Positive-strand RNA virus

<h2 id="genome">Genome</h2>
Positive-strand RNA virus genomes usually contain relatively few genes, usually between three and ten, including an RNA-dependent RNA polymerase.<a class="footnote-ref" id="fnref:7" href="#fn:7">7</a> Coronaviruses have the largest known RNA genomes, between 27 and 32 <a href="/facts/Kilobase/xmPpPQ9W">kilobases</a> in length, and likely possess replication <a href="/facts/Proofreading_(biology)/Fn8W2EhA">proofreading</a> mechanisms in the form of an <a href="/facts/Exoribonuclease/PLGzWkXl">exoribonuclease</a> within <a href="/facts/Nonstructural_protein/Nxx8FepH">nonstructural protein</a> nsp14.<a class="footnote-ref" id="fnref:8" href="#fn:8">8</a>

<h2 id="replication">Replication</h2>

Positive-strand RNA viruses have genetic material that can function both as a <a href="/facts/Genome/31Sm08JT">genome</a> and as <a href="/facts/Messenger_RNA/oUL6qxQn">messenger RNA</a>; it can be directly <a href="/facts/Translation_(biology)/JcPC1rth">translated</a> into <a href="/facts/Protein/Jxmagu3E">protein</a> in the <a href="/facts/Host_cell/L6mop0kh">host cell</a> by host <a href="/facts/Ribosome/CF2u2gtg">ribosomes</a>.<a class="footnote-ref" id="fnref:9" href="#fn:9">9</a> The first proteins to be <a href="/facts/Gene_expression/alCPohLR">expressed</a> after infection serve genome replication functions; they recruit the positive-strand viral genome to <a href="/facts/Viral_replication/btE3fu1w">viral replication</a> complexes formed in association with intracellular membranes. These complexes contain proteins of both viral and host cell origin, and may be associated with the membranes of a variety of <a href="/facts/Organelle/vhDFnGbs">organelles</a>—often the <a href="/facts/Rough_endoplasmic_reticulum/DVbLA7GE">rough endoplasmic reticulum</a>,<a class="footnote-ref" id="fnref:10" href="#fn:10">10</a> but also including membranes derived from <a href="/facts/Mitochondria/ErHqrdJ1">mitochondria</a>, <a href="/facts/Vacuole/9QjBVb7j">vacuoles</a>, the <a href="/facts/Golgi_apparatus/F91pGb3t">Golgi apparatus</a>, <a href="/facts/Chloroplast/fo8U3yTx">chloroplasts</a>, <a href="/facts/Peroxisome/sEW1RBmb">peroxisomes</a>, <a href="/facts/Plasma_membrane/vAXSD0DT">plasma membranes</a>, <a href="/facts/Autophagosome/JoRbdUjU">autophagosomal membranes</a>, and novel <a href="/facts/Cytoplasm/Wa0sE6xf">cytoplasmic</a> compartments.<a class="footnote-ref" id="fnref:11" href="#fn:11">11</a>
The replication of the positive-sense RNA genome proceeds through <a href="/facts/Double-stranded_RNA/HxPeNfNj">double-stranded RNA</a> intermediates, and the purpose of replication in these membranous invaginations may be the avoidance of cellular response to the presence of dsRNA. In many cases <a href="/facts/Subgenomic/xyuLkPbJ">subgenomic</a> RNAs are also created during replication.<a class="footnote-ref" id="fnref:12" href="#fn:12">12</a> After infection, the entirety of the host cell's translation machinery may be diverted to the production of viral proteins as a result of the very high <a href="/facts/Affinity_(pharmacology)/44QLL4eU">affinity</a> for ribosomes by the viral genome's <a href="/facts/Internal_ribosome_entry_site/5lVqcu4I">internal ribosome entry site</a> (IRES) elements; in some viruses, such as <a href="/facts/Poliovirus/M7vyrYvS">poliovirus</a> and <a href="/facts/Rhinovirus/2jDL2oWU">rhinoviruses</a>, normal protein synthesis is further disrupted by viral <a href="/facts/Protease/G1ASdVat">proteases</a> degrading components required to initiate translation of cellular mRNA.<a class="footnote-ref" id="fnref:13" href="#fn:13">13</a>
All positive-strand RNA virus genomes encode <a href="/facts/RNA-dependent_RNA_polymerase/UgPn4rjz">RNA-dependent RNA polymerase</a>, a viral protein that synthesizes RNA from an RNA template. Host cell proteins recruited by +ssRNA viruses during replication include <a href="/facts/RNA-binding_protein/qTPTczLC">RNA-binding proteins</a>, <a href="/facts/Chaperone_protein/2SAjcVnW">chaperone proteins</a>, and membrane remodeling and <a href="/facts/Lipid_synthesis/wzC4nsn8">lipid synthesis</a> proteins, which collectively participate in exploiting the cell's <a href="/facts/Secretory_pathway/VHk9eLAV">secretory pathway</a> for viral replication.<a class="footnote-ref" id="fnref:14" href="#fn:14">14</a>

