Pathogenicity and transmission of a swine influenza A(H6N6) virus
Identifieur interne : 000209 ( Pmc/Checkpoint ); précédent : 000208; suivant : 000210Pathogenicity and transmission of a swine influenza A(H6N6) virus
Auteurs : Hailiang Sun [États-Unis] ; Bryan S. Kaplan [États-Unis] ; Minhui Guan [États-Unis] ; Guihong Zhang [République populaire de Chine] ; Jianqiang Ye [États-Unis] ; Li-Ping Long [États-Unis] ; Sherry Blackmon [États-Unis] ; Chun-Kai Yang [États-Unis] ; Meng-Jung Chiang [États-Unis] ; Hang Xie [États-Unis] ; Nan Zhao [États-Unis] ; Jim Cooley [États-Unis] ; David F. Smith [États-Unis] ; Ming Liao [République populaire de Chine] ; Carol Cardona [États-Unis] ; Lei Li [États-Unis] ; George Peng Wang [États-Unis] ; Richard Webby [États-Unis] ; Xiu-Feng Wan [États-Unis]Source :
- Emerging Microbes & Infections [ 2222-1751 ] ; 2017.
Abstract
Subtype H6 influenza A viruses (IAVs) are commonly detected in wild birds and domestic poultry and can infect humans. In 2010, a H6N6 virus emerged in southern China, and since then, it has caused sporadic infections among swine. We show that this virus binds to α2,6-linked and α2,3-linked sialic acids. Mutations at residues 222 (alanine to valine) and 228 (glycine to serine) of the virus hemagglutinin (HA) affected its receptor-binding properties. Experiments showed that the virus has limited transmissibility between ferrets through direct contact or through inhalation of infectious aerosolized droplets. The internal genes of the influenza A(H1N1)pdm09 virus, which is prevalent in swine worldwide, increases the replication efficiency of H6N6 IAV in the lower respiratory tract of ferrets but not its transmissibility between ferrets. These findings suggest H6N6 swine IAV (SIV) currently poses a moderate risk to public health, but its evolution and spread should be closely monitored.
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DOI: 10.1038/emi.2017.3
PubMed: 28400591
PubMed Central: 5457681
Affiliations:
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Kaplan</name><affiliation wicri:level="1"><nlm:aff id="aff2"><institution>Department of Infectious Diseases, St Jude Children's Research Hospital</institution>, Memphis, TN 38105,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Guan, Minhui" sort="Guan, Minhui" uniqKey="Guan M" first="Minhui" last="Guan">Minhui Guan</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Zhang, Guihong" sort="Zhang, Guihong" uniqKey="Zhang G" first="Guihong" last="Zhang">Guihong Zhang</name><affiliation wicri:level="1"><nlm:aff 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of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Chiang, Meng Jung" sort="Chiang, Meng Jung" uniqKey="Chiang M" first="Meng-Jung" last="Chiang">Meng-Jung Chiang</name><affiliation wicri:level="1"><nlm:aff id="aff4"><institution>Laboratory of Respiratory Viral Diseases, Division of Viral Products, Office of Vaccines Research and Review, Center for Biologics Evaluation and Research, United States Food and Drug Administration</institution>, Silver Spring, MD 20993,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Xie, Hang" sort="Xie, Hang" uniqKey="Xie H" first="Hang" last="Xie">Hang Xie</name><affiliation wicri:level="1"><nlm:aff id="aff4"><institution>Laboratory of Respiratory Viral Diseases, Division of Viral Products, Office of Vaccines Research and Review, Center for Biologics Evaluation and Research, United States Food and Drug Administration</institution>, Silver Spring, MD 20993,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Zhao, Nan" sort="Zhao, Nan" uniqKey="Zhao N" first="Nan" last="Zhao">Nan Zhao</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Cooley, Jim" sort="Cooley, Jim" uniqKey="Cooley J" first="Jim" last="Cooley">Jim Cooley</name><affiliation wicri:level="1"><nlm:aff id="aff5"><institution>Department of Population Medicine and Pathobiology, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Smith, David F" sort="Smith, David F" uniqKey="Smith D" first="David F" last="Smith">David F. Smith</name><affiliation wicri:level="1"><nlm:aff id="aff6"><institution>Department of Biochemistry, Glycomics Center, School of Medicine, Emory University</institution>, Atlanta 30322, GA,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Liao, Ming" sort="Liao, Ming" uniqKey="Liao M" first="Ming" last="Liao">Ming Liao</name><affiliation wicri:level="1"><nlm:aff id="aff3"><institution>College of Veterinary Medicine, South China Agricultural University</institution>, Guangzhou 510462, Guangdong,<country>China</country></nlm:aff><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Cardona, Carol" sort="Cardona, Carol" uniqKey="Cardona C" first="Carol" last="Cardona">Carol Cardona</name><affiliation wicri:level="1"><nlm:aff id="aff7"><institution>College of Veterinary Medicine, University of Minnesota</institution>, St Paul, MN 30322,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Li, Lei" sort="Li, Lei" uniqKey="Li L" first="Lei" last="Li">Lei Li</name><affiliation wicri:level="1"><nlm:aff id="aff8"><institution>Department of Chemistry, Georgia State University</institution>, Atlanta, GA 