"><h2 id="Role_in_aquatic_ecosystems">Role in aquatic ecosystems</h2></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/Marine_virus" class="mw-redirect" title="Marine virus">Marine virus</a></div> Viruses are the most abundant biological entity in aquatic environments.<a href="#cite_note-Koonin_2006-2">[2]</a> There are about ten million of them in a teaspoon of seawater.<a href="#cite_note-Dávila-Ramos_2019-198">[198]</a> Most of these viruses are <a href="/wiki/Bacteriophages" class="mw-redirect" title="Bacteriophages">bacteriophages</a> infecting heterotrophic bacteria and <a href="/wiki/Cyanophages" class="mw-redirect" title="Cyanophages">cyanophages</a> infecting cyanobacteria and they are essential to the regulation of saltwater and freshwater ecosystems.<a href="#cite_note-Zhang_2018-199">[199]</a> Bacteriophages are harmless to plants and animals, and are essential to the regulation of marine and freshwater ecosystems<a href="#cite_note-200">[200]</a> are important mortality agents of <a href="/wiki/Phytoplankton" title="Phytoplankton">phytoplankton</a>, the base of the <a href="/wiki/Foodchain" class="mw-redirect" title="Foodchain">foodchain</a> in aquatic environments.<a href="#cite_note-Suttle_2005-201">[201]</a> They infect and destroy bacteria in aquatic microbial communities, and are one of the most important mechanisms of <a href="/wiki/Carbon_cycle" title="Carbon cycle">recycling carbon</a> and <a href="/wiki/Nutrient_cycling" class="mw-redirect" title="Nutrient cycling">nutrient cycling</a> in marine environments. The organic molecules released from the dead bacterial cells stimulate fresh bacterial and algal growth, in a process known as the <a href="/wiki/Viral_shunt" title="Viral shunt">viral shunt</a>.<a href="#cite_note-202">[202]</a> In particular, lysis of bacteria by viruses has been shown to enhance nitrogen cycling and stimulate phytoplankton growth.<a href="#cite_note-203">[203]</a> Viral activity may also affect the <a href="/wiki/Biological_pump" title="Biological pump">biological pump</a>, the process whereby <a href="/wiki/Carbon" title="Carbon">carbon</a> is <a href="/wiki/Carbon_sequestration" title="Carbon sequestration">sequestered</a> in the deep ocean.<a href="#cite_note-Suttle_2007-204">[204]</a> Microorganisms constitute more than 90% of the biomass in the sea. It is estimated that viruses kill approximately 20% of this biomass each day and that there are 10 to 15 times as many viruses in the oceans as there are bacteria and archaea.<a href="#cite_note-Wigington_2016-205">[205]</a> Viruses are also major agents responsible for the destruction of <a href="/wiki/Phytoplankton" title="Phytoplankton">phytoplankton</a> including <a href="/wiki/Harmful_algal_bloom" title="Harmful algal bloom">harmful algal blooms</a>,<a href="#cite_note-206">[206]</a> The number of viruses in the oceans decreases further offshore and deeper into the water, where there are fewer host organisms.<a href="#cite_note-Suttle_2007-204">[204]</a> In January 2018, scientists reported that 800 million viruses, mainly of marine origin, are deposited daily from the <a href="/wiki/Earth" title="Earth">Earth</a>'s <a href="/wiki/Atmosphere" title="Atmosphere">atmosphere</a> onto every square meter of the planet's surface, as the result of a global atmospheric stream of viruses, circulating above the weather system but below the altitude of usual airline travel, distributing viruses around the planet.<a href="#cite_note-Robbins_2018-207">[207]</a><a href="#cite_note-ISME-2018-208">[208]</a> Like any organism, <a href="/wiki/Marine_mammal" title="Marine mammal">marine mammals</a> are susceptible to viral infections. In 1988 and 2002, thousands of <a href="/wiki/Harbor_seal" title="Harbor seal">harbour seals</a> were killed in Europe by <a href="/wiki/Phocine_distemper_virus" class="mw-redirect" title="Phocine distemper virus">phocine distemper virus</a>.<a href="#cite_note-209">[209]</a> Many other viruses, including <a href="/wiki/Calicivirus" class="mw-redirect" title="Calicivirus">caliciviruses</a>, <a href="/wiki/Herpesvirus" class="mw-redirect" title="Herpesvirus">herpesviruses</a>, <a href="/wiki/Adenovirus" class="mw-redirect" title="Adenovirus">adenoviruses</a> and <a href="/wiki/Parvovirus" class="mw-redirect" title="Parvovirus">parvoviruses</a>, circulate in marine mammal populations.<a href="#cite_note-Suttle_2007-204">[204]</a> In December 2022, scientists reported the first observation of <a href="/wiki/Virovore" class="mw-redirect" title="Virovore">virovory</a> via an experiment on pond water containing <a href="/wiki/Chlorovirus" title="Chlorovirus">chlorovirus</a>, which commonly infects green algae in freshwater environments. When all other microbial food sources were removed from the water, the <a href="/wiki/Ciliate" title="Ciliate">ciliate</a> <a href="/wiki/Halteria" title="Halteria">Halteria</a> was observed to have increased in number due to the active consumption of chlorovirus as a food source instead of its typical <a href="/wiki/Bacterivore" title="Bacterivore">bacterivore</a> diet.<a href="#cite_note-DeLong_2023-210">[210]</a><a href="#cite_note-Irving_2022-211">[211]</a> <div class="mw-heading mw-heading2"><h2 id="Role_in_evolution">Role in evolution</h2></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/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></div> Viruses are an important natural means of transferring genes between different species, which increases <a href="/wiki/Genetic_diversity" title="Genetic diversity">genetic diversity</a> and drives evolution.