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Defective interfering particle - Wikipedia

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<span>Interference</span> </div> </a> <ul id="toc-Interference-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Pathogenesis" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Pathogenesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Pathogenesis</span> </div> </a> <ul id="toc-Pathogenesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Types_of_defective_RNA_genomes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Types_of_defective_RNA_genomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Types of defective RNA genomes</span> </div> </a> <ul id="toc-Types_of_defective_RNA_genomes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Research" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Research"> <div 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class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </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"><p class="mw-empty-elt"> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:RF00496.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/RF00496.jpg/220px-RF00496.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/RF00496.jpg/330px-RF00496.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5b/RF00496.jpg/440px-RF00496.jpg 2x" data-file-width="452" data-file-height="452" /></a><figcaption>Predicted secondary structure of the <a href="/w/index.php?title=Coronavirus_SL-III_cis-acting_replication_element&amp;action=edit&amp;redlink=1" class="new" title="Coronavirus SL-III cis-acting replication element (page does not exist)">Coronavirus SL-III cis-acting replication element</a>, a genomic structure required for BCoV DI RNA replication<sup id="cite_ref-BovineCorona_1-0" class="reference"><a href="#cite_note-BovineCorona-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup></figcaption></figure> <p><b>Defective interfering particles</b> (<b>DIPs</b>), also known as <b>defective interfering viruses</b>, are spontaneously generated <a href="/wiki/Virus" title="Virus">virus</a> mutants in which a critical portion of the particle's genome has been lost due to defective replication or <a href="/wiki/Non-homologous_recombination" class="mw-redirect" title="Non-homologous recombination">non-homologous recombination</a>.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-AntiviralDIPs_3-0" class="reference"><a href="#cite_note-AntiviralDIPs-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup> The mechanism of their formation is presumed to be as a result of template-switching during replication of the viral genome, although non-replicative mechanisms involving direct ligation of genomic RNA fragments have also been proposed.<sup id="cite_ref-Pathak_4-0" class="reference"><a href="#cite_note-Pathak-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> DIPs are derived from and associated with their parent virus, and particles are classed as DIPs if they are rendered non-infectious due to at least one essential gene of the virus being lost or severely damaged as a result of the defection.<sup id="cite_ref-Pathak_4-1" class="reference"><a href="#cite_note-Pathak-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> A DIP can usually still penetrate host cells, but requires another fully functional virus particle (the 'helper' virus) to <a href="/wiki/Coinfection" title="Coinfection">co-infect</a> a cell with it, in order to provide the lost factors.<sup id="cite_ref-Makino_6-0" class="reference"><a href="#cite_note-Makino-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Textbook_7-0" class="reference"><a href="#cite_note-Textbook-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> </p><p>DIPs were first <a href="/wiki/Von_Magnus_phenomenon" title="Von Magnus phenomenon">observed</a> as early as the 1950s by Von Magnus and Schlesinger, both working with influenza viruses.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> However, direct evidence for DIPs was only found in the 1960s by Hackett who noticed presence of ‘stumpy’ particles of vesicular stomatitis virus in electron micrographs<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> and the formalization of DIPs terminology was in 1970 by Huang and Baltimore.<sup id="cite_ref-Huang_10-0" class="reference"><a href="#cite_note-Huang-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> DIPs can occur within nearly every class of both DNA and RNA viruses both in clinical and laboratory settings including <a href="/wiki/Poliovirus" title="Poliovirus">poliovirus</a>, <a href="/wiki/SARS_coronavirus" class="mw-redirect" title="SARS coronavirus">SARS coronavirus</a>, <a href="/wiki/Measles_virus" title="Measles virus">measles</a>, <a href="/wiki/Alphavirus" title="Alphavirus">alphaviruses</a>, <a href="/wiki/Human_respiratory_syncytial_virus" class="mw-redirect" title="Human respiratory syncytial virus">respiratory syncytial virus</a> and <a href="/wiki/Influenza" title="Influenza">influenza virus</a>.<sup id="cite_ref-ReferenceA_11-0" class="reference"><a href="#cite_note-ReferenceA-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Defection">Defection</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Defective_interfering_particle&amp;action=edit&amp;section=1" title="Edit section: Defection"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>DIPs are a naturally occurring phenomenon that can be recreated under experimental conditions in the lab and can also be synthesized for experimental use. They are spontaneously produced by error-prone <a href="/wiki/Viral_replication" title="Viral replication">viral replication</a>, something particularly prevalent in RNA viruses over DNA viruses due to the enzyme used (replicase, or <a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA-dependent RNA polymerase</a>.)<sup id="cite_ref-Pathak_4-2" class="reference"><a href="#cite_note-Pathak-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Chiba_19-0" class="reference"><a href="#cite_note-Chiba-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> DI genomes typically retain the termini sequences needed for recognition by viral polymerases, and sequences for packaging of their genome into new particles, but little else.