30302,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Wang, George Peng" sort="Wang, George Peng" uniqKey="Wang G" first="George Peng" last="Wang">George Peng Wang</name><affiliation wicri:level="1"><nlm:aff id="aff8"><institution>Department of Chemistry, Georgia State University</institution>, Atlanta, GA 30302,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Webby, Richard" sort="Webby, Richard" uniqKey="Webby R" first="Richard" last="Webby">Richard Webby</name><affiliation wicri:level="1"><nlm:aff id="aff2"><institution>Department of Infectious Diseases, St Jude Children's Research Hospital</institution>, Memphis, TN 38105,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Wan, Xiu Feng" sort="Wan, Xiu Feng" uniqKey="Wan X" first="Xiu-Feng" last="Wan">Xiu-Feng Wan</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">28400591</idno><idno type="pmc">5457681</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5457681</idno><idno type="RBID">PMC:5457681</idno><idno type="doi">10.1038/emi.2017.3</idno><date when="2017">2017</date><idno type="wicri:Area/Pmc/Corpus">000090</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000090</idno><idno type="wicri:Area/Pmc/Curation">000090</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Curation">000090</idno><idno type="wicri:Area/Pmc/Checkpoint">000209</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Checkpoint">000209</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Pathogenicity and transmission of a swine influenza A(H6N6) virus</title><author><name sortKey="Sun, Hailiang" sort="Sun, Hailiang" uniqKey="Sun H" first="Hailiang" last="Sun">Hailiang Sun</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Kaplan, Bryan S" sort="Kaplan, Bryan S" uniqKey="Kaplan B" first="Bryan S" last="Kaplan">Bryan S. Kaplan</name><affiliation wicri:level="1"><nlm:aff id="aff2"><institution>Department of Infectious Diseases, St Jude Children's Research Hospital</institution>, Memphis, TN 38105,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Guan, Minhui" sort="Guan, Minhui" uniqKey="Guan M" first="Minhui" last="Guan">Minhui Guan</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Zhang, Guihong" sort="Zhang, Guihong" uniqKey="Zhang G" first="Guihong" last="Zhang">Guihong Zhang</name><affiliation wicri:level="1"><nlm:aff id="aff3"><institution>College of Veterinary Medicine, South China Agricultural University</institution>, Guangzhou 510462, Guangdong,<country>China</country></nlm:aff><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Ye, Jianqiang" sort="Ye, Jianqiang" uniqKey="Ye J" first="Jianqiang" last="Ye">Jianqiang Ye</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Long, Li Ping" sort="Long, Li Ping" uniqKey="Long L" first="Li-Ping" last="Long">Li-Ping Long</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Blackmon, Sherry" sort="Blackmon, Sherry" uniqKey="Blackmon S" first="Sherry" last="Blackmon">Sherry Blackmon</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Yang, Chun Kai" sort="Yang, Chun Kai" uniqKey="Yang C" first="Chun-Kai" last="Yang">Chun-Kai Yang</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Chiang, Meng Jung" sort="Chiang, Meng Jung" uniqKey="Chiang M" first="Meng-Jung" last="Chiang">Meng-Jung Chiang</name><affiliation wicri:level="1"><nlm:aff id="aff4"><institution>Laboratory of Respiratory Viral Diseases, Division of Viral Products, Office of Vaccines Research and Review, Center for Biologics Evaluation and Research, United States Food and Drug Administration</institution>, Silver Spring, MD 20993,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Xie, Hang" sort="Xie, Hang" uniqKey="Xie H" first="Hang" last="Xie">Hang Xie</name><affiliation wicri:level="1"><nlm:aff id="aff4"><institution>Laboratory of Respiratory Viral Diseases, Division of Viral Products, Office of Vaccines Research and Review, Center for Biologics Evaluation and Research, United States Food and Drug Administration</institution>, Silver Spring, MD 20993,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Zhao, Nan" sort="Zhao, Nan" uniqKey="Zhao N" first="Nan" last="Zhao">Nan Zhao</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Cooley, Jim" sort="Cooley, Jim" uniqKey="Cooley J" first="Jim" last="Cooley">Jim Cooley</name><affiliation wicri:level="1"><nlm:aff id="aff5"><institution>Department of Population Medicine and Pathobiology, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Smith, David F" sort="Smith, David F" uniqKey="Smith D" first="David F" last="Smith">David F. Smith</name><affiliation wicri:level="1"><nlm:aff id="aff6"><institution>Department of Biochemistry, Glycomics Center, School of Medicine, Emory University</institution>, Atlanta 30322, GA,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Liao, Ming" sort="Liao, Ming" uniqKey="Liao M" first="Ming" last="Liao">Ming Liao</name><affiliation wicri:level="1"><nlm:aff id="aff3"><institution>College of Veterinary Medicine, South China Agricultural University</institution>, Guangzhou 510462, Guangdong,<country>China</country></nlm:aff><country xml:lang="fr">République populaire de