<a href="#cite_note-Canchaya_2003-9">[9]</a><a href="#cite_note-Broecker2019-212">[212]</a><a href="#cite_note-213">[213]</a> It is thought that viruses played a central role in early evolution, before the diversification of the <a href="/wiki/Last_universal_ancestor" class="mw-redirect" title="Last universal ancestor">last universal common ancestor</a> into bacteria, archaea and eukaryotes.<a href="#cite_note-Forterre_1999-214">[214]</a> Viruses are still one of the largest reservoirs of unexplored genetic diversity on Earth.<a href="#cite_note-Suttle_2007-204">[204]</a> <div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2></div> <div class="mw-heading mw-heading3"><h3 id="Life_sciences_and_medicine">Life sciences and medicine</h3></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Influenza_virus_research.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Influenza_virus_research.jpg/170px-Influenza_virus_research.jpg" decoding="async" width="170" height="241" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Influenza_virus_research.jpg/255px-Influenza_virus_research.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Influenza_virus_research.jpg/340px-Influenza_virus_research.jpg 2x" data-file-width="1966" data-file-height="2786" /></a><figcaption>Scientist studying the <a href="/wiki/H5N1" class="mw-redirect" title="H5N1">H5N1</a> influenza virus</figcaption></figure> Viruses are important to the study of <a href="/wiki/Molecular_biology" title="Molecular biology">molecular</a> and <a href="/wiki/Cell_biology" title="Cell biology">cell biology</a> as they provide simple systems that can be used to manipulate and investigate the functions of cells.<a href="#cite_note-Collier_1998-26">[26]</a>: 8  The study and use of viruses have provided valuable information about aspects of cell biology.<a href="#cite_note-215">[215]</a> For example, viruses have been useful in the study of <a href="/wiki/Genetics" title="Genetics">genetics</a> and helped our understanding of the basic mechanisms of <a href="/wiki/Molecular_genetics" title="Molecular genetics">molecular genetics</a>, such as <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>, <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a>, <a href="/wiki/RNA_processing" class="mw-redirect" title="RNA processing">RNA processing</a>, <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">translation</a>, <a href="/wiki/Protein" title="Protein">protein</a> transport, and <a href="/wiki/Immunology" title="Immunology">immunology</a>. Geneticists often use viruses as <a href="/wiki/Vector_(molecular_biology)" title="Vector (molecular biology)">vectors</a> to introduce genes into cells that they are studying. This is useful for making the cell produce a foreign substance, or to study the effect of introducing a new gene into the genome. Similarly, <a href="/wiki/Virotherapy" title="Virotherapy">virotherapy</a> uses viruses as vectors to treat various diseases, as they can specifically target cells and DNA. It shows promising use in the treatment of cancer and in <a href="/wiki/Gene_therapy" title="Gene therapy">gene therapy</a>. Eastern European scientists have used <a href="/wiki/Phage_therapy" title="Phage therapy">phage therapy</a> as an alternative to antibiotics for some time, and interest in this approach is increasing, because of the high level of <a href="/wiki/Antibiotic_resistance" class="mw-redirect" title="Antibiotic resistance">antibiotic resistance</a> now found in some pathogenic bacteria.<a href="#cite_note-Matsuzaki_2005-216">[216]</a> The expression of heterologous proteins by viruses is the basis of several manufacturing processes that are currently being used for the production of various proteins such as vaccine <a href="/wiki/Antigen" title="Antigen">antigens</a> and antibodies. Industrial processes have been recently developed using viral vectors and several pharmaceutical proteins are currently in pre-clinical and clinical trials.<a href="#cite_note-GlebaYY-217">[217]</a> <div class="mw-heading mw-heading4"><h4 id="Virotherapy">Virotherapy</h4></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/Virotherapy" title="Virotherapy">Virotherapy</a></div> Virotherapy involves the use of genetically modified viruses to treat diseases.<a href="#cite_note-218">[218]</a> Viruses have been modified by scientists to reproduce in cancer cells and destroy them but not infect healthy cells. <a href="/wiki/Talimogene_laherparepvec" title="Talimogene laherparepvec">Talimogene laherparepvec</a> (T-VEC), for example, is a modified <a href="/wiki/Herpes_simplex_virus" title="Herpes simplex virus">herpes simplex virus</a> that has had a gene, which is required for viruses to replicate in healthy cells, deleted and replaced with a human gene (<a href="/wiki/GM-CSF" class="mw-redirect" title="GM-CSF">GM-CSF</a>) that stimulates immunity. When this virus infects cancer cells, it destroys them and in doing so the presence the GM-CSF gene attracts <a href="/wiki/Dendritic_cells" class="mw-redirect" title="Dendritic cells">dendritic cells</a> from the surrounding tissues of the body. The dendritic cells process the dead cancer cells and present components of them to other cells of the <a href="/wiki/Immune_system" title="Immune system">immune system</a>.<a href="#cite_note-219">[219]</a> Having completed successful <a href="/wiki/Clinical_trials" class="mw-redirect" title="Clinical trials">clinical trials</a>, the virus gained approval for the treatment of <a href="/wiki/Melanoma" title="Melanoma">melanoma</a> in late 2015.