<sup id="cite_ref-DimmockBook_20-0" class="reference"><a href="#cite_note-DimmockBook-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Resende_21-0" class="reference"><a href="#cite_note-Resende-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> The size of the genomic deletion event can vary greatly, with one such example in a DIP derived from rabies virus exhibiting a 6.1 kb deletion.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> In another example, the size of several DI-DNA plant virus genomes varied from one tenth of the size of the original genome to one half.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Interference">Interference</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Defective_interfering_particle&amp;action=edit&amp;section=2" title="Edit section: Interference"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The particles are considered interfering when they affect the function of the parent virus through <a href="/wiki/Competitive_inhibition" title="Competitive inhibition">competitive inhibition</a><sup id="cite_ref-Pathak_4-3" class="reference"><a href="#cite_note-Pathak-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> during coinfection. In other words, defective and non-defective viruses replicate simultaneously, but when defective particles increase, the amount of replicated non-defective virus is decreased. The extent of interference depends on the type and size of defection in the genome; large deletions of genomic data allow rapid replication of the defective genome.<sup id="cite_ref-DimmockBook_20-1" class="reference"><a href="#cite_note-DimmockBook-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> In SARS-CoV-2, synthetic DIPs made by removing 90% of the genome replicate three times faster than the virus.<sup id="cite_ref-:0_24-0" class="reference"><a href="#cite_note-:0-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> During the coinfection of a host cell, a critical ratio will eventually be reached in which more viral factors are being used to produce the non-infectious DIPs than infectious particles.<sup id="cite_ref-DimmockBook_20-2" class="reference"><a href="#cite_note-DimmockBook-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> Defective particles and defective genomes have also been demonstrated to stimulate the host innate immune responses and their presence during a viral infection correlates with the strength of the antiviral response.<sup id="cite_ref-ReferenceA_11-1" class="reference"><a href="#cite_note-ReferenceA-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> However, in some viruses such as SARS-CoV-2, the effect of competitive inhibition by interfering particles reduces viral-mediated innate immune responses and inflammation producing a therapeutic effect.<sup id="cite_ref-Chaturvedi_2021_25-0" class="reference"><a href="#cite_note-Chaturvedi_2021-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> </p><p>This interfering nature is becoming more and more important for research on virus therapies.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Thompson_27-0" class="reference"><a href="#cite_note-Thompson-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup> It is thought that because of their specificity, DIPs will be targeted to sites of infection. In one example, scientists have used DIPs to create "protecting viruses", which attenuated the <a href="/wiki/Pathogenicity" class="mw-redirect" title="Pathogenicity">pathogenicity</a> of an influenza A infection in mice, through inducing an <a href="/wiki/Interferon" title="Interferon">interferon</a> response, to a point that it was no longer lethal.<sup id="cite_ref-Easton_28-0" class="reference"><a href="#cite_note-Easton-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> For <a href="/wiki/SARS-CoV-2" title="SARS-CoV-2">SARS-CoV-2</a>, the first synthetic DIPs were made in 2020 <sup id="cite_ref-:0_24-1" class="reference"><a href="#cite_note-:0-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> and the interference effect was used to generate <a href="/wiki/Therapeutic_interfering_particle" title="Therapeutic interfering particle">therapeutic interfering particles (TIPs)</a> that reduced pathogenesis and protected hamsters from serious disease.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">&#91;</span>29<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Pathogenesis">Pathogenesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Defective_interfering_particle&amp;action=edit&amp;section=3" title="Edit section: Pathogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>DIPs have been shown to play a role in pathogenesis of certain viruses. One study demonstrates the relationship between a pathogen and its defective variant, showing how regulation of DI production allowed the virus to attenuate its own infectious replication, decreasing viral load and thus enhance its parasitic efficiency by preventing the host from dying too fast.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">&#91;</span>30<span class="cite-bracket">&#93;</span></a></sup> This also provides the virus with more time to spread and infect new hosts. DIP generation is regulated within viruses: the <a href="/w/index.php?title=Coronavirus_SL-III_cis-acting_replication_element&amp;action=edit&amp;redlink=1" class="new" title="Coronavirus SL-III cis-acting replication element (page does not exist)">Coronavirus SL-III cis-acting replication element</a> (shown in the image) is a higher-order genomic structure implicated in the mediation of DIP production in <a href="/wiki/Bovine_coronavirus" title="Bovine coronavirus">bovine coronavirus</a>, with apparent homologs detected in other <a href="/wiki/Coronavirus" title="Coronavirus">coronavirus</a> groups.<sup id="cite_ref-BovineCorona_1-1" class="reference"><a href="#cite_note-BovineCorona-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> A more in-depth introduction can be found in Alice Huang and David Baltimore's work from 1970.