Chine</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Cardona, Carol" sort="Cardona, Carol" uniqKey="Cardona C" first="Carol" last="Cardona">Carol Cardona</name><affiliation wicri:level="1"><nlm:aff id="aff7"><institution>College of Veterinary Medicine, University of Minnesota</institution>, St Paul, MN 30322,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Li, Lei" sort="Li, Lei" uniqKey="Li L" first="Lei" last="Li">Lei Li</name><affiliation wicri:level="1"><nlm:aff id="aff8"><institution>Department of Chemistry, Georgia State University</institution>, Atlanta, GA 30302,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Wang, George Peng" sort="Wang, George Peng" uniqKey="Wang G" first="George Peng" last="Wang">George Peng Wang</name><affiliation wicri:level="1"><nlm:aff id="aff8"><institution>Department of Chemistry, Georgia State University</institution>, Atlanta, GA 30302,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Webby, Richard" sort="Webby, Richard" uniqKey="Webby R" first="Richard" last="Webby">Richard Webby</name><affiliation wicri:level="1"><nlm:aff id="aff2"><institution>Department of Infectious Diseases, St Jude Children's Research Hospital</institution>, Memphis, TN 38105,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Wan, Xiu Feng" sort="Wan, Xiu Feng" uniqKey="Wan X" first="Xiu-Feng" last="Wan">Xiu-Feng Wan</name><affiliation wicri:level="1"><nlm:aff id="aff1"><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author></analytic><series><title level="j">Emerging Microbes & Infections</title><idno type="eISSN">2222-1751</idno><imprint><date when="2017">2017</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Subtype H6 influenza A viruses (IAVs) are commonly detected in wild birds and domestic poultry and can infect humans. In 2010, a H6N6 virus emerged in southern China, and since then, it has caused sporadic infections among swine. We show that this virus binds to α2,6-linked and α2,3-linked sialic acids. Mutations at residues 222 (alanine to valine) and 228 (glycine to serine) of the virus hemagglutinin (HA) affected its receptor-binding properties. Experiments showed that the virus has limited transmissibility between ferrets through direct contact or through inhalation of infectious aerosolized droplets. The internal genes of the influenza A(H1N1)pdm09 virus, which is prevalent in swine worldwide, increases the replication efficiency of H6N6 IAV in the lower respiratory tract of ferrets but not its transmissibility between ferrets. These findings suggest H6N6 swine IAV (SIV) currently poses a moderate risk to public health, but its evolution and spread should be closely monitored.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct><biblStruct></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Emerg Microbes Infect</journal-id><journal-id journal-id-type="iso-abbrev">Emerg Microbes Infect</journal-id><journal-title-group><journal-title>Emerging Microbes & Infections</journal-title></journal-title-group><issn pub-type="epub">2222-1751</issn><publisher><publisher-name>Nature Publishing Group</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">28400591</article-id><article-id pub-id-type="pmc">5457681</article-id><article-id pub-id-type="pii">emi20173</article-id><article-id pub-id-type="doi">10.1038/emi.2017.3</article-id><article-categories><subj-group subj-group-type="heading"><subject>Original Article</subject></subj-group></article-categories><title-group><article-title>Pathogenicity and transmission of a swine influenza A(H6N6) virus</article-title><alt-title alt-title-type="running">Risk assessment of H6N6 swine influenza A virus</alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Sun</surname><given-names>Hailiang</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Kaplan</surname><given-names>Bryan S</given-names></name><xref ref-type="aff" rid="aff2">2</xref></contrib><contrib contrib-type="author"><name><surname>Guan</surname><given-names>Minhui</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Zhang</surname><given-names>Guihong</given-names></name><xref ref-type="aff" rid="aff3">3</xref></contrib><contrib contrib-type="author"><name><surname>Ye</surname><given-names>Jianqiang</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Long</surname><given-names>Li-Ping</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Blackmon</surname><given-names>Sherry</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Yang</surname><given-names>Chun-Kai</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Chiang</surname><given-names>Meng-Jung</given-names></name><xref ref-type="aff" rid="aff4">4</xref></contrib><contrib contrib-type="author"><name><surname>Xie</surname><given-names>Hang</given-names></name><xref ref-type="aff" rid="aff4">4</xref></contrib><contrib contrib-type="author"><name><surname>Zhao</surname><given-names>Nan</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><contrib contrib-type="author"><name><surname>Cooley</surname><given-names>Jim</given-names></name><xref ref-type="aff" rid="aff5">5</xref></contrib><contrib contrib-type="author"><name><surname>Smith</surname><given-names>David F</given-names></name><xref