<a href="#cite_note-220">[220]</a> Viruses that have been reprogrammed to kill cancer cells are called <a href="/wiki/Oncolytic_virus" title="Oncolytic virus">oncolytic viruses</a>.<a href="#cite_note-221">[221]</a> <div class="mw-heading mw-heading3"><h3 id="Materials_science_and_nanotechnology">Materials science and nanotechnology</h3></div> From the viewpoint of a materials scientist, viruses can be regarded as organic <a href="/wiki/Nanoparticle" title="Nanoparticle">nanoparticles</a>.<a href="#cite_note-222">[222]</a> Their surface carries specific tools that enable them to cross the barriers of their host cells. The size and shape of viruses and the number and nature of the functional groups on their surface are precisely defined. As such, viruses are commonly used in materials science as scaffolds for covalently linked surface modifications. A particular quality of viruses is that they can be tailored by directed evolution. The powerful techniques developed by life sciences are becoming the basis of engineering approaches towards nanomaterials, opening a wide range of applications far beyond biology and medicine.<a href="#cite_note-Fischlechner_2007-223">[223]</a> Because of their size, shape, and well-defined chemical structures, viruses have been used as templates for organising materials on the nanoscale. Examples include the work at the <a href="/wiki/Naval_Research_Laboratory" class="mw-redirect" title="Naval Research Laboratory">Naval Research Laboratory</a> in Washington, D.C., using <a href="/wiki/Cowpea_mosaic_virus" title="Cowpea mosaic virus">Cowpea mosaic virus</a> (CPMV) particles to amplify signals in <a href="/wiki/DNA_microarray" title="DNA microarray">DNA microarray</a> based sensors. In this application, the virus particles separate the <a href="/wiki/Fluorescence" title="Fluorescence">fluorescent</a> <a href="/wiki/Dye" title="Dye">dyes</a> used for signalling to prevent the formation of non-fluorescent <a href="/wiki/Dimer_(chemistry)" class="mw-redirect" title="Dimer (chemistry)">dimers</a> that act as <a href="/wiki/Quenching_(fluorescence)" title="Quenching (fluorescence)">quenchers</a>.<a href="#cite_note-224">[224]</a> Another example is the use of CPMV as a nanoscale breadboard for molecular electronics.<a href="#cite_note-225">[225]</a> <div class="mw-heading mw-heading3"><h3 id="Synthetic_viruses">Synthetic viruses</h3></div> Many viruses can be synthesised de novo ("from scratch"). The first synthetic virus was created in 2002.<a href="#cite_note-226">[226]</a> Although somewhat of a misconception, it is not the actual virus that is synthesised, but rather its DNA genome (in case of a DNA virus), or a <a href="/wiki/CDNA" class="mw-redirect" title="CDNA">cDNA</a> copy of its genome (in case of RNA viruses). For many virus families the naked synthetic DNA or RNA (once enzymatically converted back from the synthetic cDNA) is infectious when introduced into a cell. That is, they contain all the necessary information to produce new viruses. This technology is now being used to investigate novel vaccine strategies.<a href="#cite_note-Coleman_2008-227">[227]</a> The ability to synthesise viruses has far-reaching consequences, since viruses can no longer be regarded as extinct, as long as the information of their genome sequence is known and <a href="/wiki/Permissive" title="Permissive">permissive</a> cells are available. As of June 2021, the full-length genome sequences of 11,464 different viruses, including smallpox, are publicly available in an online database maintained by the <a href="/wiki/National_Institutes_of_Health" title="National Institutes of Health">National Institutes of Health</a>.<a href="#cite_note-228">[228]</a> <div class="mw-heading mw-heading3"><h3 id="Weapons">Weapons</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Biological_warfare" title="Biological warfare">Biological warfare</a></div> The ability of viruses to cause devastating epidemics in human societies has led to the concern that viruses could be weaponised for <a href="/wiki/Biological_warfare" title="Biological warfare">biological warfare</a>. Further concern was raised by the successful recreation of the infamous <a href="/wiki/Spanish_flu" title="Spanish flu">1918 influenza</a> virus in a laboratory.<a href="#cite_note-Zilinskas_2017-229">[229]</a> The smallpox virus devastated numerous societies throughout history before its eradication. There are only two centres in the world authorised by the WHO to keep stocks of smallpox virus: the <a href="/wiki/State_Research_Center_of_Virology_and_Biotechnology_VECTOR" title="State Research Center of Virology and Biotechnology VECTOR">State Research Center of Virology and Biotechnology VECTOR</a> in Russia and the <a href="/wiki/Centers_for_Disease_Control_and_Prevention" title="Centers for Disease Control and Prevention">Centers for Disease Control and Prevention</a> in the United States.<a href="#cite_note-Artenstein_2008-230">[230]</a> It may be used as a weapon,<a href="#cite_note-Artenstein_2008-230">[230]</a> as the vaccine for smallpox sometimes had severe side-effects, it is no longer used routinely in any country. Thus, much of the modern human population has almost no established resistance to smallpox and would be vulnerable to the virus.