{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,9]],"date-time":"2026-06-09T04:58:34Z","timestamp":1780981114788,"version":"3.54.1"},"reference-count":92,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2026,6,5]],"date-time":"2026-06-05T00:00:00Z","timestamp":1780617600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"The Science Committee of the Ministry of Science and Higher Education of the Republic of Ka-zakhstan.","award":["Grant No. AP23487141"],"award-info":[{"award-number":["Grant No. AP23487141"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>Seasonal human influenza A viruses (H1N1 and H3N2) are frequently transmitted to swine populations. Continued transmission of these viruses among swine results in genetic and phenotypic changes, creating new lineages of swine influenza viruses that differ from their parent strains. This contributes to increased viral genetic diversity and the emergence of zoonotic risks. The aim of this study was to assess the prevalence of influenza viruses, including presumably human-origin A(H3N2) strains, on swine farms in Kazakhstan from 2022 to 2025. Real-time RT-PCR was used to test nasal swab antigens, and hemagglutinin inhibition test, and ELISA were used to determine serum antibody levels. Influenza A\/H1N1 and A\/H3N2 viruses, as well as the A(H7) hemagglutinin genes, were detected in swine. Serological analysis confirmed the presence of antibodies to the A\/H1N1pdm09 and A\/H3N2 viruses. The influenza A(H3N2) isolates were found to be genetically similar to human viruses of the J.2 or 3C.2a1b.2a.2a.3a.1 clade, possibly indicating transmission from humans to animals. The ongoing exchange of influenza viruses between humans and swine poses a threat of future pandemics. These studies highlight the need for continuous monitoring of swine influenza in Kazakhstan to understand virus evolution and protect public health.<\/jats:p>","DOI":"10.3390\/ani16111752","type":"journal-article","created":{"date-parts":[[2026,6,5]],"date-time":"2026-06-05T17:18:16Z","timestamp":1780679896000},"page":"1752","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Human-Origin Influenza A(H3N2) Viruses Revealed in Swine Farms During the Period 2022\u20132025 in Kazakhstan"],"prefix":"10.3390","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4094-137X","authenticated-orcid":false,"given":"Nailya","family":"Klivleyeva","sequence":"first","affiliation":[{"name":"The Research and Production Center for Microbiology and Virology, Almaty 050010, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1139-9868","authenticated-orcid":false,"given":"Tatyana","family":"Glebova","sequence":"additional","affiliation":[{"name":"The Research and Production Center for Microbiology and Virology, Almaty 050010, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6526-956X","authenticated-orcid":false,"given":"Nurbol","family":"Saktaganov","sequence":"additional","affiliation":[{"name":"The Research and Production Center for Microbiology and Virology, Almaty 050010, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8959-126X","authenticated-orcid":false,"given":"Assem","family":"Baimukhametova","sequence":"additional","affiliation":[{"name":"The Research and Production Center for Microbiology and Virology, Almaty 050010, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3247-2761","authenticated-orcid":false,"given":"Mereke","family":"Kalkozhayeva","sequence":"additional","affiliation":[{"name":"The Research and Production Center for Microbiology and Virology, Almaty 050010, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9022-331X","authenticated-orcid":false,"given":"Nuray","family":"Ongarbayeva","sequence":"additional","affiliation":[{"name":"The Research and Production Center for Microbiology and Virology, Almaty 050010, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Indira","family":"Ibragimova","sequence":"additional","affiliation":[{"name":"Postgraduate Cardiology Course, Kazakhstan-Russian Medical University, Almaty 050000, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Primkul","family":"Ibragimov","sequence":"additional","affiliation":[{"name":"Non-Profit