ref-type="aff" rid="aff6">6</xref></contrib><contrib contrib-type="author"><name><surname>Liao</surname><given-names>Ming</given-names></name><xref ref-type="aff" rid="aff3">3</xref></contrib><contrib contrib-type="author"><name><surname>Cardona</surname><given-names>Carol</given-names></name><xref ref-type="aff" rid="aff7">7</xref></contrib><contrib contrib-type="author"><name><surname>Li</surname><given-names>Lei</given-names></name><xref ref-type="aff" rid="aff8">8</xref></contrib><contrib contrib-type="author"><name><surname>Wang</surname><given-names>George Peng</given-names></name><xref ref-type="aff" rid="aff8">8</xref></contrib><contrib contrib-type="author"><name><surname>Webby</surname><given-names>Richard</given-names></name><xref ref-type="aff" rid="aff2">2</xref></contrib><contrib contrib-type="author"><name><surname>Wan</surname><given-names>Xiu-Feng</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="corresp" rid="caf1">*</xref></contrib><aff id="aff1"><label>1</label><institution>Department of Basic Sciences, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></aff><aff id="aff2"><label>2</label><institution>Department of Infectious Diseases, St Jude Children's Research Hospital</institution>, Memphis, TN 38105,<country>USA</country></aff><aff id="aff3"><label>3</label><institution>College of Veterinary Medicine, South China Agricultural University</institution>, Guangzhou 510462, Guangdong,<country>China</country></aff><aff id="aff4"><label>4</label><institution>Laboratory of Respiratory Viral Diseases, Division of Viral Products, Office of Vaccines Research and Review, Center for Biologics Evaluation and Research, United States Food and Drug Administration</institution>, Silver Spring, MD 20993,<country>USA</country></aff><aff id="aff5"><label>5</label><institution>Department of Population Medicine and Pathobiology, College of Veterinary Medicine, Mississippi State University</institution>, Mississippi State, MS 39759,<country>USA</country></aff><aff id="aff6"><label>6</label><institution>Department of Biochemistry, Glycomics Center, School of Medicine, Emory University</institution>, Atlanta 30322, GA,<country>USA</country></aff><aff id="aff7"><label>7</label><institution>College of Veterinary Medicine, University of Minnesota</institution>, St Paul, MN 30322,<country>USA</country></aff><aff id="aff8"><label>8</label><institution>Department of Chemistry, Georgia State University</institution>, Atlanta, GA 30302,<country>USA</country></aff></contrib-group><author-notes><corresp id="caf1"><label>*</label>E-mail: <email>wan@cvm.msstate.edu</email></corresp></author-notes><pub-date pub-type="ppub"><month>04</month><year>2017</year></pub-date><pub-date pub-type="epub"><day>12</day><month>04</month><year>2017</year></pub-date><pub-date pub-type="pmc-release"><day>1</day><month>4</month><year>2017</year></pub-date><volume>6</volume><issue>4</issue><fpage>e17</fpage><lpage></lpage><history><date date-type="received"><day>18</day><month>10</month><year>2016</year></date><date date-type="rev-recd"><day>15</day><month>12</month><year>2016</year></date><date date-type="accepted"><day>29</day><month>12</month><year>2016</year></date></history><permissions><copyright-statement>Copyright © 2017 The Author(s)</copyright-statement><copyright-year>2017</copyright-year><copyright-holder>The Author(s)</copyright-holder><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/4.0/"><pmc-comment>author-paid</pmc-comment>
<license-p>This work is licensed under a Creative Commons Attribution 4.0 International License. The images or other third party material in this article are included in the article's Creative Commons license, unless indicated otherwise in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. To view a copy of this license, visit <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">http://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><abstract><p>Subtype H6 influenza A viruses (IAVs) are commonly detected in wild birds and domestic poultry and can infect humans. In 2010, a H6N6 virus emerged in southern China, and since then, it has caused sporadic infections among swine. We show that this virus binds to α2,6-linked and α2,3-linked sialic acids. Mutations at residues 222 (alanine to valine) and 228 (glycine to serine) of the virus hemagglutinin (HA) affected its receptor-binding properties. Experiments showed that the virus has limited transmissibility between ferrets through direct contact or through inhalation of infectious aerosolized droplets. The internal genes of the influenza A(H1N1)pdm09 virus, which is prevalent in swine worldwide, increases the replication efficiency of H6N6 IAV in the lower respiratory tract of ferrets but not its transmissibility between ferrets. These findings suggest H6N6 swine IAV (SIV) currently poses a moderate risk to public health, but its evolution and spread should be closely monitored.