<a href="#cite_note-Artenstein_2008-230">[230]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 18em;"> <ul><li><a href="/wiki/Cross-species_transmission" title="Cross-species transmission">Cross-species transmission</a></li> <li><a href="/wiki/Glossary_of_virology" title="Glossary of virology">Glossary of virology</a></li> <li><a href="/wiki/Law_of_declining_virulence" class="mw-redirect" title="Law of declining virulence">Law of declining virulence</a> – Disproved hypothesis of epidemiologist Theobald Smith</li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Gene_transfer_agent" title="Gene transfer agent">Gene transfer agent</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li> <li><a href="/wiki/Smallest_organisms" title="Smallest organisms">Smallest organisms</a></li> <li><a href="/wiki/Theory_of_virulence" class="mw-redirect" title="Theory of virulence">Theory of virulence</a> – Theory by biologist Paul W. Ewald</li> <li><a href="/wiki/Viral_metagenomics" title="Viral metagenomics">Viral metagenomics</a></li> <li><a href="/wiki/Viroplasm" title="Viroplasm">Viroplasm</a></li> <li><a href="/wiki/Zoonosis" title="Zoonosis">Zoonosis</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2></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-WU_2020-1"><a href="#cite_ref-WU_2020_1-0">^</a> <style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFWu2020" class="citation news cs1">Wu KJ (15 April 2020). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200415140127/https://www.nationalgeographic.com/science/2020/04/factors-allow-viruses-infect-humans-coronavirus/">"There are more viruses than stars in the universe. 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//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/80px-Wiktionary-logo-en-v2.svg.png 2x" data-file-width="512" data-file-height="512" /></div> <div class="side-box-text plainlist">Look up <a href="https://en.wiktionary.org/wiki/Special:Search/virus" class="extiw" title="wiktionary:Special:Search/virus">virus</a> in Wiktionary, the free dictionary.</div></div> </div> <ul><li><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> Media related to <a href="https://commons.wikimedia.org/wiki/Category:Viruses" class="extiw" title="commons:Category:Viruses">Viruses</a> at Wikimedia Commons</li> <li><a href="/wiki/File:Wikispecies-logo.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/14px-Wikispecies-logo.svg.png" decoding="async" width="14" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/20px-Wikispecies-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/27px-Wikispecies-logo.svg.png 2x" data-file-width="941" data-file-height="1103" /></a> Data related to <a href="https://species.wikimedia.org/wiki/Special:Search/Virus" class="extiw" title="wikispecies:Special:Search/Virus">Virus</a> at Wikispecies</li> <li><a rel="nofollow" class="external text" href="http://viralzone.expasy.org/">ViralZone</a> A Swiss Institute of Bioinformatics resource for all viral families, providing general molecular and epidemiological 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.navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .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="Microbiology:_Virus" 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" 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abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a 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:Virus_topics" title="Template:Virus topics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Virus_topics" title="Template talk:Virus topics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Virus_topics" title="Special:EditPage/Template:Virus topics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Microbiology:_Virus" style="font-size:114%;margin:0 4em"><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a>: <a class="mw-selflink selflink">Virus</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li><a href="/wiki/History_of_virology" title="History of virology">History</a></li> <li><a href="/wiki/Introduction_to_viruses" title="Introduction to viruses">Introduction</a></li> <li><a href="/wiki/Social_history_of_viruses" title="Social history of viruses">Social history of viruses</a></li> <li><a href="/wiki/Virology" title="Virology">Virology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Components</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Capsid" title="Capsid">Capsid</a></li> <li><a href="/wiki/Viral_envelope" title="Viral envelope">Viral envelope</a></li> <li><a href="/wiki/Viral_protein" title="Viral protein">Viral protein</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><a href="/wiki/File:Rotavirus_Reconstruction.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/100px-Rotavirus_Reconstruction.jpg" decoding="async" width="100" height="88" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/150px-Rotavirus_Reconstruction.