{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T10:54:32Z","timestamp":1781607272890,"version":"3.54.5"},"reference-count":42,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2024,1,1]],"date-time":"2024-01-01T00:00:00Z","timestamp":1704067200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2024,1,1]],"date-time":"2024-01-01T00:00:00Z","timestamp":1704067200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,1,1]],"date-time":"2025-01-01T00:00:00Z","timestamp":1735689600000},"content-version":"vor","delay-in-days":366,"URL":"http:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"},{"start":{"date-parts":[[2024,1,1]],"date-time":"2024-01-01T00:00:00Z","timestamp":1704067200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2024,1,1]],"date-time":"2024-01-01T00:00:00Z","timestamp":1704067200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2024,1,1]],"date-time":"2024-01-01T00:00:00Z","timestamp":1704067200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2024,1,1]],"date-time":"2024-01-01T00:00:00Z","timestamp":1704067200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2024,1,1]],"date-time":"2024-01-01T00:00:00Z","timestamp":1704067200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32160825"],"award-info":[{"award-number":["32160825"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004607","name":"Guangxi Natural Science Foundation","doi-asserted-by":"publisher","award":["2023GXNSFDA026042"],"award-info":[{"award-number":["2023GXNSFDA026042"]}],"id":[{"id":"10.13039\/501100004607","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["clinicalkey.fr","clinicalkey.jp","clinicalkey.com.au","clinicalkey.es","clinicalkey.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Virology"],"published-print":{"date-parts":[[2024,1]]},"DOI":"10.1016\/j.virol.2023.109927","type":"journal-article","created":{"date-parts":[[2023,11,7]],"date-time":"2023-11-07T18:41:09Z","timestamp":1699382469000},"page":"109927","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":3,"special_numbering":"C","title":["Reassortant H9N2 canine influenza viruses containing the pandemic H1N1\/2009 ribonucleoprotein complex circulating in pigs acquired enhanced virulence in mice"],"prefix":"10.1016","volume":"589","author":[{"given":"Min","family":"Zhu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hao","family":"Zeng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianqiao","family":"He","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yaohui","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pingping","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianing","family":"Guo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinfan","family":"Guo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huabo","family":"Zhou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yifeng","family":"Qin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kang","family":"Ouyang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zuzhang","family":"Wei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weijian","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6720-353X","authenticated-orcid":false,"given":"Ying","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.virol.2023.109927_bib1","doi-asserted-by":"crossref","first-page":"5909","DOI":"10.1038\/s41467-020-19671-3","article-title":"Dominant subtype switch in avian influenza viruses during 2016-2019 in China","volume":"11","author":"Bi","year":"2020","journal-title":"Nat. Commun."},{"key":"10.1016\/j.virol.2023.109927_bib2","doi-asserted-by":"crossref","first-page":"714","DOI":"10.1016\/S0140-6736(14)60111-2","article-title":"Clinical and epidemiological characteristics of a fatal case of avian influenza A H10N8 virus infection: a descriptive study","volume":"383","author":"Chen","year":"2014","journal-title":"Lancet"},{"key":"10.1016\/j.virol.2023.109927_bib3","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00909-18","article-title":"Emergence and evolution of novel reassortant influenza A viruses in canines in southern China","volume":"9","author":"Chen","year":"2018","journal-title":"mBio"},{"key":"10.1016\/j.virol.2023.109927_bib4","doi-asserted-by":"crossref","first-page":"2035","DOI":"10.1099\/vir.0.82783-0","article-title":"Antigenic and genetic characterization of H9N2 swine influenza viruses in China","volume":"88","author":"Cong","year":"2007","journal-title":"J. Gen. Virol."