{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,12]],"date-time":"2025-10-12T02:33:06Z","timestamp":1760236386408,"version":"build-2065373602"},"reference-count":55,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2021,11,23]],"date-time":"2021-11-23T00:00:00Z","timestamp":1637625600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32072836","31702244"],"award-info":[{"award-number":["32072836","31702244"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100012543","name":"Shanghai Science and Technology Development Foundation","doi-asserted-by":"publisher","award":["18391902200","21N41900100"],"award-info":[{"award-number":["18391902200","21N41900100"]}],"id":[{"id":"10.13039\/100012543","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Shanghai Key Laboratory of Veterinary Biotechnology and Key Laboratory of Urban Agriculture","award":["klab201607"],"award-info":[{"award-number":["klab201607"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>Reassortant variant viruses generated between 2009 H1N1 pandemic influenza virus [A(H1N1)pdm09] and endemic swine influenza viruses posed a potential risk to humans. Surprisingly, genetic analysis showed that almost all of these variant viruses contained the M segment from A(H1N1)pdm09, which originated from Eurasian avian-like swine influenza viruses. Studies have shown that the A(H1N1)pdm09 M gene is critical for the transmissibility and pathogenicity of the variant viruses. However, the M gene encodes two proteins, M1 and M2, and which of those plays a more important role in virus pathogenicity remains unknown. In this study, the M1 and M2 genes of A(H1N1)pdm09 were replaced with those of endemic H3N2 swine influenza virus, respectively. The chimeric viruses were rescued and evaluated in vitro and in mice. Both M1 and M2 of H3N2 affected the virus replication in vitro. In mice, the introduction of H3N2 M1 attenuated the chimeric virus, where all the mice survived from the infection, compared with the wild type virus that caused 100 % mortality. However, the chimeric virus containing H3N2 M2 was still virulent to mice, and caused 16.6% mortality, as well as similar body weight loss to the wild type virus infected group. Compared with the wild type virus, the chimeric virus containing H3N2 M1 induced lower levels of inflammatory cytokines and higher levels of anti-inflammatory cytokines, whereas the chimeric virus containing H3N2 M2 induced substantial pro-inflammatory responses, but higher levels of anti-inflammatory cytokines. The study demonstrated that Eurasian avian-like M1 played a more important role than M2 in the pathogenicity of A(H1N1)pdm09 in mice.<\/jats:p>","DOI":"10.3390\/v13122335","type":"journal-article","created":{"date-parts":[[2021,12,1]],"date-time":"2021-12-01T03:12:56Z","timestamp":1638328376000},"page":"2335","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Eurasian Avian-like M1 Plays More Important Role than M2 in Pathogenicity of 2009 Pandemic H1N1 Influenza Virus in Mice"],"prefix":"10.3390","volume":"13","author":[{"given":"Lixiang","family":"Xie","sequence":"first","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Guanlong","family":"Xu","sequence":"additional","affiliation":[{"name":"China Institute of Veterinary Drug Control, Beijing 100081, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lingxiang","family":"Xin","sequence":"additional","affiliation":[{"name":"China Institute of Veterinary Drug Control, Beijing 100081, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zhaofei","family":"Wang","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"},{"name":"Shanghai Key Laboratory of Veterinary Biotechnology, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rujuan","family":"Wu","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mingqing","family":"Wu","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuqiang","family":"Cheng","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"},{"name":"Shanghai Key Laboratory of Veterinary Biotechnology, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hengan","family":"Wang","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"},{"name":"Shanghai Key Laboratory of Veterinary Biotechnology, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yaxian","family":"Yan","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"},{"name":"Shanghai Key Laboratory of Veterinary Biotechnology, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1187-7249","authenticated-orcid":false,"given":"Jingjiao","family":"Ma","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"},{"name":"Shanghai Key Laboratory of Veterinary Biotechnology, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jianhe","family":"Sun","sequence":"additional","affiliation":[{"name":"School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai 200240, China"},{"name":"Shanghai Key Laboratory of Veterinary Biotechnology, Shanghai 200240, China"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2021,11,23]]},"reference":[{"key":"ref_1","unstructured":"WHO (2009). Influenza A(H1N1)\u2014Update 61, WHO."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"S44","DOI":"10.1093\/cid\/ciq010","article-title":"Influenza pandemic epidemiologic and virologic diversity: Reminding ourselves of the possibilities","volume":"52","author":"Kasowski","year":"2011","journal-title":"Clin. Infect. Dis."