</p></abstract><kwd-group><kwd>2009 H1N1 influenza virus</kwd><kwd>aerosol transmission</kwd><kwd>glycan microarray</kwd><kwd>influenza A(H1N1)pdm09 virus</kwd><kwd>receptor binding</kwd><kwd>risk assessment</kwd><kwd>subtype H6N6 influenza virus</kwd><kwd>swine influenza virus</kwd></kwd-group></article-meta></front><floats-group><fig id="fig1"><label>Figure 1</label><caption><p>Phylogenetic analyses of subtype H6N6 swine influenza virus A/swine/Guangdong/k6/2010(H6N6). (<bold>A</bold>) HA gene. (<bold>B</bold>) NA gene. (<bold>C</bold>) PB2 gene. (<bold>D</bold>) PB1 gene. (<bold>E</bold>) PA gene. (<bold>F</bold>) NP gene. (<bold>G</bold>) MP gene. (<bold>H</bold>) NS gene. The phylogenetic trees were constructed by using maximum-likelihood implemented in GARLI version 0.96;<sup><xref ref-type="bibr" rid="bib40">40</xref></sup> bootstrap values were generated, with 1000 replications, by using neighbor-joining methods implemented in PAUP* 4.0 Beta.<sup><xref ref-type="bibr" rid="bib41">41</xref></sup> Scale bars represent nucleotide substitutions per site. The virus marked in red indicates the H6N6 strain isolated from swine; the virus marked in green indicates the subtype H6N1 avian influenza strain isolated from a human in Taiwan.</p></caption><graphic xlink:href="emi20173f1a"></graphic><graphic xlink:href="emi20173f1b"></graphic><graphic xlink:href="emi20173f1c"></graphic><graphic xlink:href="emi20173f1d"></graphic></fig><fig id="fig2"><label>Figure 2</label><caption><p>Virus-binding affinities of recombinant influenza A(H6N6) viruses to glycans. Binding affinities of viruses to specific biotinylated Neu5Acα2,3-Gal glycans (<bold>A</bold>) and to specific biotinylated Neu5Acα2,6-Gal glycans (<bold>B</bold>). Four viruses were used: rgH6N6 × PR8-222V/228S, rgH6N6 × PR8-222A/228S, rgH6N6 × PR8-222V/228G and rgH6N6 × PR8-222A/228G. Six biotinlyated glycans were used, including two analogs (3′SLN and 6′SLN) and four synthetic <italic>N</italic>-linked glycans (N32, N33, N52 and N53). 3′SLN, N32 and N52 were used to represent as Neu5Acα2,3-Gal; 6′SLN, N33 and N53 were used to represent as Neu5Acα2,6-Gal.</p></caption><graphic xlink:href="emi20173f2"></graphic></fig><fig id="fig3"><label>Figure 3</label><caption><p>The binding profile of recombinant influenza A(H6N6) viruses to sialic acids on the glycan array. Colors indicate different categories of glycans on the array: red, α2,3-sulfated sialosides; green, α2,3- di-, tri- and qua-sialosides; yellow, α2,3-linear sialosides; purple, α2,3-fucosylated sialosides; blue, α2,3-internal sialosides; black, α2,3- and α2,6-sialosides; and gray, different categories of α2,6-sialosides are highlighted in the same order as listed for the α2,3-sialosides. The dashed line indicates minimum relative fluorescence units of 2000. Black bars represent error bars. The glycan sequences are detailed in <xref ref-type="supplementary-material" rid="sup1">Supplementary Table S1</xref>.</p></caption><graphic xlink:href="emi20173f3"></graphic></fig><fig id="fig4"><label>Figure 4</label><caption><p>Mean titers of influenza viruses recovered from nasal wash fluids of virus inoculated and contact ferrets in direct-contact transmission experiments. Inoculated ferrets were nasally inoculated with 10<sup>6</sup> TCID<sub>50</sub> of each virus shown: the wild-type-like rgH6N6-222V/228S (<bold>A</bold>) and rgH6N6xpdm09-222V/228S (<bold>B</bold>), and 1 day later each was housed in the same cage with a contact ferret. Virus titers were measured in nasal wash fluids (collected on indicated days) by using endpoint titration in Madin-Darby canine kidney cells; ending titers were expressed as mean log<sub>10</sub>TCID<sub>50</sub>/mL±standard deviation. The limit of virus detection was 10<sup>0.699</sup> TCID<sub>50</sub>/mL. the ferret #1, F#1; the ferret #2, F#2; the ferret #3, F#3; virus inoculated, VI; direct contact, DC.</p></caption><graphic xlink:href="emi20173f4"></graphic></fig><fig id="fig5"><label>Figure 5</label><caption><p>Mean titers of influenza viruses recovered from nasal wash fluids of virus-inoculated and contact ferrets in aerosol transmission experiments. Inoculated ferrets were inoculated with 10<sup>6</sup> 50% tissue culture infectious doses (TCID<sub>50</sub>) of each virus shown: the wild-type-like rgH6N6-222V/228S (<bold>A</bold>) and rgH6N6xpdm-222V/228S (<bold>B</bold>), and 1 day later, a contact ferret was housed in the same cage as an inoculated ferret but on a different side of a 1-cm-thick, double-layered, steel partition with 5-mm perforations (Allentown Inc.). Virus titers were measured in nasal wash fluids (collected on indicated days) by using endpoint titration in Madin-Darby canine kidney cells; ending titers were expressed as mean log<sub>10</sub>TCID<sub>50</sub>/mL±standard deviation. The limit of virus detection was 10<sup>0.699</sup> TCID<sub>50</sub>/mL. aerosol exposure, AE; the ferret #1, F#1; the ferret #2, F#2; the ferret #3, F#3; virus inoculated, VI.