Joint-Stock Company, Kazakh National Agrarian Research University, Almaty 050010, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4397-7132","authenticated-orcid":false,"given":"Richard","family":"Webby","sequence":"additional","affiliation":[{"name":"Department of Infectious Diseases, St. Jude Children\u2019s Research Hospital, Memphis, TN 38105-3678, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,6,5]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Klivleyeva, N., Saktaganov, N., Glebova, T., Lukmanova, G., Ongarbayeva, N., and Webby, R. (2024). Influenza A Viruses in the Swine Population: Ecology and Geographical Distribution. Viruses, 16.","DOI":"10.20944\/preprints202410.0234.v1"},{"key":"ref_2","first-page":"158","article-title":"The Pig as a Mixing Vessel for Influenza Viruses: Human and Veterinary Implications","volume":"3","author":"Ma","year":"2008","journal-title":"J. Mol. Genet. Med."},{"key":"ref_3","unstructured":"Knipe, D.M., and Howley, P.M. (2007). Orthomyxoviruses. Fields Virology, Lippincot Williams & Wilkins. [5th ed.]."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"73804","DOI":"10.47391\/JPMA.6348","article-title":"Serological Studies of Influenza Infection among Population in Southern Region of Kazakhstan during the 2018\u20132021 Epidemic Season","volume":"73","author":"Shamenova","year":"2023","journal-title":"J. Pak. Med. Assoc."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1896","DOI":"10.5455\/OVJ.2024.v14.i8.17","article-title":"Circulation of Influenza Viruses in the Dog Population in Kazakhstan (2023\u20132024)","volume":"14","author":"Glebova","year":"2024","journal-title":"Open Vet. J."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1128\/mr.56.1.152-179.1992","article-title":"Evolution and Ecology of Influenza A Viruses","volume":"56","author":"Webster","year":"1992","journal-title":"Microbiol. Rev."},{"key":"ref_7","first-page":"133","article-title":"History and epidemiology of Swine influenza in Europe","volume":"370","author":"Brown","year":"2013","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"2195","DOI":"10.1099\/vir.0.044974-0","article-title":"Global Transmission of Influenza Viruses from Humans to Swine","volume":"93","author":"Nelson","year":"2012","journal-title":"J. Gen. Virol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"2584","DOI":"10.1099\/vir.0.045930-0","article-title":"Genomic Reassortment of Influenza A Virus in North American Swine, 1998\u20132011","volume":"93","author":"Nelson","year":"2012","journal-title":"J. Gen. Virol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1038\/nature10004","article-title":"Long-Term Evolution and Transmission Dynamics of Swine Influenza A Virus","volume":"473","author":"Vijaykrishna","year":"2011","journal-title":"Nature"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1236","DOI":"10.1099\/vir.0.051839-0","article-title":"Genotype Patterns of Contemporary Reassorted H3N2 Virus in US Swine","volume":"94","author":"Kitikoon","year":"2013","journal-title":"J. Gen. Virol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"2498","DOI":"10.1093\/aje\/kwy150","article-title":"Origins of the 1918 Pandemic: Revisiting the Swine \u201cMixing Vessel\u201d Hypothesis","volume":"187","author":"Nelson","year":"2018","journal-title":"Am. J. Epidemiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1084\/jem.54.3.373","article-title":"Swine Influenza: III. Filtration Experiments and Etiology","volume":"54","author":"Shope","year":"1931","journal-title":"J. Exp. Med."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"11276","DOI":"10.1038\/s41598-021-90512-z","article-title":"Genetic and Antigenic Evolution of H1 Swine Influenza A Viruses Isolated in Belgium and the Netherlands from 2014 through 2019","volume":"11","author":"Chepkwony","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_15","unstructured":"(2026, April 20). History of Swine Influenza. Russian Veterinary Journal. Available online: https:\/\/cyberleninka.ru\/article\/n\/istoriya-grippa-sviney."