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Rotavirus_Reconstruction.jpg/200px-Rotavirus_Reconstruction.jpg 2x" data-file-width="1986" data-file-height="1739" /></a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Viral_life_cycle" title="Viral life cycle">Viral life cycle</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Viral_entry" title="Viral entry">Viral entry</a></li> <li><a href="/wiki/Viral_replication" title="Viral replication">Viral replication</a></li> <li><a href="/wiki/Viral_shedding" title="Viral shedding">Viral shedding</a></li> <li><a href="/wiki/Viroplasm" title="Viroplasm">Viroplasm</a></li> <li><a href="/wiki/Virus_latency" title="Virus latency">Virus latency</a></li> <li><a href="/wiki/Lytic_cycle" title="Lytic cycle">Lytic cycle</a></li> <li><a href="/wiki/Lysogenic_cycle" title="Lysogenic cycle">Lysogenic cycle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genetics</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/Antigenic_drift" title="Antigenic drift">Antigenic drift</a></li> <li><a href="/wiki/Antigenic_shift" title="Antigenic shift">Antigenic shift</a></li> <li><a href="/wiki/Phenotype_mixing" title="Phenotype mixing">Phenotype mixing</a></li> <li><a href="/wiki/Reassortment" title="Reassortment">Reassortment</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viral evolution</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By host</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_virus" title="Animal virus">Animal virus</a></li> <li><a href="/wiki/Bacteriophage" title="Bacteriophage">Bacteriophage</a></li> <li><a href="/wiki/Virophage" title="Virophage">Virophage</a></li> <li><a href="/wiki/Human_virome" title="Human virome">Human virome</a></li> <li><a href="/wiki/Mycovirus" title="Mycovirus">Mycovirus</a></li> <li><a href="/wiki/Plant_virus" title="Plant virus">Plant virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antiviral_drug" title="Antiviral drug">Antiviral drug</a></li> <li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li> <li><a href="/wiki/Helper_virus" title="Helper virus">Helper virus</a></li> <li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a> <ul><li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li><a href="/wiki/Laboratory_diagnosis_of_viral_infections" title="Laboratory diagnosis of viral infections">Laboratory diagnosis of viral infections</a></li> <li><a href="/wiki/Marine_viruses" title="Marine viruses">Marine viruses</a></li> <li><a href="/wiki/Neurotropic_virus" title="Neurotropic virus">Neurotropic virus</a></li> <li><a href="/wiki/Oncovirus" title="Oncovirus">Oncovirus</a></li> <li><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellites</a></li> <li><a href="/wiki/Viral_disease" title="Viral disease">Viral disease</a></li> <li><a href="/wiki/Viral_load" title="Viral load">Viral load</a></li> <li><a href="/wiki/Virus-like_particle" title="Virus-like particle">Virus-like particle</a></li> <li><a href="/wiki/Virus_classification" title="Virus classification">Virus classification</a></li> <li><a href="/wiki/Virus_quantification" title="Virus quantification">Virus quantification</a></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li> <li><a href="/wiki/Virosphere" title="Virosphere">Virosphere</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a> <a href="/wiki/Portal:Viruses" title="Portal:Viruses">Portal</a></li> <li><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" /> <a href="/wiki/Category:Viruses" title="Category:Viruses">Category</a></li> <li><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 href="https://commons.wikimedia.org/wiki/Category:Viruses" class="extiw" title="commons:Category:Viruses">Commons</a></li> <li><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/16px-People_icon.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/24px-People_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/32px-People_icon.svg.png 2x" data-file-width="100" data-file-height="100" /> <a href="/wiki/Wikipedia:WikiProject_Viruses" title="Wikipedia:WikiProject Viruses">WikiProject</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Self-replicating_organic_structures" 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" style="text-align: center;"><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:Self-replicating_organic_structures" title="Template:Self-replicating organic structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Self-replicating_organic_structures" title="Template talk:Self-replicating organic structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Self-replicating_organic_structures" title="Special:EditPage/Template:Self-replicating organic structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Self-replicating_organic_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Self-replication" title="Self-replication">Self-replicating</a> organic structures</div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Life" title="Life">Cellular life</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a></li> <li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li><a href="/wiki/Protist" title="Protist">Protista</a></li></ul></li> <li><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a> <ul><li><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></li> <li><a href="/wiki/Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a class="mw-selflink selflink">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a> <ul><li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li></ul></li> <li><a href="/wiki/DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li> <li><a href="/wiki/Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li> <li><a href="/wiki/Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li> <li><a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li> <li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Helper_virus" title="Helper virus">Helper-virus dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;"><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>ssRNA satellite virus</li> <li>dsDNA satellite virus (<a href="/wiki/Virophage" title="Virophage">Virophage</a>)</li> <li>ssDNA satellite virus</li> <li>ssDNA satellite</li> <li>dsRNA satellite</li> <li>ssRNA satellite (<a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>)</li> <li>Satellite-like