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1111\/irv.12515","article-title":"Mortality burden from seasonal influenza and 2009 H1N1 pandemic influenza in Beijing, China, 2007\u20132013","volume":"12","author":"Wu","year":"2018","journal-title":"Influenza Other Respir. Viruses"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1126\/science.1176225","article-title":"Antigenic and genetic characteristics of swine-origin 2009 A(H1N1) influenza viruses circulating in humans","volume":"325","author":"Garten","year":"2009","journal-title":"Science"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1093\/bja\/aep375","article-title":"Pandemic (H1N1) 2009 influenza","volume":"104","author":"Patel","year":"2010","journal-title":"Br. J. Anaesth."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1099\/vir.0.037358-0","article-title":"Natural and experimental infection of dogs with pandemic H1N1\/2009 influenza virus","volume":"93","author":"Lin","year":"2012","journal-title":"J. Gen. Virol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1186\/1743-422X-7-27","article-title":"Susceptibility of turkeys to pandemic-H1N1 virus by reproductive tract insemination","volume":"7","author":"Wasilenko","year":"2010","journal-title":"Virol. J."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1111\/j.1863-2378.2011.01406.x","article-title":"Influenza A pandemic (H1N1) 2009 virus outbreak in a cat colony in Italy","volume":"58","author":"Fiorentini","year":"2011","journal-title":"Zoonoses Public Health"},{"key":"ref_9","first-page":"1153","article-title":"An investigation into human pandemic influenza virus (H1N1) 2009 on an Alberta swine farm","volume":"50","author":"Howden","year":"2009","journal-title":"Can. Vet. J."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"747","DOI":"10.3201\/eid1704.101355","article-title":"Pandemic (H1N1) 2009 virus in 3 wildlife species, San Diego, California, USA","volume":"17","author":"Schrenzel","year":"2011","journal-title":"Emerg. Infect. Dis."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"642","DOI":"10.1136\/vr.4851","article-title":"Initial incursion of pandemic (H1N1) 2009 influenza A virus into European pigs","volume":"166","author":"Welsh","year":"2010","journal-title":"Vet. Rec."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1587","DOI":"10.3201\/eid1610.100665","article-title":"Pandemic (H1N1) 2009 virus on commercial swine farm, Thailand","volume":"16","author":"Sreta","year":"2010","journal-title":"Emerg. Infect. Dis."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1624","DOI":"10.3201\/1709.110338","article-title":"Multiple reassortment between pandemic (H1N1) 2009 and endemic influenza viruses in pigs, United States","volume":"17","author":"Ducatez","year":"2011","journal-title":"Emerg. Infect. Dis."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"S4","DOI":"10.1093\/cid\/cit272","article-title":"Human infections with influenza A(H3N2) variant virus in the United States, 2011\u20132012","volume":"57","author":"Epperson","year":"2013","journal-title":"Clin. Infect. Dis."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1703","DOI":"10.1093\/cid\/cit649","article-title":"Outbreak of variant influenza A(H3N2) virus in the United States","volume":"57","author":"Jhung","year":"2013","journal-title":"Clin. Infect. Dis."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"2605","DOI":"10.1056\/NEJMoa0903810","article-title":"Emergence of a novel swine-origin influenza A (H1N1) virus in humans","volume":"360","author":"Dawood","year":"2009","journal-title":"N. Engl. J. Med."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2616","DOI":"10.1056\/NEJMoa0903812","article-title":"Triple-reassortant swine influenza A (H1) in humans in the United States, 2005\u20132009","volume":"360","author":"Shinde","year":"2009","journal-title":"N. Engl. J. Med."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1038\/s41426-018-0088-z","article-title":"Novel triple-reassortant influenza viruses in pigs, Guangxi, China","volume":"7","author":"He","year":"2018","journal-title":"Emerg. Microbes Infect."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"e01763-16","DOI":"10.1128\/JVI.01763-16","article-title":"Reassortment between Swine H3N2 and 2009 Pandemic H1N1 in the United States Resulted in Influenza A Viruses with Diverse Genetic Constellations with Variable Virulence in Pigs","volume":"91","author":"Rajao","year":"2017","journal-title":"J. Virol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Lakdawala, S.S., Lamirande, E.W., Suguitan, A.L., Wang, W., Santos, C.P., Vogel, L., Matsuoka, Y., Lindsley, W.G., Jin, H., and Subbarao, K. (2011). Eurasian-origin gene segments contribute to the transmissibility, aerosol release, and morphology of the 2009 pandemic H1N1 influenza virus. PLoS Pathog., 7.","DOI":"10.1371\/journal.ppat.1002443"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"11235","DOI":"10.1128\/JVI.05794-11","article-title":"The M segment of the 2009 new pandemic H1N1 influenza virus is critical for its high transmission efficiency in the guinea pig