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1665","DOI":"10.3201\/eid1810.120398","article-title":"Influenza A(H1N1) Pdm09 Virus in Pigs, R\u00e9union Island","volume":"18","author":"Cardinale","year":"2012","journal-title":"Emerg. Infect. Dis."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1519","DOI":"10.3201\/eid1809.120431","article-title":"Influenza A(H1N1)Pdm09 Virus among Healthy Show Pigs, United States","volume":"18","author":"Gray","year":"2012","journal-title":"Emerg. Infect. Dis."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"10864","DOI":"10.1128\/JVI.01327-14","article-title":"Expansion of Genotypic Diversity and Establishment of 2009 H1N1 Pandemic-Origin Internal Genes in Pigs in China","volume":"88","author":"Liang","year":"2014","journal-title":"J. Virol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"699","DOI":"10.4014\/jmb.1106.05062","article-title":"Identification of reassortant pandemic H1N1 influenza virus in Korean pigs","volume":"22","author":"Han","year":"2012","journal-title":"J. Microbiol. Biotechnol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Glebova, T., Klivleyeva, N., Baimukhametova, A., Lukmanova, G., Saktaganov, N., Ongarbayeva, N., Baimakhanova, B., Kassymova, G., Sagatova, M., and Rachimbayeva, A. (2025). Acute Respiratory and Influenza Viruses Circulating in Kazakhstan During 2018\u20132024. Pathogens, 14.","DOI":"10.3390\/pathogens14050493"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"8243","DOI":"10.1128\/JVI.74.18.8243-8251.2000","article-title":"Evolution of Swine H3N2 Influenza Viruses in the United States","volume":"74","author":"Webby","year":"2000","journal-title":"J. Virol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"8872","DOI":"10.1128\/JVI.00259-12","article-title":"Evolution of Novel Reassortant A\/H3N2 Influenza Viruses in North American Swine and Humans, 2009\u20132011","volume":"86","author":"Nelson","year":"2012","journal-title":"J. Virol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1007\/s00705-018-4091-4","article-title":"Genetic and Antigenic Dynamics of Influenza A Viruses of Swine on Pig Farms in Thailand","volume":"164","author":"Mine","year":"2019","journal-title":"Arch. Virol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"857","DOI":"10.1038\/228857a0","article-title":"Hong Kong A-2 Influenza Virus Infection among Swine during a Human Epidemic in Taiwan","volume":"228","author":"Kundin","year":"1970","journal-title":"Nature"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1454","DOI":"10.1126\/science.867041","article-title":"Persistence of Hong Kong Influenza Virus Variants in Pigs","volume":"196","author":"Shortridge","year":"1977","journal-title":"Science"},{"key":"ref_26","first-page":"187","article-title":"Haemagglutination-Inhibiting Antibodies against Influenza Type A Viruses in Swine Sera in India","volume":"68","author":"Singh","year":"1978","journal-title":"Indian J. Med. Res."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1099\/0022-1317-56-2-441","article-title":"Antigenic and Genetic Analysis of A\/Hong Kong (H3N2) Influenza Viruses Isolated from Swine and Man","volume":"56","author":"Nerome","year":"1981","journal-title":"J. Gen. Virol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1620\/tjem.136.353","article-title":"The Follow-Up Study of Swine and Hong Kong Influenza Virus Infection among Japanese Hogs","volume":"136","author":"Arikawa","year":"1982","journal-title":"Tohoku J. Exp. Med."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1016\/0042-6822(88)90405-9","article-title":"Origin of the Hemagglutinin Gene of H3N2 Influenza Viruses from Pigs in China","volume":"162","author":"Kida","year":"1988","journal-title":"Virology"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1099\/0022-1317-76-3-613","article-title":"Genetic Analysis of Porcine H3N2 Viruses Originating in Southern China","volume":"76","author":"Nerome","year":"1995","journal-title":"J. Gen. Virol."},{"key":"ref_31","first-page":"53","article-title":"Sero-Epidemiology and Genetic Characterization of Swine Influenza Virus","volume":"38","author":"Lyoo","year":"1998","journal-title":"Korean J. Vet. Res."