nucleic acids <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a> <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Prion" title="Prion">Mammalian prion</a></li> <li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a> self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mobilome" title="Mobilome">Mobilome</a> <ul><li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Genomic_island" title="Genomic island">Genomic island</a></li></ul></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li> <li><a href="/wiki/DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a> <ul><li><a href="/wiki/Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li> <li><a href="/wiki/R-factor" class="mw-redirect" title="R-factor">Resistance</a></li> <li><a href="/wiki/Colicin" title="Colicin">Col</a></li> <li>Degradative</li> <li><a href="/wiki/Virulence_factor" title="Virulence factor">Virulence</a>/<a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti</a></li> <li>Cryptic</li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a> <ul><li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li></ul></li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li> <li><a href="/wiki/Group_II_intron" title="Group II intron">Group II intron</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a> <ul><li><a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear</a></li> <li><a href="/wiki/Circular_chromosome" title="Circular chromosome">Circular</a></li> <li><a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li> <li><a href="/wiki/Genome" title="Genome">Genome</a> <ul><li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Prophage" title="Prophage">Prophage</a></li> <li><a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a> <ul><li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li> <li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Kappa_organism" title="Kappa organism">Kappa organism</a></li> <li>Organs <ul><li><a href="/wiki/Bacteriome" title="Bacteriome">Bacteriome</a></li> <li><a href="/wiki/Trophosome" title="Trophosome">Trophosome</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li>?<a href="/wiki/RNA_world" title="RNA world">RNA life</a> <ul><li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li> <li>†<a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Coacervate" title="Coacervate">Coacervate</a></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a></li> <li><a href="/wiki/Sulphobes" title="Sulphobes">Sulphobe</a></li> <li>Research <ul><li><a href="/wiki/Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Organism" title="Organism">Organism</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a> <ul><li><a href="/wiki/Cell_division" title="Cell division">Cell division</a></li> <li><a href="/wiki/Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Synthetic_virology" title="Synthetic virology">Synthetic virus</a> <ul><li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a></li> <li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li>?<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></li> <li>?<a href="/wiki/Nanobe" title="Nanobe">Nanobe</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Life,_non-cellular_life,_and_comparable_structures" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Life,_non-cellular_life,_and_comparable_structures" title="Template:Life, non-cellular life, and comparable structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Life,_non-cellular_life,_and_comparable_structures" title="Template talk:Life, non-cellular life, and comparable structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Life,_non-cellular_life,_and_comparable_structures" title="Special:EditPage/Template:Life, non-cellular life, and comparable structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Life,_non-cellular_life,_and_comparable_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Life" title="Life">Life</a>, <a href="/wiki/Non-cellular_life" title="Non-cellular life">non-cellular life</a>, and comparable structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">"<a href="/wiki/Prokaryote" title="Prokaryote">Prokaryota</a>"</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li></ul></li> <li><a href="/wiki/LUCA" class="mw-redirect" title="LUCA">LUCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li>'<a href="/wiki/Protist" title="Protist">Protista</a>'</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></li> <li><a href="/wiki/Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Virus</a> <hr /><a href="/wiki/Viroid" title="Viroid">Viroid</a> <hr /><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Realms</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Adnaviria" title="Adnaviria">Adnaviria</a></li> <li><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></li> <li><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></li> <li><a href="/wiki/Riboviria" title="Riboviria">Riboviria</a></li> <li><a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a></li> <li><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Unassigned</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Classes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><a href="/wiki/Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Families</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></li> <li><a href="/wiki/Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></li> <li><a href="/wiki/Anelloviridae" title="Anelloviridae">Anelloviridae</a></li> <li><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></li> <li><a