</p></caption><graphic xlink:href="emi20173f5"></graphic></fig><fig id="fig6"><label>Figure 6</label><caption><p>Mean titers of influenza viruses recovered from respiratory tract tissues of ferrets after nasal inoculation of virus. Ferrets were inoculated with 10<sup>6</sup> 50% tissue culture infectious doses (TCID<sub>50</sub>) of each virus shown: the wild-type-like rgH6N6-222V/228S and rgH6N6xpdm09-222V/228S. Two ferrets were killed three days after inoculation, and virus titers in nasal turbinate, trachea, bronchus and lung of each ferret were determined by using end-point titration in Madin-Darby canine kidney cells. The titers were quantified as the mean titer from at least three sections of each tissue. The limit of virus detection was 10<sup>0.699</sup> TCID<sub>50</sub>/mL. The dashed line indicates the lower limit of detection. Black bars represent error bars.</p></caption><graphic xlink:href="emi20173f6"></graphic></fig><table-wrap id="tbl1"><label>Table 1</label><caption><title>Recombinant influenza A(H6N6) viruses generated in this study</title></caption><table frame="hsides" rules="groups" border="1"><colgroup><col align="left"></col><col align="center"></col><col align="center"></col></colgroup><thead valign="bottom"><tr><th align="left" valign="top" charoff="50"><bold>Strain name<xref ref-type="fn" rid="t1-fn1">a</xref></bold></th><th align="center" valign="top" charoff="50"><bold>HA and NA genes</bold></th><th align="center" valign="top" charoff="50"><bold>PB2, PB1, PA, NP, MP and NS genes</bold></th></tr></thead><tbody valign="top"><tr><td align="left" valign="top" charoff="50">rgH6N6-222V/228S</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6-222A/228G</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × PR8-222V/228S</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/Puerto Rico/8/1934(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × PR8-222A/228S</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/Puerto Rico/8/1934(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × PR8-222V/228G</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/Puerto Rico/8/1934(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × PR8-222A/228G</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/Puerto Rico/8/1934(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × pdm09-222V/228S</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/California/4/2009(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × pdm09-222A/228S</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/California/4/2009(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × pdm09-222V/228G</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/California/4/2009(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH6N6 × pdm09-222A/228G</td><td align="center" valign="top" charoff="50">A/swine/Guangdong/K6/2010(H6N6)</td><td align="center" valign="top" charoff="50">A/California/4/2009(H1N1)</td></tr><tr><td align="left" valign="top" charoff="50">rgH3N2 × pdm09</td><td align="center" valign="top" charoff="50">A/swine/Ohio/11SW226/2011(H3N2)</td><td align="center" valign="top" charoff="50">A/California/4/2009(H1N1)</td></tr></tbody></table><table-wrap-foot><fn id="t1-fn1"><label>a</label><p>rg indicates reverse genetics.</p></fn></table-wrap-foot></table-wrap><table-wrap id="tbl2"><label>Table 2</label><caption><title>Effect of mutations A222V and G228S of influenza A(H6N6) virus hemagglutinin on binding affinity to erythrocytes from various animals<xref ref-type="fn" rid="t2-fn1">a</xref></title></caption><table frame="hsides" rules="groups" border="1"><colgroup><col align="left"></col><col align="center"></col><col align="center"></col><col align="center"></col><col align="center"></col></colgroup><thead valign="bottom"><tr><th align="left" valign="top" charoff="50">Virus</th><th colspan="4" align="center" valign="top" charoff="50">Mean hemagglutination titer<xref ref-type="fn" rid="t2-fn2">b</xref><hr></hr></th></tr><tr><th align="left" valign="top" charoff="50"> </th><th align="center" valign="top" charoff="50">Turkey</th><th align="center" valign="top" charoff="50">Chicken</th><th align="center" valign="top" charoff="50">Guinea pig</th><th align="center" valign="top" charoff="50">Horse</th></tr></thead><tbody valign="top"><tr><td align="left" valign="top" charoff="50"><italic>Types of erythrocytes</italic></td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × PR8-222V/228S</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × PR8-222A/228S</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">16</td><td align="center" valign="top" charoff="50">32</td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × PR8-222V/228G</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">64</td><td align="center" valign="top" charoff="50"><2</td><td align="center" valign="top" charoff="50">2</td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × PR8-222A/228G</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">64</td><td align="center" valign="top" charoff="50"><2</td><td align="center" valign="top" charoff="50"><2</td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × pdm09-222V/228S</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">16</td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × pdm09-222A/228S</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">32</td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × pdm09-222V/228G</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">64</td><td align="center" valign="top" charoff="50"><2</td><td align="center" valign="top" charoff="50"><2</td></tr><tr><td align="left" valign="top" charoff="50"> rgH6N6 × pdm09-222A/228G</td><td align="center" valign="top" charoff="50">32</td><td align="center" valign="top" charoff="50">128</td><td align="center" valign="top" charoff="50"><2</td><td align="center" valign="top" charoff="50">2</td></tr></tbody></table><table-wrap-foot><fn id="t2-fn1"><label>a</label><p>rg indicates reverse genetics.