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"9679","DOI":"10.1128\/JVI.75.20.9679-9686.2001","article-title":"Cocirculation of Avian H9N2 and Contemporary \u201cHuman\u201d H3N2 Influenza A Viruses in Pigs in Southeastern China: Potential for Genetic Reassortment?","volume":"75","author":"Peiris","year":"2001","journal-title":"J. Virol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1177\/104063870301500107","article-title":"Isolation of H3N2 Swine Influenza Virus in South Korea","volume":"15","author":"Song","year":"2003","journal-title":"J. Vet. Diagn. Investig."},{"key":"ref_34","unstructured":"Richt, J., and Webby, R. (2011). Swine Influenza Viruses: An Asian Perspective. Swine Influenza, Springer. Current Topics in Microbiology and Immunology."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1128\/JCM.01257-07","article-title":"Genetic Evolution of Swine Influenza A (H3N2) Viruses in China from 1970 to 2006","volume":"46","author":"Yu","year":"2008","journal-title":"J. Clin. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Tochetto, C., Junqueira, D.M., Anderson, T.K., Gava, D., Haach, V., Cant\u00e3o, M.E., Baker, A.L.V., and Schaefer, R. (2023). Introductions of Human-Origin Seasonal H3N2, H1N2 and Pre-2009 H1N1 Influenza Viruses to Swine in Brazil. Viruses, 15.","DOI":"10.3390\/v15020576"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.virusres.2008.08.005","article-title":"Seroprevalence and Genetic Evolutions of Swine Influenza Viruses under Vaccination Pressure in Korean Swine Herds","volume":"138","author":"Pascua","year":"2008","journal-title":"Virus Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"481","DOI":"10.3201\/eid1903.120945","article-title":"Swine Influenza in Sri Lanka","volume":"19","author":"Perera","year":"2013","journal-title":"Emerg. Infect. Dis."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1077","DOI":"10.4269\/ajtmh.2012.11-0195","article-title":"Serologic Study of Pig-Associated Viral Zoonoses in Laos","volume":"86","author":"Conlan","year":"2012","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1557","DOI":"10.1111\/tbed.12547","article-title":"Influenza A Virus in Backyard Pigs and Poultry in Rural Cambodia","volume":"64","author":"Osbjer","year":"2017","journal-title":"Transbound. Emerg. Dis."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1292\/jvms.57.1023","article-title":"Antigenic and Genetic Analyses of the Hemagglutinin of Influenza Viruses Isolated from Pigs in 1993","volume":"57","author":"Katsuda","year":"1995","journal-title":"J. Vet. Med. Sci."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/S0168-1702(00)00154-4","article-title":"Genetic Characterization of H3N2 Influenza Viruses Isolated from Pigs in North America, 1977\u20131999: Evidence for Wholly Human and Reassortant Virus Genotypes","volume":"68","author":"Karasin","year":"2000","journal-title":"Virus Res."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1829","DOI":"10.1098\/rstb.2001.1020","article-title":"Integrating Historical, Clinical and Molecular Genetic Data in Order to Explain the Origin and Virulence of the 1918 Spanish Influenza Virus","volume":"356","author":"Taubenberger","year":"2001","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"ref_44","first-page":"207","article-title":"Diversity of Influenza Viruses in Swine and the Emergence of a Novel Human Pandemic Influenza A (H1N1). Influenza Other Respir","volume":"3","author":"Webster","year":"2009","journal-title":"Viruses"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1111\/zph.12210","article-title":"Antigenic Detection of Human Strain of Influenza Virus A (H3N2) in Swine Populations at Three Locations in Nigeria and Ghana during the Dry Early Months of 2014","volume":"63","author":"Adeola","year":"2016","journal-title":"Zoonoses Public Health"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Rajao, D.S., Vincent, A.L., and Perez, D.R. (2019). Adaptation of Human Influenza Viruses to Swine. Front. Vet. Sci., 