model","volume":"85","author":"Chou","year":"2011","journal-title":"J. Virol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Calderon, B.M., Danzy, S., Delima, G.K., Jacobs, N.T., Ganti, K., Hockman, M.R., Conn, G.L., Lowen, A.C., and Steel, J. (2019). Dysregulation of M segment gene expression contributes to influenza A virus host restriction. PLoS Pathog., 15.","DOI":"10.1101\/599886"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1016\/j.virol.2003.12.009","article-title":"The M1 matrix protein controls the filamentous phenotype of influenza A virus","volume":"321","author":"Elleman","year":"2004","journal-title":"Virology"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"7569","DOI":"10.1128\/JVI.00119-14","article-title":"Residue 41 of the Eurasian avian-like swine influenza a virus matrix protein modulates virion filament length and efficiency of contact transmission","volume":"88","author":"Campbell","year":"2014","journal-title":"J. Virol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"3802","DOI":"10.1128\/JVI.03607-13","article-title":"The M segment of the 2009 pandemic influenza virus confers increased neuraminidase activity, filamentous morphology, and efficient contact transmissibility to A\/Puerto Rico\/8\/1934-based reassortant viruses","volume":"88","author":"Campbell","year":"2014","journal-title":"J. Virol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1113\/jphysiol.1996.sp021495","article-title":"Selective proton permeability and pH regulation of the influenza virus M2 channel expressed in mouse erythroleukaemia cells","volume":"494","author":"Chizhmakov","year":"1996","journal-title":"J. Physiol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1038\/nature06531","article-title":"Structure and mechanism of the M2 proton channel of influenza A virus","volume":"451","author":"Schnell","year":"2008","journal-title":"Nature"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"13029","DOI":"10.1128\/JVI.01430-14","article-title":"Stepwise priming by acidic pH and a high K+ concentration is required for efficient uncoating of influenza A virus cores after penetration","volume":"88","author":"Stauffer","year":"2014","journal-title":"J. Virol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1016\/j.virol.2010.12.003","article-title":"Influenza virus assembly and budding","volume":"411","author":"Rossman","year":"2011","journal-title":"Virology"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/j.virol.2005.09.049","article-title":"Influenza virus assembly and budding in raft-derived microdomains: A quantitative analysis of the surface distribution of HA, NA and M2 proteins","volume":"342","author":"Leser","year":"2005","journal-title":"Virology"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"11538","DOI":"10.1128\/JVI.74.24.11538-11547.2000","article-title":"Influenza virus matrix protein is the major driving force in virus budding","volume":"74","author":"Albo","year":"2000","journal-title":"J. Virol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"902","DOI":"10.1016\/j.cell.2010.08.029","article-title":"Influenza virus M2 protein mediates ESCRT-independent membrane scission","volume":"142","author":"Rossman","year":"2010","journal-title":"Cell"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1425","DOI":"10.1042\/BST20140136","article-title":"Alterations of membrane curvature during influenza virus budding","volume":"42","author":"Martyna","year":"2014","journal-title":"Biochem. Soc. Trans."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"2226","DOI":"10.1128\/JVI.01565-12","article-title":"Virulence determinants of pandemic A(H1N1)2009 influenza virus in a mouse model","volume":"87","author":"Uraki","year":"2013","journal-title":"J. Virol."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Kiso, M., Shinya, K., Shimojima, M., Takano, R., Takahashi, K., Katsura, H., Kakugawa, S., Le, M.T., Yamashita, M., and Furuta, Y. (2010). Characterization of oseltamivir-resistant 2009 H1N1 pandemic influenza A viruses. PLoS Pathog., 6.","DOI":"10.1371\/journal.ppat.1001079"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2314","DOI":"10.1099\/vir.0.021402-0","article-title":"Viral reassortment and transmission after co-infection of pigs with classical H1N1 and triple-reassortant H3N2 swine influenza viruses","volume":"91","author":"Ma","year":"2010","journal-title":"J. Gen. Virol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2275","DOI":"10.1007\/s007050170002","article-title":"Universal primer set for the full-length amplification of all influenza A viruses","volume":"146","author":"Hoffmann","year":"2001","journal-title":"Arch. Virol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.virol.2017.01.015","article-title":"Impacts of different expressions of PA-X protein on 2009 pandemic H1N1 virus replication, pathogenicity and host immune responses","volume":"504","author":"Lee","year":"2017","journal-title":"Virology"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"3198","DOI":"10.1128\/JCM.41.7.3198-3205.2003","article-title":"Pathogenic and antigenic properties of phylogenetically distinct reassortant H3N2 swine influenza viruses cocirculating in the United States","volume":"41","author":"Richt","year":"2003","journal-title":"J. Clin. Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Zhang, Y., Zhu, J., Li, Y., Bradley, K.C., Cao, J., Chen, H., Jin, M., and Zhou, H. (2013). Glycosylation on hemagglutinin affects the virulence and pathogenicity of pandemic H1N1\/2009 influenza A virus in mice. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0061397"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"10","DOI":"10.30699\/ijp.13.1.10","article-title":"The Modulatory Effects of Vitamin D on the Expression of IL-12 and TGF-beta in the Spinal Cord and Serum of Mice with Experimental Autoimmune Encephalomyelitis","volume":"13","author":"Azizi","year":"2018","journal-title":"Iran. J. Pathol."},{"key":"ref_42","unstructured":"CDC (2019). 2009 H1N1 Pandemic Timeline."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1111\/irv.12559","article-title":"Regional patterns of genetic diversity in swine influenza A viruses in the United States from 2010 to 2016","volume":"13","author":"Walia","year":"2019","journal-title":"Influenza Other Respir. Viruses"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1016\/j.virusres.2013.04.012","article-title":"Reassortment ability of the 2009 pandemic H1N1 influenza virus with circulating human and avian influenza viruses: Public health risk implications","volume":"175","author":"Stincarelli","year":"2013","journal-title":"Virus Res."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1261","DOI":"10.1099\/vir.0.040535-0","article-title":"The neuraminidase and matrix genes of the 2009 pandemic influenza H1N1 virus cooperate functionally to facilitate efficient replication and transmissibility in pigs","volume":"93","author":"Ma","year":"2012","journal-title":"J. Gen. Virol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e00095-18","DOI":"10.1128\/JVI.00095-18","article-title":"Antigenically Diverse Swine Origin H1N1 Variant Influenza Viruses Exhibit Differential Ferret Pathogenesis and Transmission Phenotypes","volume":"92","author":"Jones","year":"2018","journal-title":"J. Virol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"e01019-21","DOI":"10.1128\/JVI.01019-21","article-title":"Key amino acids of M1-41 and M2-27 determine growth and pathogenicity of chimeric H17 bat influenza virus in cells and in mice","volume":"95","author":"Yang","year":"2021","journal-title":"J. Virol."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Bialas, K.M., Desmet, E.A., and Takimoto, T. (2012). Specific residues in the 2009 H1N1 swine-origin influenza matrix protein influence virion morphology and efficiency of viral spread in vitro. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0050595"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1023\/A:1008131312306","article-title":"Genomic analysis of matrix gene and antigenic studies of its gene product (M1) of a swine influenza virus (H1N1) causing chronic respiratory disease in pigs","volume":"21","author":"Welman","year":"2000","journal-title":"Virus Genes"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"10059","DOI":"10.1128\/JVI.01184-08","article-title":"The influenza virus M2 protein cytoplasmic tail interacts with the M1 protein and influences virus assembly at the site of virus budding","volume":"82","author":"Chen","year":"2008","journal-title":"J. Virol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1038\/nrneph.2016.41","article-title":"Role of TLRs and DAMPs in allograft inflammation and transplant outcomes","volume":"12","author":"Braza","year":"2016","journal-title":"Nat. Rev. Nephrol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1128\/CMR.00046-08","article-title":"Pathogen recognition and inflammatory signaling in innate immune defenses","volume":"22","author":"Mogensen","year":"2009","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1923","DOI":"10.1093\/infdis\/jiv330","article-title":"A Newly Emerged Swine-Origin Influenza A(H3N2) Variant Dampens Host Antiviral Immunity but Induces Potent Inflammasome Activation","volume":"212","author":"Cao","year":"2015","journal-title":"J. Infect. Dis."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1163","DOI":"10.1080\/15548627.2019.1580089","article-title":"Influenza M2 protein regulates MAVS-mediated signaling pathway through interacting with MAVS and increasing ROS production","volume":"15","author":"Wang","year":"2019","journal-title":"Autophagy"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"12341","DOI":"10.1128\/JVI.01725-12","article-title":"Mutations in the M-gene segment can substantially increase replication efficiency of NS1 deletion influenza A virus in MDCK cells","volume":"86","author":"Harmsen","year":"2012","journal-title":"J. Virol."}],"container-title":["Viruses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1999-4915\/13\/12\/2335\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T07:34:24Z","timestamp":1760168064000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1999-4915\/13\/12\/2335"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,11,23]]},"references-count":55,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["v13122335"],"URL":"https:\/\/doi.org\/10.3390\/v13122335","relation":{},"ISSN":["1999-4915"],"issn-type":[{"type":"electronic","value":"1999-4915"}],"subject":[],"published":{"date-parts":[[2021,11,23]]}}}