<sup id="cite_ref-Huang_10-1" class="reference"><a href="#cite_note-Huang-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Types_of_defective_RNA_genomes">Types of defective RNA genomes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Defective_interfering_particle&amp;action=edit&amp;section=4" title="Edit section: Types of defective RNA genomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ol><li>Deletions defections are when a fragment of the template is skipped. Examples of this type of defection can be found in tomato spotted wilt virus and Flock House virus.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">&#91;</span>31<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Resende_21-1" class="reference"><a href="#cite_note-Resende-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup></li> <li>Snapbacks defections are when replicase transcribes part of one strand then uses this new strand as a template. The result of this can produce a hairpin. Snapback defections have been observed in <a href="/wiki/Human_respiratory_syncytial_virus" class="mw-redirect" title="Human respiratory syncytial virus">vesicular stomatitis virus</a>.<sup id="cite_ref-Schubert_32-0" class="reference"><a href="#cite_note-Schubert-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup></li> <li>Panhandle defections are when the polymerase carries a partially made strand and then switches back to transcribe the 5' end, forming the panhandle shape. Panhandle defections are found in influenza viruses.<sup id="cite_ref-Fodor_33-0" class="reference"><a href="#cite_note-Fodor-33"><span class="cite-bracket">&#91;</span>33<span class="cite-bracket">&#93;</span></a></sup></li> <li>Compound defections are when both a deletion and snapback defection happens together.</li> <li>Mosaic or complex DI genome, in which the various regions may come from the same helper virus genome but in the wrong order; from different helper genome segments, or could include segments of host RNA. Duplications may also occur.<sup id="cite_ref-AntiviralDIPs_3-1" class="reference"><a href="#cite_note-AntiviralDIPs-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup></li></ol> <div class="mw-heading mw-heading2"><h2 id="Research">Research</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Defective_interfering_particle&amp;action=edit&amp;section=5" title="Edit section: Research"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Research has been conducted by virologists to learn more about the interference in infection of host cells and how DI genomes could potentially work as <a href="/wiki/Immunostimulatory" class="mw-redirect" title="Immunostimulatory">immunostimulatory</a> antiviral agents.<sup id="cite_ref-AntiviralDIPs_3-2" class="reference"><a href="#cite_note-AntiviralDIPs-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup> Another branch of research has pursued the concept of engineering DIPs into antiviral <a href="/wiki/Therapeutic_interfering_particle" title="Therapeutic interfering particle">therapeutic interfering particles (TIPs)</a>,<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">&#91;</span>34<span class="cite-bracket">&#93;</span></a></sup> a purely theoretical concept until recently.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">&#91;</span>35<span class="cite-bracket">&#93;</span></a></sup> A 2014 article describes the pre-clinical work to test the immunostimulatory effectiveness of a DIP against influenza viruses by inducing the innate antiviral immune responses (i.e., interferon).<sup id="cite_ref-Dimmock_36-0" class="reference"><a href="#cite_note-Dimmock-36"><span class="cite-bracket">&#91;</span>36<span class="cite-bracket">&#93;</span></a></sup> Subsequent work tested the pre-clinical efficacy of TIPs against <a href="/wiki/HIV" title="HIV">HIV</a><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">&#91;</span>37<span class="cite-bracket">&#93;</span></a></sup> and <a href="/wiki/SARS-CoV-2" title="SARS-CoV-2">SARS-CoV-2</a>.<sup id="cite_ref-:0_24-2" class="reference"><a href="#cite_note-:0-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Chaturvedi_2021_25-1" class="reference"><a href="#cite_note-Chaturvedi_2021-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> DI-RNAs have also been found to aid in the infection of fungi via viruses of the family <i><a href="/wiki/Partitiviridae" title="Partitiviridae">Partitiviridae</a></i> for the first time, which makes room for more interdisciplinary work.<sup id="cite_ref-Chiba_19-1" class="reference"><a href="#cite_note-Chiba-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> </p><p>Several tools as ViReMa<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">&#91;</span>38<span class="cite-bracket">&#93;</span></a></sup> and DI-tector<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">&#91;</span>39<span class="cite-bracket">&#93;</span></a></sup> have been developed to help to detect defective viral genomes in next-generation sequencing data and <a href="/wiki/High-throughput_screening" title="High-throughput screening">high-throughput</a> approaches, such as random-deletion library <a href="/wiki/Sequencing" title="Sequencing">sequencing</a> (RanDeL-Seq),<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">&#91;</span>40<span class="cite-bracket">&#93;</span></a></sup> allow rational mapping of the viral genetic elements that are required for DI-particle propagation. </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Defective_interfering_particle&amp;action=edit&amp;section=6" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-BovineCorona-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-BovineCorona_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-BovineCorona_1-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFRamanBoumaWilliamsBrian2003" class="citation journal cs1">Raman S, Bouma P, Williams GD, Brian DA (June 2003). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC156170">"Stem-loop III in the 5' untranslated region is a cis-acting element in bovine coronavirus defective interfering RNA replication"</a>. <i>Journal of Virology</i>. <b>77</b> (12): 6720–6730. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1128%2FJVI.77.12.6720-6730.2003">10.1128/JVI.77.12.6720-6730.2003</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&#160;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC156170">156170</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12767992">12767992</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Journal+of+Virology&amp;rft.atitle=Stem-loop+III+in+the+5%27+untranslated+region+is+a+cis-acting+element+in+bovine+coronavirus+defective+interfering+RNA+replication&amp;rft.volume=77&amp;rft.issue=12&amp;rft.pages=6720-6730&amp;rft.date=2003-06&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC156170%23id-name%3DPMC&amp;rft_id=info%3Apmid%2F12767992&amp;rft_id=info%3Adoi%2F10.1128%2FJVI.77.12.6720-6730.2003&amp;rft.aulast=Raman&amp;rft.aufirst=S&amp;rft.au=Bouma%2C+P&amp;rft.au=Williams%2C+GD&amp;rft.au=Brian%2C+DA&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC156170&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ADefective+interfering+particle" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link 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(identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/8254723">8254723</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Journal+of+Virology&amp;rft.atitle=Nonhomologous+RNA+recombination+in+tombusviruses%3A+generation+and+evolution+of+defective+interfering+RNAs+by+stepwise+deletions&amp;rft.volume=68&amp;rft.issue=1&amp;rft.pages=14-24&amp;rft.date=1994-01&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC236259%23id-name%3DPMC&amp;rft_id=info%3Apmid%2F8254723&amp;rft_id=info%3Adoi%2F10.1128%2FJVI.68.1.14-24.1994&amp;rft.aulast=White&amp;rft.aufirst=KA&amp;rft.au=Morris%2C+TJ&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC236259&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ADefective+interfering+particle" class="Z3988"></span></span> </li> <li id="cite_note-AntiviralDIPs-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-AntiviralDIPs_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-AntiviralDIPs_3-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-AntiviralDIPs_3-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMarriottDimmock2010" class="citation journal cs1">Marriott AC, Dimmock NJ (January 2010). 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</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<br />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><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></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<br />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 class="mw-selflink selflink">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><br />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<br />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%"><i><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></i></li> <li><a href="/wiki/Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Virus" title="Virus">Virus</a> <hr /><a 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><i><a href="/wiki/Adnaviria" title="Adnaviria">Adnaviria</a></i></li> <li><i><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></i></li> <li><i><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></i></li> <li><i><a href="/wiki/Riboviria" title="Riboviria">Riboviria</a></i></li> <li><i><a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a></i></li> <li><i><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></i></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"><i><a href="/wiki/Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></i></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><i><a href="/wiki/Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></i></li> <li><i><a href="/wiki/Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></i></li> <li><i><a href="/wiki/Anelloviridae" title="Anelloviridae">Anelloviridae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li> <li><i><a href="/wiki/Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></i></li> <li><i><a href="/wiki/Clavaviridae" title="Clavaviridae">Clavaviridae</a></i></li> <li><i><a href="/wiki/Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></i></li> <li><i><a href="/wiki/Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></i></li> <li><i><a href="/wiki/Globuloviridae" title="Globuloviridae">Globuloviridae</a></i></li> <li><i><a href="/wiki/Guttaviridae" title="Guttaviridae">Guttaviridae</a></i></li> <li><i><a href="/wiki/Halspiviridae" title="Halspiviridae">Halspiviridae</a></i></li> <li><i><a href="/wiki/Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></i></li> <li><i><a href="/wiki/Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></i></li> <li><i><a href="/wiki/Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></i></li> <li><i><a href="/wiki/Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></i></li> <li><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Spiraviridae" title="Spiraviridae">Spiraviridae</a></i></li> <li><i><a href="/wiki/Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></i></li> <li><i><a href="/wiki/Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></i></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><i><a href="/wiki/Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></i></li> <li><i><a href="/wiki/Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></i></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<br />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 class="mw-selflink selflink">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>"<i><a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></i>"</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&#39;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> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐f69cdc8f6‐67nx9 Cached time: 20241124085644 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 0.589 seconds Real time usage: 0.754 seconds Preprocessor visited node count: 2415/1000000 Post‐expand include size: 176368/2097152 bytes Template argument size: 94/2097152 bytes Highest expansion depth: 10/100 Expensive parser function count: 1/500 Unstrip recursion depth: 1/20 Unstrip post‐expand size: 186700/5000000 bytes Lua time usage: 0.365/10.000 seconds Lua memory usage: 3885558/52428800 bytes Number of Wikibase entities loaded: 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