},{"key":"10.1016\/j.virol.2023.109927_bib5","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1007\/s11262-008-0227-z","article-title":"Swine infection with H9N2 influenza viruses in China in 2004","volume":"36","author":"Cong","year":"2008","journal-title":"Virus Gene."},{"key":"10.1016\/j.virol.2023.109927_bib6","doi-asserted-by":"crossref","first-page":"18590","DOI":"10.1073\/pnas.0507415102","article-title":"The viral polymerase mediates adaptation of an avian influenza virus to a mammalian host","volume":"102","author":"Gabriel","year":"2005","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.virol.2023.109927_bib7","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":"10.1016\/j.virol.2023.109927_bib8","doi-asserted-by":"crossref","first-page":"9363","DOI":"10.1073\/pnas.96.16.9363","article-title":"Molecular characterization of H9N2 influenza viruses: were they the donors of the \"internal\" genes of H5N1 viruses in Hong Kong?","volume":"96","author":"Guan","year":"1999","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.virol.2023.109927_bib9","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. Microb. Infect."},{"key":"10.1016\/j.virol.2023.109927_bib10","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1007\/s00430-018-0548-z","article-title":"PA-X: a key regulator of influenza A virus pathogenicity and host immune responses","volume":"207","author":"Hu","year":"2018","journal-title":"Med. Microbiol. Immunol."},{"key":"10.1016\/j.virol.2023.109927_bib11","doi-asserted-by":"crossref","first-page":"12084","DOI":"10.1073\/pnas.1108058108","article-title":"Compatibility of H9N2 avian influenza surface genes and 2009 pandemic H1N1 internal genes for transmission in the ferret model","volume":"108","author":"Kimble","year":"2011","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.virol.2023.109927_bib12","doi-asserted-by":"crossref","first-page":"1611","DOI":"10.1128\/JVI.01491-08","article-title":"Influenza a virus polymerase is an integral component of the CPSF30-NS1A protein complex in infected cells","volume":"83","author":"Kuo","year":"2009","journal-title":"J. Virol."},{"key":"10.1016\/j.virol.2023.109927_bib13","doi-asserted-by":"crossref","first-page":"142","DOI":"10.1007\/s11262-015-1272-z","article-title":"Isolation of a novel H3N2 influenza virus containing a gene of H9N2 avian influenza in a dog in South Korea in 2015","volume":"52","author":"Lee","year":"2016","journal-title":"Virus Gene."},{"key":"10.1016\/j.virol.2023.109927_bib14","first-page":"185","article-title":"Enhancement of influenza virus transmission by gene reassortment","volume":"385","author":"Li","year":"2014","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"10.1016\/j.virol.2023.109927_bib15","doi-asserted-by":"crossref","DOI":"10.1101\/cshperspect.a038349","article-title":"H7N9 influenza virus in China","volume":"11","author":"Li","year":"2021","journal-title":"Cold Spring Harb Perspect Med"},{"key":"10.1016\/j.virol.2023.109927_bib16","doi-asserted-by":"crossref","first-page":"e187","DOI":"10.1016\/j.jinf.2022.08.033","article-title":"Human infection with H3N8 avian influenza virus: a novel H9N2-original reassortment virus","volume":"85","author":"Li","year":"2022","journal-title":"J. Infect."},{"key":"10.1016\/j.virol.2023.109927_bib17","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1004508","article-title":"Genetics, receptor binding property, and transmissibility in mammals of naturally isolated H9N2 Avian Influenza viruses","volume":"10","author":"Li","year":"2014","journal-title":"PLoS Pathog."},{"key":"10.1016\/j.virol.2023.109927_bib18","doi-asserted-by":"crossref","first-page":"1024","DOI":"10.1007\/s11427-020-1981-5","article-title":"Emergence of a novel reassortant avian influenza virus (H10N3) in Eastern China with high pathogenicity and respiratory droplet transmissibility to mammals","volume":"65","author":"Liu","year":"2022","journal-title":"Sci. China Life Sci."