</p></fn><fn id="t2-fn2"><label>b</label><p>All virus concentrations were normalized to a hemagglutination titer of 32 in turkey erythrocytes. Each assay was repeated three times, and in each case, the standard deviation was 0.</p></fn></table-wrap-foot></table-wrap><table-wrap id="tbl3"><label>Table 3</label><caption><title>Hemagglutination inhibition titers in serum samples from ferrets inoculated with or exposed to animals inoculated with influenza A(H6N6) virus</title></caption><table frame="hsides" rules="groups" border="1"><colgroup><col align="left"></col><col align="center"></col><col align="center"></col><col align="center"></col><col align="center"></col></colgroup><thead valign="bottom"><tr><th align="center" valign="top" charoff="50">Virus and experiment group<xref ref-type="fn" rid="t3-fn1">a</xref></th><th align="center" valign="top" charoff="50">Time of serum collection<xref ref-type="fn" rid="t3-fn2">b</xref></th><th colspan="3" align="center" valign="top" charoff="50">Titer<xref ref-type="fn" rid="t3-fn3">c</xref></th></tr></thead><tbody valign="top"><tr><td colspan="5" align="left" valign="top" charoff="50"><italic>rgH6N6-222V/228S</italic></td></tr><tr><td align="left" valign="top" charoff="50"> Aerosol transmission study</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="left" valign="top" charoff="50"> Virus inoculated</td><td align="center" valign="top" charoff="50">14 DPI</td><td align="center" valign="top" charoff="50">640</td><td align="center" valign="top" charoff="50">320</td><td align="center" valign="top" charoff="50">1280</td></tr><tr><td align="left" valign="top" charoff="50"> Exposure by aerosol</td><td align="center" valign="top" charoff="50">14 DPE</td><td align="center" valign="top" charoff="50"><10</td><td align="center" valign="top" charoff="50"><10</td><td align="center" valign="top" charoff="50"><10</td></tr><tr><td align="left" valign="top" charoff="50"> Contact transmission study</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="left" valign="top" charoff="50"> Virus inoculated</td><td align="center" valign="top" charoff="50">14 DPI</td><td align="center" valign="top" charoff="50">640</td><td align="center" valign="top" charoff="50">640</td><td align="center" valign="top" charoff="50">1280</td></tr><tr><td align="left" valign="top" charoff="50"> Exposure by direct contact</td><td align="center" valign="top" charoff="50">14 DPE</td><td align="center" valign="top" charoff="50"><10</td><td align="center" valign="top" charoff="50"><10</td><td align="center" valign="top" charoff="50">20</td></tr><tr><td colspan="5" align="left" valign="top" charoff="50"> </td></tr><tr><td colspan="5" align="left" valign="top" charoff="50"><italic>rgH6N6 × pdm09-222V/228S</italic></td></tr><tr><td align="left" valign="top" charoff="50"> Aerosol transmission study</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="left" valign="top" charoff="50"> Virus inoculated</td><td align="center" valign="top" charoff="50">14 DPI</td><td align="center" valign="top" charoff="50">1280</td><td align="center" valign="top" charoff="50">1280</td><td align="center" valign="top" charoff="50">1280</td></tr><tr><td align="left" valign="top" charoff="50"> Exposure by aerosol</td><td align="center" valign="top" charoff="50">14 DPE</td><td align="center" valign="top" charoff="50"><10</td><td align="center" valign="top" charoff="50"><10</td><td align="center" valign="top" charoff="50"><10</td></tr><tr><td align="left" valign="top" charoff="50"> Contact transmission study</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="left" valign="top" charoff="50"> Virus inoculated</td><td align="center" valign="top" charoff="50">14 DPI</td><td align="center" valign="top" charoff="50">1280</td><td align="center" valign="top" charoff="50">1280</td><td align="center" valign="top" charoff="50">640</td></tr><tr><td align="left" valign="top" charoff="50"> Exposure by direct contact</td><td align="center" valign="top" charoff="50">14 DPE</td><td align="center" valign="top" charoff="50">640</td><td align="center" valign="top" charoff="50">640</td><td align="center" valign="top" charoff="50">640</td></tr></tbody></table><table-wrap-foot><fn id="t3-fn1"><label>a</label><p>rg indicates reverse genetics.</p></fn><fn id="t3-fn2"><label>b</label><p>Day post exposure, DPE; day post inoculation, DPI.</p></fn><fn id="t3-fn3"><label>c</label><p>Hemagglutination inhibition assays were performed using turkey erythrocytes. The titers were generated from sera collected from three individual ferrets.