<h2 id="recombination">Recombination</h2>

Numerous positive-strand RNA viruses can undergo <a href="/facts/Genetic_recombination/uocNH0m7">genetic recombination</a> when at least two viral genomes are present in the same host cell.<a class="footnote-ref" id="fnref:15" href="#fn:15">15</a> The capability for recombination among +ssRNA virus pathogens of humans is common. RNA recombination appears to be a major driving force in determining genome architecture and the course of viral evolution among <a href="/facts/Picornavirus/QmeWDCDr">Picornaviridae</a> (e.g. poliovirus).<a class="footnote-ref" id="fnref:16" href="#fn:16">16</a> In the <a href="/facts/Retrovirus/GWAx3C3o">Retroviridae</a> (e.g. <a href="/facts/HIV/ulY2rgzu">HIV</a>), genome damage appears to be avoided during <a href="/facts/Reverse_transcriptase/Vc1lGk8P">reverse transcription</a> by strand switching, a form of recombination.<a class="footnote-ref" id="fnref:17" href="#fn:17">17</a><a class="footnote-ref" id="fnref:18" href="#fn:18">18</a><a class="footnote-ref" id="fnref:19" href="#fn:19">19</a> Recombination occurs in the <a href="/facts/Coronaviridae/NVjnbxYV">Coronaviridae</a> (e.g. <a href="/facts/Severe_acute_respiratory_syndrome/F5b7XSMS">SARS</a>).<a class="footnote-ref" id="fnref:20" href="#fn:20">20</a> Recombination in RNA viruses appears to be an adaptation for coping with genome damage.<a class="footnote-ref" id="fnref:21" href="#fn:21">21</a> Recombination can also occur infrequently between +ssRNA viruses of the same species but of divergent lineages. The resulting recombinant viruses may sometimes cause an outbreak of infection in humans, as in the case of SARS and MERS.<a class="footnote-ref" id="fnref:22" href="#fn:22">22</a>
Positive-strand RNA viruses are common in plants. In <a href="/facts/Tombusvirus/QsVCV9lm">tombusviruses</a> and <a href="/facts/Carmovirus/fnHULh78">carmoviruses</a>, RNA recombination occurs frequently during replication.<a class="footnote-ref" id="fnref:23" href="#fn:23">23</a> The ability of the RNA-dependent RNA polymerase of these viruses to switch RNA templates suggests a copy choice model of RNA recombination that may be an adaptive mechanism for coping with damage in the viral genome.<a class="footnote-ref" id="fnref:24" href="#fn:24">24</a> Other +ssRNA viruses of plants have also been reported to be capable of recombination, such as Brom mosaic <a href="/facts/Bromovirus/1cYzzJY2">bromovirus</a><a class="footnote-ref" id="fnref:25" href="#fn:25">25</a> and <a href="/facts/Sindbis_virus/9vQwqwSb">Sindbis virus</a>.<a class="footnote-ref" id="fnref:26" href="#fn:26">26</a>

<h2 id="classification">Classification</h2>

Positive-strand RNA viruses are found in three phyla: Kitrinoviricota, Lenarviricota, and Pisuviricota, each of which are assigned to the kingdom <a href="/facts/Orthornavirae/qG4evU2T">Orthornavirae</a> in the realm <a href="/facts/Riboviria/WBD92F1n">Riboviria</a>. In the <a href="/facts/Baltimore_classification/XTV7bbGJ">Baltimore classification</a> system, which groups viruses together based on their manner of mRNA synthesis, +ssRNA viruses are group IV.