},{"key":"10.1016\/j.virol.2023.109927_bib19","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1128\/JVI.05699-11","article-title":"Combination of PB2 271A and SR polymorphism at positions 590\/591 is critical for viral replication and virulence of swine influenza virus in cultured cells and in vivo","volume":"86","author":"Liu","year":"2012","journal-title":"J. Virol."},{"key":"10.1016\/j.virol.2023.109927_bib20","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1038\/s41579-018-0115-z","article-title":"Host and viral determinants of influenza A virus species specificity","volume":"17","author":"Long","year":"2019","journal-title":"Nat. Rev. Microbiol."},{"key":"10.1016\/j.virol.2023.109927_bib21","doi-asserted-by":"crossref","first-page":"1323","DOI":"10.1038\/s41598-017-01512-x","article-title":"A reassortant H9N2 influenza virus containing 2009 pandemic H1N1 internal-protein genes acquired enhanced pig-to-pig transmission after serial passages in swine","volume":"7","author":"Mancera Gracia","year":"2017","journal-title":"Sci. Rep."},{"key":"10.1016\/j.virol.2023.109927_bib22","doi-asserted-by":"crossref","first-page":"21312","DOI":"10.1073\/pnas.0911915106","article-title":"Adaptive strategies of the influenza virus polymerase for replication in humans","volume":"106","author":"Mehle","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.virol.2023.109927_bib23","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2203919119","article-title":"A Eurasian avian-like H1N1 swine influenza reassortant virus became pathogenic and highly transmissible due to mutations in its PA gene","volume":"119","author":"Meng","year":"2022","journal-title":"P Natl Acad Sci USA"},{"key":"10.1016\/j.virol.2023.109927_bib24","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1128\/CMR.00037-06","article-title":"Avian influenza virus (H5N1): a threat to human health","volume":"20","author":"Peiris","year":"2007","journal-title":"Clin. Microbiol. Rev."},{"key":"10.1016\/j.virol.2023.109927_bib25","doi-asserted-by":"crossref","first-page":"12411","DOI":"10.1128\/JVI.01677-12","article-title":"Evolutionary conservation of the PA-X open reading frame in segment 3 of influenza A virus","volume":"86","author":"Shi","year":"2012","journal-title":"J. Virol."},{"key":"10.1016\/j.virol.2023.109927_bib26","doi-asserted-by":"crossref","first-page":"1122","DOI":"10.1038\/nature08182","article-title":"Origins and evolutionary genomics of the 2009 swine-origin H1N1 influenza A epidemic","volume":"459","author":"Smith","year":"2009","journal-title":"Nature"},{"key":"10.1016\/j.virol.2023.109927_bib27","doi-asserted-by":"crossref","first-page":"741","DOI":"10.3201\/eid1405.071471","article-title":"Transmission of avian influenza virus (H3N2) to dogs","volume":"14","author":"Song","year":"2008","journal-title":"Emerg. Infect. Dis."},{"key":"10.1016\/j.virol.2023.109927_bib28","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.3201\/eid2807.220642","article-title":"Natural reassortment of eurasian avian-like swine H1N1 and avian H9N2 influenza viruses in pigs, China","volume":"28","author":"Sun","year":"2022","journal-title":"Emerg. Infect. Dis."},{"key":"10.1016\/j.virol.2023.109927_bib29","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1016\/j.meegid.2013.10.012","article-title":"Evidence of avian-like H9N2 influenza A virus among dogs in Guangxi, China","volume":"20","author":"Sun","year":"2013","journal-title":"Infect. Genet. Evol."},{"key":"10.1016\/j.virol.2023.109927_bib30","doi-asserted-by":"crossref","first-page":"4164","DOI":"10.1073\/pnas.1019109108","article-title":"High genetic compatibility and increased pathogenicity of reassortants derived from avian H9N2 and pandemic H1N1\/2009 influenza viruses","volume":"108","author":"Sun","year":"2011","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.virol.2023.109927_bib31","doi-asserted-by":"crossref","first-page":"1017","DOI":"10.1080\/22221751.2019.1637284","article-title":"Characterization of swine-origin H1N1 canine influenza viruses","volume":"8","author":"Wang","year":"2019","journal-title":"Emerg. Microb. Infect."},{"key":"10.1016\/j.virol.2023.109927_bib32","doi-asserted-by":"crossref","first-page":"3506","DOI":"10.1128\/JVI.02605-15","article-title":"Infectivity and transmissibility of avian H9N2 influenza viruses in pigs","volume":"90","author":"Wang","year":"2016","journal-title":"J. Virol."