Characterization of a Novel Reassortant Influenza A Virus (H2N2) from a Domestic Duck in Eastern China
Identifieur interne : 000008 ( Pmc/Curation ); précédent : 000007; suivant : 000009Characterization of a Novel Reassortant Influenza A Virus (H2N2) from a Domestic Duck in Eastern China
Auteurs : Mai-Juan Ma [République populaire de Chine] ; Xiao-Xian Yang [République populaire de Chine] ; Yan-Hua Qian [République populaire de Chine] ; Si-Yan Zhao [République populaire de Chine] ; Sha Hua [République populaire de Chine] ; Tie-Cheng Wang [République populaire de Chine] ; Shan-Hui Chen [République populaire de Chine] ; Guang-Yuan Ma [République populaire de Chine] ; Xiao-Yu Sang [République populaire de Chine] ; Lin-Na Liu [République populaire de Chine] ; Ai-Ping Wu [République populaire de Chine] ; Tai-Jiao Jiang [République populaire de Chine] ; Yu-Wei Gao [République populaire de Chine] ; Gregory C. Gray [États-Unis] ; Teng Zhao [République populaire de Chine] ; Xia Ling [République populaire de Chine] ; Jing-Lin Wang [République populaire de Chine] ; Bing Lu [République populaire de Chine] ; Jun Qian [République populaire de Chine] ; Wu-Chun Cao [République populaire de Chine]Source :
- Scientific Reports [ 2045-2322 ] ; 2014.
Abstract
While H2N2 viruses have been sporadically isolated from wild and domestic birds, H2N2 viruses have not been detected among human populations since 1968. Should H2N2 viruses adapt to domestic poultry they may pose a risk of infection to people, as most anyone born after 1968 would likely be susceptible to their infection. We report the isolation of a novel influenza A virus (H2N2) cultured in 2013 from a healthy domestic duck at a live poultry market in Wuxi City, China. Sequence data revealed that the novel H2N2 virus was similar to Eurasian avian lineage avian influenza viruses, the virus had been circulating for ≥ two years among poultry, had an increase in α2,6 binding affinity, and was not highly pathogenic. Approximately 9% of 100 healthy chickens sampled from the same area had elevated antibodies against the H2 antigen. Fortunately, there was sparse serological evidence that the virus was infecting poultry workers or had adapted to infect other mammals. These findings suggest that a novel H2N2 virus has been circulating among domestic poultry in Wuxi City, China and has some has increased human receptor affinity. It seems wise to conduct better surveillance for novel influenza viruses at Chinese live bird markets.
Url:
DOI: 10.1038/srep07588
PubMed: 25533850
PubMed Central: 4274511
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<author><name sortKey="Yang, Xiao Xian" sort="Yang, Xiao Xian" uniqKey="Yang X" first="Xiao-Xian" last="Yang">Xiao-Xian Yang</name>
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, Beijing, 100071, P. R. China</nlm:aff>
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, Changchun, 130122, P. R. China</nlm:aff>
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<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
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<author><name sortKey="Sang, Xiao Yu" sort="Sang, Xiao Yu" uniqKey="Sang X" first="Xiao-Yu" last="Sang">Xiao-Yu Sang</name>
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<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
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<author><name sortKey="Liu, Lin Na" sort="Liu, Lin Na" uniqKey="Liu L" first="Lin-Na" last="Liu">Lin-Na Liu</name>
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<author><name sortKey="Wu, Ai Ping" sort="Wu, Ai Ping" uniqKey="Wu A" first="Ai-Ping" last="Wu">Ai-Ping Wu</name>
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<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
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<author><name sortKey="Jiang, Tai Jiao" sort="Jiang, Tai Jiao" uniqKey="Jiang T" first="Tai-Jiao" last="Jiang">Tai-Jiao Jiang</name>
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<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
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<author><name sortKey="Gao, Yu Wei" sort="Gao, Yu Wei" uniqKey="Gao Y" first="Yu-Wei" last="Gao">Yu-Wei Gao</name>
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<wicri:regionArea>, Changchun, 130122</wicri:regionArea>
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<affiliation wicri:level="1"><nlm:aff id="a8"><institution>Duke Infectious Disease & Duke Health Institute, Duke University Medical Center</institution>
, Durham, NC, 27710,<country>USA</country>
</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea># see nlm:aff country strict</wicri:regionArea>
</affiliation>
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<author><name sortKey="Zhao, Teng" sort="Zhao, Teng" uniqKey="Zhao T" first="Teng" last="Zhao">Teng Zhao</name>