<h3>Kitrinoviricota</h3>
The first +ssRNA phylum is <a href="/facts/Kitrinoviricota/x0ceeDAA">Kitrinoviricota</a>. The phylum contains what have been referred to as the "<a href="/facts/Alphavirus/3SkPGkEd">alphavirus</a> supergroup" and "<a href="/facts/Flavivirus/J1DqLnX8">flavivirus</a> supergroup" along with various other short-genome viruses. Four classes in the phylum are recognized: <a href="/facts/Alsuviricetes/wmkwPVdD">Alsuviricetes</a>, the alphavirus supergroup, which contains a large number of <a href="/facts/Plant_virus/gTtPbVCB">plant viruses</a> and arthropod viruses; Flasuviricetes, which contains flaviviruses, <a href="/facts/Magsaviricetes/pg6K6MKq">Magsaviricetes</a>, which contains <a href="/facts/Nodaviridae/KASEHmHJ">nodaviruses</a> and sinhaliviruses; and <a href="/facts/Tolucaviricetes/VplPwV2s">Tolucaviricetes</a>, which primarily contains plant viruses.<a class="footnote-ref" id="fnref:27" href="#fn:27">27</a><a class="footnote-ref" id="fnref:28" href="#fn:28">28</a>

<h3>Lenarviricota</h3>
<a href="/facts/Lenarviricota/kaHSjwQB">Lenarviricota</a> is the second +ssRNA phylum. It contains the class <a href="/facts/Leviviricetes/2Mu3poup">Leviviricetes</a>, which infect <a href="/facts/Prokaryote/oYyvaDDH">prokaryotes</a>, and the apparent descendants of leviviruses, which infect <a href="/facts/Eukaryote/SkwyPLMA">eukaryotes</a>. The phylum is divided into four classes: Leviviricetes, which contains leviviruses and their relatives, Amabiliviricetes, which contains <a href="/facts/Narnaviridae/39nU7yg9">narnaviruses</a> and their relatives, Howeltoviricetes, which contains <a href="/facts/Mitoviridae/trd2Rryc">mitoviruses</a> and their relatives, and Miaviricetes, which contains <a href="/facts/Botourmiaviridae/C5aRYDw2">botourmiaviruses</a> and their relatives. Based on phylogenetic analysis of RdRp, all other RNA viruses are considered to comprise a sister clade in relation to Lenarviricota.<a class="footnote-ref" id="fnref:29" href="#fn:29">29</a><a class="footnote-ref" id="fnref:30" href="#fn:30">30</a>

<h3>Pisuviricota</h3>

The third phylum that contains +ssRNA viruses is <a href="/facts/Pisuviricota/SPAcFwMe">Pisuviricota</a>, which has been informally called the "picornavirus supergroup". The phylum contains a large assemblage of eukaryotic viruses known to infect animals, plants, fungi, and protists. The phylum contains three classes, two of which contain only +ssRNA viruses: <a href="/facts/Pisoniviricetes/AU6dRE5G">Pisoniviricetes</a>, which contains <a href="/facts/Nidovirales/31HeBgDI">nidoviruses</a>, <a href="/facts/Picornavirales/02dhRELV">picornaviruses</a>, and <a href="/facts/Sobelivirales/eDRg5Mdv">sobeliviruses</a>, and <a href="/facts/Stelpaviricetes/jBpx6pvK">Stelpaviricetes</a>, which contains <a href="/facts/Potyviridae/lR4UMIMN">potyviruses</a> and <a href="/facts/Astroviridae/5vGTNb7e">astroviruses</a>. The third class is <a href="/facts/Duplopiviricetes/NYYmwuLI">Duplopiviricetes</a>, whose members are double-stranded RNA viruses that are descended from +ssRNA viruses.<a class="footnote-ref" id="fnref:31" href="#fn:31">31</a><a class="footnote-ref" id="fnref:32" href="#fn:32">32</a>

<h2 id="see-also">See also</h2>
<ul><li><a href="/facts/Double-stranded_RNA_virus/pF5FwT74">Double-stranded RNA virus</a></li>
<li><a href="/facts/Negative-strand_RNA_virus/PVAyRbNM">Negative-strand RNA virus</a></li>
<li><a href="/facts/Sense_(molecular_biology)/hcKgNvy6">Sense (molecular biology)</a></li></ul>