5.","DOI":"10.3389\/fvets.2018.00347"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"e00822-19","DOI":"10.1128\/mSphere.00822-19","article-title":"Influenza A Virus Field Surveillance at a Swine-Human Interface","volume":"5","author":"Keller","year":"2020","journal-title":"mSphere"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1111\/zph.13104","article-title":"Evolution of Influenza A Viruses in Exhibition Swine and Transmission to Humans, 2013\u20132015","volume":"71","author":"Szablewski","year":"2024","journal-title":"Zoonoses Public Health"},{"key":"ref_49","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_50","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1016\/j.virusres.2004.02.015","article-title":"Multiple Lineages of Antigenically and Genetically Diverse Influenza A Virus Co-Circulate in the United States Swine Population","volume":"103","author":"Webby","year":"2004","journal-title":"Virus Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1007\/978-1-0716-0346-8_18","article-title":"A Brief Introduction to Influenza A Virus in Swine","volume":"2123","author":"Vincent","year":"2020","journal-title":"Methods Mol. Biol."},{"key":"ref_52","first-page":"201","article-title":"Serologic and Genetic Characterization of North American H3N2 Swine Influenza A Viruses","volume":"71","author":"Gramer","year":"2007","journal-title":"Can. J. Vet. Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1132","DOI":"10.3201\/eid1207.060268","article-title":"Triple Reassortant H3N2 Influenza A Viruses, Canada, 2005","volume":"12","author":"Olsen","year":"2006","journal-title":"Emerg. Infect. Dis."},{"key":"ref_54","first-page":"262","article-title":"Regional Patterns of Genetic Diversity in Swine Influenza A Viruses in the United States from 2010 to 2016. Influenza Other Respir","volume":"13","author":"Walia","year":"2019","journal-title":"Viruses"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"8851","DOI":"10.1128\/JVI.73.10.8851-8856.1999","article-title":"Genetic Reassortment of Avian, Swine, and Human Influenza A Viruses in American Pigs","volume":"73","author":"Zhou","year":"1999","journal-title":"J. Virol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1111\/zph.12049","article-title":"Review of influenza A virus in swine worldwide: A call for increased surveillance and research","volume":"61","author":"Vincent","year":"2014","journal-title":"Zoonoses Public Health"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1178","DOI":"10.1086\/651507","article-title":"Human Infection with a Triple Reassortant Swine Influenza A(H1N1) Virus Containing the Hemagglutinin and Neuraminidase Genes of Seasonal Influenza Virus","volume":"201","author":"Bastien","year":"2010","journal-title":"J. Infect. Dis."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1529","DOI":"10.1126\/science.1189132","article-title":"Reassortment of Pandemic H1N1\/2009 Influenza A Virus in Swine","volume":"328","author":"Vijaykrishna","year":"2010","journal-title":"Science"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1073","DOI":"10.1128\/JCM.40.3.1073-1079.2002","article-title":"Genetic Characterization of H1N2 Influenza A Viruses Isolated from Pigs throughout the United States","volume":"40","author":"Karasin","year":"2002","journal-title":"J. Clin. Microbiol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1016\/S0168-1702(02)00053-9","article-title":"Phylogenetic Analysis of H1N2 Isolates of Influenza A Virus from Pigs in the United States","volume":"87","author":"Choi","year":"2002","journal-title":"Virus Res."},{"key":"ref_61","first-page":"188","article-title":"Genetic Composition of Contemporary Swine Influenza Viruses in the West Central Region of the United States of America. Influenza Other Respir","volume":"5","author":"Evseenko","year":"2011","journal-title":"Viruses"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1099\/vir.0.027557-0","article-title":"Genetic and Antigenic Characterization of H1 Influenza Viruses from United States Swine from 2008","volume":"92","author":"Lorusso","year":"2011","journal-title":"J. Gen. Virol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1111\/j.1865-1682.2011.01219.x","article-title":"Molecular and Antigenic Characterization of Triple-Reassortant H3N2 Swine Influenza Viruses Isolated from Pigs, Turkey and Quail in Canada","volume":"58","author":"Nfon","year":"2011","journal-title":"Transbound. Emerg. Dis."