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Adv."],"published-print":{"date-parts":[[2026,5,8]]},"abstract":"<jats:p>H5Nx clade 2.3.4.4b viruses are evolving rapidly, expanding host ranges and threatening animal and public health. In the US, genotype B3.13 dominates dairy outbreaks, while D1.1 is linked to fewer cases. In the UK, an asymptomatic ewe infected with genotype DI.2 raised concerns about ruminant susceptibility. We inoculated lactating and nonlactating sheep with D1.1 (H5N1) and A6 (H5N5) viruses. Intramammary inoculation in lactating sheep caused clinical mastitis, high viral loads in milk, and transmission to suckling lambs, which further spread infection to the uninoculated mammary glands. Both ewes and their lambs seroconverted. Aerosol exposure of nonlactating sheep led to transient respiratory infection, with low-level viral replication, and seroconversion. In vitro, both viruses replicated in sheep mammary epithelial cells. These findings establish sheep as a viable ruminant model for H5N1 and H5N5 infection and highlight previously unidentified transmission dynamics, including milk-mediated and lamb-to-ewe spread, relevant for surveillance and biosecurity in ruminant populations.<\/jats:p>","DOI":"10.1126\/sciadv.aed1287","type":"journal-article","created":{"date-parts":[[2026,5,12]],"date-time":"2026-05-12T09:53:19Z","timestamp":1778579599000},"update-policy":"https:\/\/doi.org\/10.34133\/aaas_crossmark","source":"Crossref","is-referenced-by-count":1,"title":["Mammary and respiratory infection of sheep with H5Nx clade 2.3.4.4b viruses with milk-mediated transmission to lambs"],"prefix":"10.1126","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0009-0008-6507-6635","authenticated-orcid":true,"given":"Tamiru N.","family":"Alkie","sequence":"first","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carissa","family":"Embury-Hyatt","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0664-0864","authenticated-orcid":true,"given":"Anthony V.","family":"Signore","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-9483-3606","authenticated-orcid":true,"given":"Estella","family":"Moffat","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Darwin","family":"Ramos","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sugandha","family":"Raj","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0216-6496","authenticated-orcid":true,"given":"Tamiko","family":"Hisanaga","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6742-3065","authenticated-orcid":true,"given":"Ifeoluwa","family":"Ayilara","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel S.","family":"Sullivan","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patrick","family":"Lloyd","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lemarie","family":"Pama","sequence":"additional","affiliation":[{"name":"Canadian Food Inspection Agency, National Centre for Foreign Animal Disease, Winnipeg, Manitoba, Canada."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8893-7804","authenticated-orcid":true,"given":"Cassidy N. 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Alexakis, H. Buczkowski, M. Ducatez, A. Fusaro, J. L. Gonzales, T. Kuiken, K. St\u00e5hl, C. Staubach, O. Svartstr\u00f6m, C. Terregino, K. Willgert, M. Melo, L. Kohnle, Avian influenza overview December 2024\u2013March 2025. EFSA J. 23, e9352 (2025).","journal-title":"EFSA J."},{"key":"e_1_3_2_21_2","unstructured":"Pan American Health Organization\/World Organization Epidemiological update: Avian influenza A(H5N1) in the Americas region 4 March 2025 (PAHO\/WHO 2025); www.paho.org\/sites\/default\/files\/2025-03\/2025-mar-4-phe-epidupdate-avianinfluenza-eng-final.pdf."},{"key":"e_1_3_2_22_2","unstructured":"L. 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Smith Highly pathogenic avian influenza H5 virus exposure in goats and sheep. bioRxiv 610397 [Preprint] (2024); https:\/\/doi.org\/10.1101\/2024.08.31.610397.","DOI":"10.1101\/2024.08.31.610397"},{"key":"e_1_3_2_31_2","doi-asserted-by":"crossref","unstructured":"J. H. Fosse G. R\u00f8mo F. Bonfante I. K. Myhrvold K. S. Soetart K. Udjus R. 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Hulscher P. A. McCullough \u201cH5N1 genotype D1.1: Understanding its evolution risks and public health solutions \u201d Focal Points Courageous Discourse 2 March 2025; www.thefocalpoints.com\/p\/h5n1-genotype-d11-understanding-its."