},{"key":"10.1016\/j.virol.2023.109927_bib33","doi-asserted-by":"crossref","DOI":"10.1101\/cshperspect.a038489","article-title":"The ecology and evolution of influenza viruses","volume":"10","author":"Wille","year":"2020","journal-title":"Cold Spring Harb Perspect Med"},{"key":"10.1016\/j.virol.2023.109927_bib34","article-title":"Genesis and dissemination of highly pathogenic H5N6 avian influenza viruses","volume":"91","author":"Yang","year":"2017","journal-title":"J. Virol."},{"key":"10.1016\/j.virol.2023.109927_bib35","doi-asserted-by":"crossref","first-page":"57","DOI":"10.3389\/fmicb.2016.00057","article-title":"Phylogenetic analysis and pathogenicity assessment of two strains of avian influenza virus subtype H9N2 isolated from migratory birds: high homology of internal genes with human H10N8 virus","volume":"7","author":"Ye","year":"2016","journal-title":"Front. Microbiol."},{"key":"10.1016\/j.virol.2023.109927_bib36","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1016\/j.vetmic.2010.11.008","article-title":"Genetic diversity of H9N2 influenza viruses from pigs in China: a potential threat to human health?","volume":"149","author":"Yu","year":"2011","journal-title":"Vet. Microbiol."},{"key":"10.1016\/j.virol.2023.109927_bib37","doi-asserted-by":"crossref","first-page":"6553","DOI":"10.1038\/ncomms7553","article-title":"Mammalian adaptation of influenza A(H7N9) virus is limited by a narrow genetic bottleneck","volume":"6","author":"Zaraket","year":"2015","journal-title":"Nat. Commun."},{"key":"10.1016\/j.virol.2023.109927_bib38","first-page":"3266","article-title":"Genetic evolution and pathogenicity analysis of a canine-origin H9N2 subtype influenza virus","volume":"53","author":"Zeng","year":"2022","journal-title":"Journal of Southern Agriculture"},{"key":"10.1016\/j.virol.2023.109927_bib39","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1126\/science.1240532","article-title":"H7N9 influenza viruses are transmissible in ferrets by respiratory droplet","volume":"341","author":"Zhang","year":"2013","journal-title":"Science"},{"key":"10.1016\/j.virol.2023.109927_bib40","doi-asserted-by":"crossref","first-page":"1459","DOI":"10.1126\/science.1229455","article-title":"H5N1 hybrid viruses bearing 2009\/H1N1 virus genes transmit in Guinea pigs by respiratory droplet","volume":"340","author":"Zhang","year":"2013","journal-title":"Science"},{"key":"10.1016\/j.virol.2023.109927_bib41","doi-asserted-by":"crossref","first-page":"2260","DOI":"10.1128\/JVI.03024-13","article-title":"PB2-588I enhances 2009 H1N1 pandemic influenza virus virulence by increasing viral replication and exacerbating PB2 inhibition of beta interferon expression","volume":"88","author":"Zhao","year":"2014","journal-title":"J. Virol."},{"key":"10.1016\/j.virol.2023.109927_bib42","doi-asserted-by":"crossref","DOI":"10.1016\/j.vetmic.2022.109615","article-title":"Synergistic HA and NS mutations enhanced the virulence of a mouse-adapted H1N1 influenza A virus","volume":"276","author":"Zhu","year":"2023","journal-title":"Vet. Microbiol."}],"container-title":["Virology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0042682223002465?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0042682223002465?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,1,1]],"date-time":"2025-01-01T04:52:35Z","timestamp":1735707155000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0042682223002465"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,1]]},"references-count":42,"alternative-id":["S0042682223002465"],"URL":"https:\/\/doi.org\/10.1016\/j.virol.2023.109927","relation":{},"ISSN":["0042-6822"],"issn-type":[{"value":"0042-6822","type":"print"}],"subject":[],"published":{"date-parts":[[2024,1]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Reassortant H9N2 canine influenza viruses containing the pandemic H1N1\/2009 ribonucleoprotein complex circulating in pigs acquired enhanced virulence in mice","name":"articletitle","label":"Article Title"},{"value":"Virology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.virol.2023.109927","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2023 Elsevier Inc.","name":"copyright","label":"Copyright"}],"article-number":"109927"}}