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"8667","DOI":"10.1128\/JVI.00801-11","article-title":"Characterization of H1N1 Swine Influenza Viruses Circulating in Canadian Pigs in 2009","volume":"85","author":"Nfon","year":"2011","journal-title":"J. Virol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1016\/j.virol.2020.11.006","article-title":"Characterization of Contemporary 2010.1 H3N2 Swine Influenza A Viruses Circulating in United States Pigs","volume":"553","author":"Powell","year":"2021","journal-title":"Virology"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"e16777","DOI":"10.7554\/eLife.16777","article-title":"Origins of the 2009 H1N1 Influenza Pandemic in Swine in Mexico","volume":"5","author":"Mena","year":"2016","journal-title":"Elife"},{"key":"ref_67","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_68","doi-asserted-by":"crossref","unstructured":"Klivleyeva, N., Glebova, T., Saktaganov, N., and Webby, R. (2025). Cases of Interspecies Transmission of Influenza A Virus from Swine to Humans. Vet. Sci., 12.","DOI":"10.3390\/vetsci12090873"},{"key":"ref_69","unstructured":"WOAH (2023). Terrestrial Animal Health Code. Section 7: Animal Welfare, World Organisation for Animal Health."},{"key":"ref_70","unstructured":"WOAH (2024). Collection, submission and storage of diagnostic specimens. Manual of Diagnostic Tests and Vaccines for Terrestrial Animals, World Organisation for Animal Health. [2024 ed.]. Available online: http:\/\/woah.org."},{"key":"ref_71","unstructured":"OFFLU (2021). Standard Operating Procedure: Sample Collection and Transport for Swine Influenza Surveillance, FAO\/WOAH Network of Expertise on Animal Influenza. Available online: http:\/\/surl.lt."},{"key":"ref_72","unstructured":"World Health Organization (2024). Manual for the Laboratory Diagnosis and Virological Surveillance of Influenza, WHO. Available online: http:\/\/who.int."},{"key":"ref_73","unstructured":"Centers for Disease Control and Prevention (2025, May 13). CDC Influenza Virus Real-Time RT-PCR Diagnostic Panel\u2014Instructions for Use, Available online: http:\/\/fda.gov."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1007\/978-1-61779-621-0_3","article-title":"Influenza Virus Titration, Antigenic Characterization, and Serological Methods for Antibody Detection","volume":"865","author":"Klimov","year":"2012","journal-title":"Methods Mol. Biol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1007\/978-1-61779-621-0_11","article-title":"Influenza A Virus Molecular Virology Techniques","volume":"865","author":"Zhou","year":"2012","journal-title":"Methods Mol. Biol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1821","DOI":"10.3201\/eid2707.210462","article-title":"Multiplex Real-Time Reverse Transcription PCR for Influenza A Virus, Influenza B Virus, and Severe Acute Respiratory Syndrome Coronavirus 2","volume":"27","author":"Shu","year":"2021","journal-title":"Emerg. Infect. Dis."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1002\/sim.4780121305","article-title":"A Comparison of the Power of Two Tests for Qualitative Interactions","volume":"12","author":"Piantadosi","year":"1993","journal-title":"Stat. Med."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"msae263","DOI":"10.1093\/molbev\/msae263","article-title":"MEGA12: Molecular Evolutionary Genetic Analysis Version 12 for Adaptive and Green Computing","volume":"41","author":"Kumar","year":"2024","journal-title":"Mol. Biol. Evol."},{"key":"ref_79","first-page":"406","article-title":"The Neighbor-Joining Method: A New Method for Reconstructing Phylogenetic Trees","volume":"4","author":"Saitou","year":"1987","journal-title":"Mol. Biol. Evol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1007\/BF02101694","article-title":"Dating of the Human-Ape Splitting by a Molecular Clock of Mitochondrial DNA","volume":"22","author":"Hasegawa","year":"1985","journal-title":"J. Mol. Evol."