href="/wiki/Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></li> <li><a href="/wiki/Clavaviridae" title="Clavaviridae">Clavaviridae</a></li> <li><a href="/wiki/Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></li> <li><a href="/wiki/Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></li> <li><a href="/wiki/Globuloviridae" title="Globuloviridae">Globuloviridae</a></li> <li><a href="/wiki/Guttaviridae" title="Guttaviridae">Guttaviridae</a></li> <li><a href="/wiki/Halspiviridae" title="Halspiviridae">Halspiviridae</a></li> <li><a href="/wiki/Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></li> <li><a href="/wiki/Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></li> <li><a href="/wiki/Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></li> <li><a href="/wiki/Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></li> <li><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></li> <li><a href="/wiki/Spiraviridae" title="Spiraviridae">Spiraviridae</a></li> <li><a href="/wiki/Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></li> <li><a href="/wiki/Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genera</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></li> <li><a href="/wiki/Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nanobe" title="Nanobe">Nanobe</a> (?)</li> <li><a href="/wiki/Obelisk_(biology)" title="Obelisk (biology)">Obelisk</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Comparable structures</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bio-like_structure" title="Bio-like structure">Bio-like structure</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Integrative_and_conjugative_element" title="Integrative and conjugative element">Integrative and conjugative element</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li> <li>"<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a>"</li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Prion" title="Prion">Prion</a> <ul><li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a> <a href="/wiki/Microparticle#Biological_protocells" title="Microparticle">microsphere</a></li> <li>Retroelements not elsewhere classified <ul><li><a href="/wiki/Retron" title="Retron">Retron</a></li> <li><a href="/wiki/Diversity-generating_retroelement" title="Diversity-generating retroelement">Diversity-generating retroelement</a></li> <li><a href="/wiki/Telomerase_reverse_transcriptase" title="Telomerase reverse transcriptase">Telomerase reverse transcriptase</a></li> <li><a href="/wiki/Reverse_transcriptase-related_cellular_gene" class="mw-redirect" title="Reverse transcriptase-related cellular gene">Reverse transcriptase-related cellular gene</a></li></ul></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Spiegelman%27s_Monster" title="Spiegelman's Monster">Spiegelman's Monster</a></li> <li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retroposon" title="Retroposon">Retroposon</a></li></ul></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li> <li><a href="/wiki/Xenobot" title="Xenobot">Xenobot</a></li></ul> </div></td></tr></tbody></table></div> <style data-mw-deduplicate="TemplateStyles:r1130092004">.mw-parser-output .portal-bar{font-size:88%;font-weight:bold;display:flex;justify-content:center;align-items:baseline}.mw-parser-output .portal-bar-bordered{padding:0 2em;background-color:#fdfdfd;border:1px solid #a2a9b1;clear:both;margin:1em auto 0}.mw-parser-output .portal-bar-related{font-size:100%;justify-content:flex-start}.mw-parser-output .portal-bar-unbordered{padding:0 1.7em;margin-left:0}.mw-parser-output .portal-bar-header{margin:0 1em 0 0.5em;flex:0 0 auto;min-height:24px}.mw-parser-output .portal-bar-content{display:flex;flex-flow:row wrap;flex:0 1 auto;padding:0.15em 0;column-gap:1em;align-items:baseline;margin:0;list-style:none}.mw-parser-output .portal-bar-content-related{margin:0;list-style:none}.mw-parser-output .portal-bar-item{display:inline-block;margin:0.15em 0.2em;min-height:24px;line-height:24px}@media screen and (max-width:768px){.mw-parser-output .portal-bar{font-size:88%;font-weight:bold;display:flex;flex-flow:column wrap;align-items:baseline}.mw-parser-output .portal-bar-header{text-align:center;flex:0;padding-left:0.5em;margin:0 auto}.mw-parser-output .portal-bar-related{font-size:100%;align-items:flex-start}.mw-parser-output .portal-bar-content{display:flex;flex-flow:row wrap;align-items:center;flex:0;column-gap:1em;border-top:1px solid #a2a9b1;margin:0 auto;list-style:none}.mw-parser-output .portal-bar-content-related{border-top:none;margin:0;list-style:none}}.mw-parser-output .navbox+link+.portal-bar,.mw-parser-output .navbox+style+.portal-bar,.mw-parser-output .navbox+link+.portal-bar-bordered,.mw-parser-output .navbox+style+.portal-bar-bordered,.mw-parser-output .sister-bar+link+.portal-bar,.mw-parser-output .sister-bar+style+.portal-bar,.mw-parser-output .portal-bar+.navbox-styles+.navbox,.mw-parser-output .portal-bar+.navbox-styles+.sister-bar{margin-top:-1px}</style><div class="portal-bar noprint metadata noviewer portal-bar-bordered" role="navigation" aria-label="Portals"><a href="/wiki/Wikipedia:Contents/Portals" title="Wikipedia:Contents/Portals">Portals</a>:<ul class="portal-bar-content"><li class="portal-bar-item"><a href="/wiki/File:Issoria_lathonia.jpg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/21px-Issoria_lathonia.jpg" decoding="async" width="21" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/32px-Issoria_lathonia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/42px-Issoria_lathonia.jpg 2x" data-file-width="629" data-file-height="445" /></a> <a href="/wiki/Portal:Biology" title="Portal:Biology">Biology</a></li><li class="portal-bar-item"><a href="/wiki/File:Tree_of_life.