</p></fn></table-wrap-foot></table-wrap><table-wrap id="tbl4"><label>Table 4</label><caption><title>Virus titers in nasal wash fluids of ferrets in aerosol transmission experiment nasally inoculated with influenza A(H6N6) virus</title></caption><table frame="hsides" rules="groups" border="1"><colgroup><col align="left"></col><col align="center"></col><col align="center"></col><col align="center"></col></colgroup><thead valign="bottom"><tr><th align="left" valign="top" charoff="50">Virus and time (DPI or DPE) of nasal wash fluid collection<xref ref-type="fn" rid="t4-fn2">a</xref></th><th colspan="3" align="center" valign="top" charoff="50">Mean virus titer (log<sub>10</sub>EID<sub>50</sub>/mL)<xref ref-type="fn" rid="t4-fn3">b</xref></th></tr></thead><tbody valign="top"><tr><td colspan="4" align="left" valign="top" charoff="50"><italic>rgH6N6-222V/228S</italic></td></tr><tr><td align="left" valign="top" charoff="50"> 2</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td></tr><tr><td align="left" valign="top" charoff="50"> 4</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td></tr><tr><td align="left" valign="top" charoff="50"> 6</td><td align="char" valign="top" char="." charoff="50">1.00</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td></tr><tr><td align="left" valign="top" charoff="50"> 8</td><td align="char" valign="top" char="." charoff="50">1.00</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"> </td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"><italic>rgH6N6 × pdm09-222V/228S</italic></td></tr><tr><td align="left" valign="top" charoff="50"> 2</td><td align="center" valign="top" charoff="50">ND</td><td align="char" valign="top" char="." charoff="50">1.00</td><td align="center" valign="top" charoff="50">ND</td></tr><tr><td align="left" valign="top" charoff="50"> 4</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td></tr><tr><td align="left" valign="top" charoff="50"> 6</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td></tr><tr><td align="left" valign="top" charoff="50"> 8</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td><td align="center" valign="top" charoff="50">ND</td></tr></tbody></table><table-wrap-foot><fn id="t4-fn1"><p>Abbreviations: day post exposure, DPE; day post inoculation, DPI; egg infectious dose, EID; not detected, ND.</p></fn><fn id="t4-fn2"><label>a</label><p>Data are for virus inoculated ferrets and direct- and aerosol-contact exposure ferrets at various DPI or DPE.</p></fn><fn id="t4-fn3"><label>b</label><p>Virus titers were determined by 50% egg infectious dose (EID<sub>50</sub>) in embryonated chicken eggs. The titers were generated from nasal swabs collected from three individual ferrets.</p></fn></table-wrap-foot></table-wrap></floats-group></pmc><affiliations><list><country><li>République populaire de Chine</li><li>États-Unis</li></country></list><tree><country name="États-Unis"><noRegion><name sortKey="Sun, Hailiang" sort="Sun, Hailiang" uniqKey="Sun H" first="Hailiang" last="Sun">Hailiang Sun</name></noRegion><name sortKey="Blackmon, Sherry" sort="Blackmon, Sherry" uniqKey="Blackmon S" first="Sherry" last="Blackmon">Sherry Blackmon</name><name sortKey="Cardona, Carol" sort="Cardona, Carol" uniqKey="Cardona C" first="Carol" last="Cardona">Carol Cardona</name><name sortKey="Chiang, Meng Jung" sort="Chiang, Meng Jung" uniqKey="Chiang M" first="Meng-Jung" last="Chiang">Meng-Jung Chiang</name><name sortKey="Cooley, Jim" sort="Cooley, Jim" uniqKey="Cooley J" first="Jim" last="Cooley">Jim Cooley</name><name sortKey="Guan, Minhui" sort="Guan, Minhui" uniqKey="Guan M" first="Minhui" last="Guan">Minhui Guan</name><name sortKey="Kaplan, Bryan S" sort="Kaplan, Bryan S" uniqKey="Kaplan B" first="Bryan S" last="Kaplan">Bryan S. Kaplan</name><name sortKey="Li, Lei" sort="Li, Lei" uniqKey="Li L" first="Lei" last="Li">Lei Li</name><name sortKey="Long, Li Ping" sort="Long, Li Ping" uniqKey="Long L" first="Li-Ping" last="Long">Li-Ping Long</name><name sortKey="Smith, David F" sort="Smith, David F" uniqKey="Smith D" first="David F" last="Smith">David F. Smith</name><name sortKey="Wan, Xiu Feng" sort="Wan, Xiu Feng" uniqKey="Wan X" first="Xiu-Feng" last="Wan">Xiu-Feng Wan</name><name sortKey="Wang, George Peng" sort="Wang, George Peng" uniqKey="Wang G" first="George Peng" last="Wang">George Peng Wang</name><name sortKey="Webby, Richard" sort="Webby, Richard" uniqKey="Webby R" first="Richard" last="Webby">Richard Webby</name><name sortKey="Xie, Hang" sort="Xie, Hang" uniqKey="Xie H" first="Hang" last="Xie">Hang Xie</name><name sortKey="Yang, Chun Kai" sort="Yang, Chun Kai" uniqKey="Yang C" first="Chun-Kai" last="Yang">Chun-Kai Yang</name><name sortKey="Ye, Jianqiang" sort="Ye, Jianqiang" uniqKey="Ye J" first="Jianqiang" last="Ye">Jianqiang Ye</name><name sortKey="Zhao, Nan" sort="Zhao, Nan" uniqKey="Zhao N" first="Nan" last="Zhao">Nan Zhao</name></country><country name="République populaire de Chine"><noRegion><name sortKey="Zhang, Guihong" sort="Zhang, Guihong" uniqKey="Zhang G" first="Guihong" last="Zhang">Guihong Zhang</name></noRegion><name sortKey="Liao, Ming" sort="Liao, Ming" uniqKey="Liao M" first="Ming" last="Liao">Ming Liao</name></country></tree></affiliations></record>Pour manipuler ce document sous Unix (Dilib)
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