},{"key":"ref_81","first-page":"678","article-title":"Estimation of the Number of Nucleotide Substitutions When There Are Strong Transition-Transversion and G + C-Content Biases","volume":"9","author":"Tamura","year":"1992","journal-title":"Mol. Biol. Evol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"3773","DOI":"10.21105\/joss.03773","article-title":"Nextclade: Clade Assignment, Mutation Calling and Quality Control for Viral Genomes","volume":"6","author":"Aksamentov","year":"2021","journal-title":"J. Open Source Softw."},{"key":"ref_83","unstructured":"(2026, April 20). Northern Kazakhstan Has the Largest Pig Population in the Country. Available online: https:\/\/pkzsk.info\/."},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Ag\u00fcero, B.M., Ariyama, N., Berrios, F., Enciso, N., Quezada, B., Medina, R.A., and Neira, V. (2025). Novel Introductions of Human-Origin H3N2 Influenza Viruses in Swine, Chile. Front. Vet. Sci., 11.","DOI":"10.3389\/fvets.2024.1505497"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"e0080921","DOI":"10.1128\/msphere.00809-21","article-title":"Characterization of a 2016\u20132017 Human Seasonal H3 Influenza A Virus Spillover Now Endemic to U.S. Swine","volume":"7","author":"Sharma","year":"2022","journal-title":"mSphere"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"e20230403","DOI":"10.1590\/0103-8478cr20230403","article-title":"Influenza A Virus Surveillance in Domestic Pigs in Kazakhstan 2018\u20132021","volume":"54","author":"Lukmanova","year":"2024","journal-title":"Ci\u00eanc. Rural."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"355","DOI":"10.15389\/agrobiology.2020.2.355eng","article-title":"Study on Antigenic Relationships and Biological Properties of Swine Influenza a\/H1n1 Virus Strains Isolated in Northern Kazakhstan in 2018","volume":"55","author":"Saktaganov","year":"2020","journal-title":"Sel\u2019skokhozyaistvennaya Biol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"e0078621","DOI":"10.1128\/MRA.00786-21","article-title":"Coding-Complete Genome Sequence of Swine Influenza Virus Isolate A\/Swine\/Karaganda\/04\/2020 (H1N1) from Kazakhstan","volume":"10","author":"Klivleyeva","year":"2021","journal-title":"Microbiol. Resour. Announc."},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Glebova, T., Klivleyeva, N., Ongarbayeva, N., Baimukhametova, A., Saktaganov, N., Kalkozhayeva, M., Karamendin, K., Ibragimova, I., Sagatova, M., and Mutaliyeva, A. (2026). Surveillance of Seasonal Influenza Viruses in Kazakhstan (2020\u20132025). Viruses, 18.","DOI":"10.3390\/v18040441"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"1169","DOI":"10.15585\/mmwr.mm6742a1","article-title":"Influenza A(H3N2) Variant Virus Outbreak at Three Fairs\u2014Maryland, 2017","volume":"67","author":"Duwell","year":"2018","journal-title":"MMWR Morb. Mortal. Wkly. Rep."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1551","DOI":"10.3201\/eid2309.170847","article-title":"Influenza A(H3N2) Virus in Swine at Agricultural Fairs and Transmission to Humans, Michigan and Ohio, USA, 2016","volume":"23","author":"Bowman","year":"2017","journal-title":"Emerg. Infect. Dis."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"001623","DOI":"10.1099\/jmm.0.001623","article-title":"JMM Profile: Swine Influenza A Virus: A Neglected Virus with Pandemic Potential","volume":"72","author":"Mollett","year":"2023","journal-title":"J. Med. Microbiol."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/16\/11\/1752\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,9]],"date-time":"2026-06-09T04:18:58Z","timestamp":1780978738000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/16\/11\/1752"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,5]]},"references-count":92,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2026,6]]}},"alternative-id":["ani16111752"],"URL":"https:\/\/doi.org\/10.3390\/ani16111752","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,5]]}}}