<h2 id="references">References</h2>

<ol>
<li id="fn:1">"Current ICTV Taxonomy Release | ICTV". ictv.global. Retrieved 3 April 2023. <a href="https://ictv.global/taxonomy" target="_blank">https://ictv.global/taxonomy</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></li>
<li id="fn:2">Baltimore D (September 1971). "Expression of animal virus genomes". Bacteriological Reviews. 35 (3): 235–41. doi:10.1128/MMBR.35.3.235-241.1971. PMC 378387. PMID 4329869. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC378387" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC378387</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></li>
<li id="fn:3">Lu R, Zhao X, Li J, Niu P, Yang B, Wu H, Wang W, Song H, Huang B, Zhu N, Bi Y, Ma X, Zhan F, Wang L, Hu T, Zhou H, Hu Z, Zhou W, Zhao L, Chen J, Meng Y, Wang J, Lin Y, Yuan J, Xie Z, Ma J, Liu WJ, Wang D, Xu W, Holmes EC, Gao GF, Wu G, Chen W, Shi W, Tan W (February 2020). "Genomic characterisation and epidemiology of 2019 novel coronavirus: implications for virus origins and receptor binding". Lancet. 395 (10224): 565–574. doi:10.1016/S0140-6736(20)30251-8. PMC 7159086. PMID 32007145. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7159086" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7159086</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></li>
<li id="fn:4">Nagy PD, Pogany J (December 2011). "The dependence of viral RNA replication on co-opted host factors". Nature Reviews. Microbiology. 10 (2): 137–149. doi:10.1038/nrmicro2692. PMC 7097227. PMID 22183253. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></li>
<li id="fn:5">Ahlquist P, Noueiry AO, Lee WM, Kushner DB, Dye BT (August 2003). "Host factors in positive-strand RNA virus genome replication". Journal of Virology. 77 (15): 8181–8186. doi:10.1128/JVI.77.15.8181-8186.2003. PMC 165243. PMID 12857886. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC165243" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC165243</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></li>
<li id="fn:6">Modrow S, Falke D, Truyen U, Schätzl H (2013). "Viruses with Single-Stranded, Positive-Sense RNA Genomes". Molecular Virology. Berlin, Heidelberg: Springer. pp. 185–349. doi:10.1007/978-3-642-20718-1_14. ISBN 978-3-642-20718-1. S2CID 82608215. <a href="978-3-642-20718-1" target="_blank">978-3-642-20718-1</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></li>
<li id="fn:7">Nagy PD, Pogany J (December 2011). "The dependence of viral RNA replication on co-opted host factors". Nature Reviews. Microbiology. 10 (2): 137–149. doi:10.1038/nrmicro2692. PMC 7097227. PMID 22183253. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></li>
<li id="fn:8">Smith EC, Denison MR (5 December 2013). "Coronaviruses as DNA wannabes: a new model for the regulation of RNA virus replication fidelity". PLOS Pathogens. 9 (12): e1003760. doi:10.1371/journal.ppat.1003760. PMC 3857799. PMID 24348241. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3857799" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3857799</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></li>
<li id="fn:9">"Positive stranded RNA virus replication". ViralZone. Retrieved 8 September 2016. <a href="http://viralzone.expasy.org/all_by_species/1116.html" target="_blank">http://viralzone.expasy.org/all_by_species/1116.html</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></li>
<li id="fn:10">Andronov L, Han M, Zhu Y, Balaji A, Roy AR, Barentine AE, Patel P, Garhyan J, Qi LS, Moerner WE (May 2024). "Nanoscale cellular organization of viral RNA and proteins in SARS-CoV-2 replication organelles". Nature Communications. 15 (1): 4644. Bibcode:2024NatCo..15.4644A. doi:10.1038/s41467-024-48991-x. PMC 11143195. PMID 38821943. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC11143195" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC11143195</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></li>
<li id="fn:11">Nagy PD, Pogany J (December 2011). "The dependence of viral RNA replication on co-opted host factors". Nature Reviews. Microbiology. 10 (2): 137–149. doi:10.1038/nrmicro2692. PMC 7097227. PMID 22183253. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></li>