},{"key":"e_1_3_2_48_2","doi-asserted-by":"crossref","unstructured":"A. J. Campbell M. Shephard A. P. Paulos M. Pauly M. Vu C. Stenkamp-Strahm K. Bushfield B. Hunter-Binns O. Sablon E. E. Bendall W. J. Fitzimmons K. Brizuela G. Quirk N. Kumar B. McCluskey N. Shetty L. C. Marr J. J. Guthmiller K. Abernathy A. S. Lauring B. T. Melody M. Wolfe J. Lombard S. S. Lakdawala Surveillance on California dairy farms reveals multiple sources of H5N1 transmission. bioRxiv 666798 [Preprint] (2025); 10.1101\/2025.07.31.666798.","DOI":"10.1101\/2025.07.31.666798"},{"key":"e_1_3_2_49_2","doi-asserted-by":"publisher","DOI":"10.3201\/eid3008.240775"},{"key":"e_1_3_2_50_2","doi-asserted-by":"crossref","unstructured":"C. Lee N. N. Tarbuck H. J. Cochran B. M. Foreman P. 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Bell Exploring influenza A virus receptor distribution in the lactating mammary gland of domesticated livestock and in human breast tissue. bioRxiv 649193 [Preprint] (2025); https:\/\/doi.org\/10.1101\/2025.04.16.649193.","DOI":"10.1101\/2025.04.16.649193"},{"key":"e_1_3_2_53_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41564-025-01998-6"},{"key":"e_1_3_2_54_2","first-page":"97","article-title":"Preliminary experiments relating to the propagation of viruses in the bovine mammary gland","volume":"17","author":"Mitchell C. A.","year":"1953","unstructured":"C. A. Mitchell, R. V. Walker, G. L. Bannister, Preliminary experiments relating to the propagation of viruses in the bovine mammary gland. Can. J. Comp. Med. Vet. Sci. 17, 97\u2013104 (1953).","journal-title":"Can. J. Comp. Med. Vet. Sci."},{"key":"e_1_3_2_55_2","doi-asserted-by":"publisher","DOI":"10.3201\/eid3107.250134"},{"key":"e_1_3_2_56_2","unstructured":"R. Brooks \u201cSome California veterinarians say virus-hit dairies see more abortions in first-calf heifers and dry cows \u201d Bovine Veterinarian [updated 14 January] (2025); www.bovinevetonline.com\/news\/industry\/california-veterinarians-say-virus-hit-dairies-see-more-abortions-first-calf-heifer."},{"key":"e_1_3_2_57_2","doi-asserted-by":"crossref","unstructured":"C. Fiorito A. Colom A. Fern\u00e1ndez P. Alonso-Almorox M. Andrada D. Lombardo E. Sierra Pathology of Influenza A (H5N1) infection in pinnipeds reveals novel tissue tropism and vertical transmission. bioRxiv 636856 (2025); https:\/\/doi.org\/10.1101\/2025.02.07.636856.","DOI":"10.1101\/2025.02.07.636856"},{"key":"e_1_3_2_58_2","doi-asserted-by":"crossref","unstructured":"B. A. Seibert M. A. Parish S. N. Talukdar T. Zhang S. Chaulagain J. A. Liu J. Sachithanandham C. Schneider L. Coughlan A. Bowman R. J. Webby J. R. Harkema A. Pekosz S. L. Klein Trimester-dependent vertical transmission of H5N1 influenza virus through placental and mammary routes impairs offspring development. bioRxiv 663583 [Preprint] (2025); 10.1101\/2025.07.07.663583.","DOI":"10.1101\/2025.07.07.663583"},{"key":"e_1_3_2_59_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(07)61515-3"},{"key":"e_1_3_2_60_2","doi-asserted-by":"publisher","DOI":"10.4269\/ajtmh.1977.26.190"},{"key":"e_1_3_2_61_2","doi-asserted-by":"publisher","DOI":"10.3201\/eid1510.090743"},{"key":"e_1_3_2_62_2","doi-asserted-by":"crossref","unstructured":"K. S. Davila A. Baker P. Boggiatto M. Palmer E. Putz S. Olsen G. C. Zanella A. Campos A. Buckley B. Arruda Susceptibility of calves fed unpasteurized milk from cows experimentally infected with highly pathogenic avian influenza H5N1. 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Langel Intramammary infection of bovine H5N1 influenza virus in ferrets leads to transmission and mortality in suckling neonates. bioRxiv 623885 (2024); https:\/\/doi.org\/10.1101\/2024.11.15.623885.","DOI":"10.1101\/2024.11.15.623885"},{"key":"e_1_3_2_67_2","doi-asserted-by":"publisher","DOI":"10.1080\/22221751.2025.2475836"},{"key":"e_1_3_2_68_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-53766-5"},{"key":"e_1_3_2_69_2","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.02944-24"},{"key":"e_1_3_2_70_2","first-page":"426","article-title":"Persistence of neutralizing antibody in milk and blood of cows and goats following the instillation of virus into the mammary gland","volume":"18","author":"Mitchell C. A.","year":"1954","unstructured":"C. A. Mitchell, R. V. Walker, G. L. Bannister, Persistence of neutralizing antibody in milk and blood of cows and goats following the instillation of virus into the mammary gland. Can. J. Comp. Med. Vet. 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