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/21px-Tree_of_life.svg.png" decoding="async" width="21" height="17" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/32px-Tree_of_life.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/42px-Tree_of_life.svg.png 2x" data-file-width="915" data-file-height="760" /></a> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology</a></li><li class="portal-bar-item"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/8px-WHO_Rod.svg.png" decoding="async" width="8" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/12px-WHO_Rod.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/16px-WHO_Rod.svg.png 2x" data-file-width="107" data-file-height="250" /> <a href="/wiki/Portal:Medicine" title="Portal:Medicine">Medicine</a></li><li class="portal-bar-item"><a href="/wiki/File:Nuvola_apps_kalzium.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Nuvola_apps_kalzium.svg/19px-Nuvola_apps_kalzium.svg.png" decoding="async" width="19" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Nuvola_apps_kalzium.svg/29px-Nuvola_apps_kalzium.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Nuvola_apps_kalzium.svg/38px-Nuvola_apps_kalzium.svg.png 2x" data-file-width="128" data-file-height="128" /></a> <a href="/wiki/Portal:Science" title="Portal:Science">Science</a></li><li class="portal-bar-item"><a href="/wiki/File:Sida-aids.png" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/19px-Sida-aids.png" decoding="async" width="19" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/29px-Sida-aids.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Sida-aids.png/38px-Sida-aids.png 2x" data-file-width="290" data-file-height="290" /></a> <a href="/wiki/Portal:Viruses" title="Portal:Viruses">Viruses</a></li></ul></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Taxon_identifiers" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Taxon_identifiers" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Taxon_identifiers" title="Help:Taxon identifiers">Taxon identifiers</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;">Virus</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/Wikidata" title="Wikidata">Wikidata</a>: <a href="https://www.wikidata.org/wiki/Q808" class="extiw" title="wikidata:Q808">Q808</a></li> <li><a href="/wiki/Wikispecies" title="Wikispecies">Wikispecies</a>: <a href="https://species.wikimedia.org/wiki/Virus" class="extiw" title="wikispecies:Virus">Virus</a></li> <li><a href="/wiki/Barcode_of_Life_Data_System" title="Barcode of Life Data System">BOLD</a>: <a rel="nofollow" class="external text" href="http://www.boldsystems.org/index.php/TaxBrowser_TaxonPage?taxid=782766">782766</a></li> <li><a href="/wiki/Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <a rel="nofollow" class="external text" href="https://eol.org/pages/5006">5006</a></li> <li><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <a rel="nofollow" class="external text" href="https://www.gbif.org/species/8">8</a></li> <li><a href="/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/131236">131236</a></li> <li><a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=10239">10239</a></li> <li><a href="/wiki/World_Register_of_Marine_Species" title="World Register of Marine Species">WoRMS</a>: <a rel="nofollow" class="external text" href="https://www.marinespecies.org/aphia.php?p=taxdetails&id=10">10</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" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q808#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" 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="2"><div id="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q808#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <a href="https://www.wikidata.org/wiki/Q808#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">International</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 rel="nofollow" class="external text" href="http://id.worldcat.org/fast/1167771/">FAST</a></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</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 rel="nofollow" class="external text" href="https://d-nb.info/gnd/4063589-2">Germany</a></li><li><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85143833">United States</a></li><li><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb11938341t">France</a></li><li><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb11938341t">BnF data</a></li><li><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/00560678">Japan</a></li><li><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=ph116578&CON_LNG=ENG">Czech Republic</a></li><li><a rel="nofollow" class="external text" href="http://catalogo.bne.es/uhtbin/authoritybrowse.cgi?action=display&authority_id=XX527219">Spain</a></li><li><a rel="nofollow" class="external text" href="https://kopkatalogs.lv/F?func=direct&local_base=lnc10&doc_number=000094794&P_CON_LNG=ENG">Latvia</a></li><li><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007541431905171">Israel</a></li></ul></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Virus&oldid=1253984986">https://en.wikipedia.org/w/index.php?title=Virus&oldid=1253984986</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" 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