<li id="fn:12">"Positive stranded RNA virus replication". ViralZone. Retrieved 8 September 2016. <a href="http://viralzone.expasy.org/all_by_species/1116.html" target="_blank">http://viralzone.expasy.org/all_by_species/1116.html</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></li>
<li id="fn:13">Modrow S, Falke D, Truyen U, Schätzl H (2013). "Viruses with Single-Stranded, Positive-Sense RNA Genomes". Molecular Virology. Berlin, Heidelberg: Springer. pp. 185–349. doi:10.1007/978-3-642-20718-1_14. ISBN 978-3-642-20718-1. S2CID 82608215. <a href="978-3-642-20718-1" target="_blank">978-3-642-20718-1</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></li>
<li id="fn:14">Nagy PD, Pogany J (December 2011). "The dependence of viral RNA replication on co-opted host factors". Nature Reviews. Microbiology. 10 (2): 137–149. doi:10.1038/nrmicro2692. PMC 7097227. PMID 22183253. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7097227</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></li>
<li id="fn:15">Barr JN, Fearns R (June 2010). "How RNA viruses maintain their genome integrity". The Journal of General Virology. 91 (Pt 6): 1373–87. doi:10.1099/vir.0.020818-0. PMID 20335491. <a href="https://doi.org/10.1099%2Fvir.0.020818-0" target="_blank">https://doi.org/10.1099%2Fvir.0.020818-0</a> <a href="#fnref:15" class="footnote-back-ref">↩</a></li>
<li id="fn:16">Muslin C, Mac Kain A, Bessaud M, Blondel B, Delpeyroux F (September 2019). "Recombination in Enteroviruses, a Multi-Step Modular Evolutionary Process". Viruses. 11 (9): 859. doi:10.3390/v11090859. PMC 6784155. PMID 31540135. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6784155" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6784155</a> <a href="#fnref:16" class="footnote-back-ref">↩</a></li>
<li id="fn:17">Hu WS, Temin HM (November 1990). "Retroviral recombination and reverse transcription". Science. 250 (4985): 1227–33. Bibcode:1990Sci...250.1227H. doi:10.1126/science.1700865. PMID 1700865. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:17" class="footnote-back-ref">↩</a></li>
<li id="fn:18">Rawson JM, Nikolaitchik OA, Keele BF, Pathak VK, Hu WS (November 2018). "Recombination is required for efficient HIV-1 replication and the maintenance of viral genome integrity". Nucleic Acids Research. 46 (20): 10535–10545. doi:10.1093/nar/gky910. PMC 6237782. PMID 30307534. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6237782" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6237782</a> <a href="#fnref:18" class="footnote-back-ref">↩</a></li>
<li id="fn:19">Bernstein H, Bernstein C, Michod RE (January 2018). "Sex in microbial pathogens". Infection, Genetics and Evolution. 57: 8–25. Bibcode:2018InfGE..57....8B. doi:10.1016/j.meegid.2017.10.024. PMID 29111273. <a href="https://doi.org/10.1016%2Fj.meegid.2017.10.024" target="_blank">https://doi.org/10.1016%2Fj.meegid.2017.10.024</a> <a href="#fnref:19" class="footnote-back-ref">↩</a></li>
<li id="fn:20">Su S, Wong G, Shi W, Liu J, Lai AC, Zhou J, et al. (June 2016). "Epidemiology, Genetic Recombination, and Pathogenesis of Coronaviruses". Trends in Microbiology. 24 (6): 490–502. doi:10.1016/j.tim.2016.03.003. PMC 7125511. PMID 27012512. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7125511" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7125511</a> <a href="#fnref:20" class="footnote-back-ref">↩</a></li>
<li id="fn:21">Barr JN, Fearns R (June 2010). "How RNA viruses maintain their genome integrity". The Journal of General Virology. 91 (Pt 6): 1373–87. doi:10.1099/vir.0.020818-0. PMID 20335491. <a href="https://doi.org/10.1099%2Fvir.0.020818-0" target="_blank">https://doi.org/10.1099%2Fvir.0.020818-0</a> <a href="#fnref:21" class="footnote-back-ref">↩</a></li>
<li id="fn:22">Su S, Wong G, Shi W, Liu J, Lai AC, Zhou J, et al. (June 2016). "Epidemiology, Genetic Recombination, and Pathogenesis of Coronaviruses". Trends in Microbiology. 24 (6): 490–502. doi:10.1016/j.tim.2016.03.003. PMC 7125511. PMID 27012512. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7125511" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7125511</a> <a href="#fnref:22" class="footnote-back-ref">↩</a></li>
<li id="fn:23">Cheng CP, Nagy PD (November 2003). "Mechanism of RNA recombination in carmo- and tombusviruses: evidence for template switching by the RNA-dependent RNA polymerase in vitro". Journal of Virology. 77 (22): 12033–47. doi:10.1128/jvi.77.22.12033-12047.2003. PMC 254248. PMID 14581540. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC254248" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC254248</a> <a href="#fnref:23" class="footnote-back-ref">↩</a></li>
<li id="fn:24">Cheng CP, Nagy PD (November 2003). "Mechanism of RNA recombination in carmo- and tombusviruses: evidence for template switching by the RNA-dependent RNA polymerase in vitro". Journal of Virology. 77 (22): 12033–47. doi:10.1128/jvi.77.22.12033-12047.2003. PMC 254248. PMID 14581540. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC254248" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC254248</a> <a href="#fnref:24" class="footnote-back-ref">↩</a></li>
<li id="fn:25">Kolondam B, Rao P, Sztuba-Solinska J, Weber PH, Dzianott A, Johns MA, Bujarski JJ (2015). "Co-infection with two strains of Brome mosaic bromovirus reveals common RNA recombination sites in different hosts". Virus Evolution. 1 (1): vev021. doi:10.1093/ve/vev021. PMC 5014487. PMID 27774290. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5014487" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5014487</a> <a href="#fnref:25" class="footnote-back-ref">↩</a></li>
<li id="fn:26">Weiss BG, Schlesinger S (August 1991). "Recombination between Sindbis virus RNAs". Journal of Virology. 65 (8): 4017–25. doi:10.1128/JVI.65.8.4017-4025.1991. PMC 248832. PMID 2072444. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC248832" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC248832</a> <a href="#fnref:26" class="footnote-back-ref">↩</a></li>
<li id="fn:27">Koonin EV, Dolja VV, Krupovic M, Varsani A, Wolf YI, Yutin N, Zerbini M, Kuhn JH (18 October 2019). "Create a megataxonomic framework, filling all principal taxonomic ranks, for realm Riboviria" (docx). International Committee on Taxonomy of Viruses (ICTV). Retrieved 14 August 2020. <a href="https://ictv.global/ictv/proposals/2019.006G.zip" target="_blank">https://ictv.global/ictv/proposals/2019.006G.zip</a> <a href="#fnref:27" class="footnote-back-ref">↩</a></li>
<li id="fn:28">Wolf YI, Kazlauskas D, Iranzo J, Lucia-Sanz A, Kuhn JH, Krupovic M, Dolja VV, Koonin EV (27 November 2018). "Origins and Evolution of the Global RNA Virome". mBio. 9 (6): e02329-18. doi:10.1128/mBio.02329-18. PMC 6282212. PMID 30482837. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6282212" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6282212</a> <a href="#fnref:28" class="footnote-back-ref">↩</a></li>
<li id="fn:29">Koonin EV, Dolja VV, Krupovic M, Varsani A, Wolf YI, Yutin N, Zerbini M, Kuhn JH (18 October 2019). "Create a megataxonomic framework, filling all principal taxonomic ranks, for realm Riboviria" (docx). International Committee on Taxonomy of Viruses (ICTV). Retrieved 14 August 2020. <a href="https://ictv.global/ictv/proposals/2019.006G.zip" target="_blank">https://ictv.global/ictv/proposals/2019.006G.zip</a> <a href="#fnref:29" class="footnote-back-ref">↩</a></li>
<li id="fn:30">Wolf YI, Kazlauskas D, Iranzo J, Lucia-Sanz A, Kuhn JH, Krupovic M, Dolja VV, Koonin EV (27 November 2018). "Origins and Evolution of the Global RNA Virome". mBio. 9 (6): e02329-18. doi:10.1128/mBio.02329-18. PMC 6282212. PMID 30482837. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6282212" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6282212</a> <a href="#fnref:30" class="footnote-back-ref">↩</a></li>
<li id="fn:31">Koonin EV, Dolja VV, Krupovic M, Varsani A, Wolf YI, Yutin N, Zerbini M, Kuhn JH (18 October 2019). "Create a megataxonomic framework, filling all principal taxonomic ranks, for realm Riboviria" (docx). International Committee on Taxonomy of Viruses (ICTV). Retrieved 14 August 2020. <a href="https://ictv.global/ictv/proposals/2019.006G.zip" target="_blank">https://ictv.global/ictv/proposals/2019.006G.zip</a> <a href="#fnref:31" class="footnote-back-ref">↩</a></li>
<li id="fn:32">Wolf YI, Kazlauskas D, Iranzo J, Lucia-Sanz A, Kuhn JH, Krupovic M, Dolja VV, Koonin EV (27 November 2018). "Origins and Evolution of the Global RNA Virome". mBio. 9 (6): e02329-18. doi:10.1128/mBio.02329-18. PMC 6282212. PMID 30482837. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6282212" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6282212</a> <a href="#fnref:32" class="footnote-back-ref">↩</a></li>
</ol>

Positive-strand RNA virus open-in-new

Positive-strand RNA virus