<affiliation wicri:level="1"><nlm:aff id="a1"><institution>State Key Laboratory of Pathogen and Biosecurity, Beijing Institute of Microbiology and Epidemiology</institution>
, Beijing, 100071, P. R. China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>, Beijing, 100071</wicri:regionArea>
</affiliation>
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<author><name sortKey="Ling, Xia" sort="Ling, Xia" uniqKey="Ling X" first="Xia" last="Ling">Xia Ling</name>
<affiliation wicri:level="1"><nlm:aff id="a3"><institution>Wuxi Center for Disease Control and Prevention</institution>
, Wuxi, 214023, P. R. China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>, Wuxi, 214023</wicri:regionArea>
</affiliation>
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<author><name sortKey="Wang, Jing Lin" sort="Wang, Jing Lin" uniqKey="Wang J" first="Jing-Lin" last="Wang">Jing-Lin Wang</name>
<affiliation wicri:level="1"><nlm:aff id="a1"><institution>State Key Laboratory of Pathogen and Biosecurity, Beijing Institute of Microbiology and Epidemiology</institution>
, Beijing, 100071, P. R. China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>, Beijing, 100071</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Lu, Bing" sort="Lu, Bing" uniqKey="Lu B" first="Bing" last="Lu">Bing Lu</name>
<affiliation wicri:level="1"><nlm:aff id="a3"><institution>Wuxi Center for Disease Control and Prevention</institution>
, Wuxi, 214023, P. R. China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>, Wuxi, 214023</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Qian, Jun" sort="Qian, Jun" uniqKey="Qian J" first="Jun" last="Qian">Jun Qian</name>
<affiliation wicri:level="1"><nlm:aff id="a4"><institution>Military Veterinary Research Institute, Academy of Military Medical Sciences</institution>
, Changchun, 130122, P. R. China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>, Changchun, 130122</wicri:regionArea>
</affiliation>
</author>
<author><name sortKey="Cao, Wu Chun" sort="Cao, Wu Chun" uniqKey="Cao W" first="Wu-Chun" last="Cao">Wu-Chun Cao</name>
<affiliation wicri:level="1"><nlm:aff id="a1"><institution>State Key Laboratory of Pathogen and Biosecurity, Beijing Institute of Microbiology and Epidemiology</institution>
, Beijing, 100071, P. R. China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>, Beijing, 100071</wicri:regionArea>
</affiliation>
<affiliation wicri:level="1"><nlm:aff id="a2"><institution>School of Public Health, Central South University</institution>
, Changsha, Hunan, 410000 P. R. China</nlm:aff>
<country xml:lang="fr">République populaire de Chine</country>
<wicri:regionArea>, Changsha, Hunan</wicri:regionArea>
</affiliation>
</author>
</analytic>
<series><title level="j">Scientific Reports</title>
<idno type="eISSN">2045-2322</idno>
<imprint><date when="2014">2014</date>
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<front><div type="abstract" xml:lang="en"><p>While H2N2 viruses have been sporadically isolated from wild and domestic birds, H2N2 viruses have not been detected among human populations since 1968. Should H2N2 viruses adapt to domestic poultry they may pose a risk of infection to people, as most anyone born after 1968 would likely be susceptible to their infection. We report the isolation of a novel influenza A virus (H2N2) cultured in 2013 from a healthy domestic duck at a live poultry market in Wuxi City, China. Sequence data revealed that the novel H2N2 virus was similar to Eurasian avian lineage avian influenza viruses, the virus had been circulating for ≥ two years among poultry, had an increase in α2,6 binding affinity, and was not highly pathogenic. Approximately 9% of 100 healthy chickens sampled from the same area had elevated antibodies against the H2 antigen. Fortunately, there was sparse serological evidence that the virus was infecting poultry workers or had adapted to infect other mammals. These findings suggest that a novel H2N2 virus has been circulating among domestic poultry in Wuxi City, China and has some has increased human receptor affinity. It seems wise to conduct better surveillance for novel influenza viruses at Chinese live bird markets.</p>
</div>
</front>
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<pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Sci Rep</journal-id>
<journal-id journal-id-type="iso-abbrev">Sci Rep</journal-id>
<journal-title-group><journal-title>Scientific Reports</journal-title>
</journal-title-group>
<issn pub-type="epub">2045-2322</issn>
<publisher><publisher-name>Nature Publishing Group</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">25533850</article-id>
<article-id pub-id-type="pmc">4274511</article-id>
<article-id pub-id-type="pii">srep07588</article-id>
<article-id pub-id-type="doi">10.1038/srep07588</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Article</subject>
</subj-group>
</article-categories>
<title-group><article-title>Characterization of a Novel Reassortant Influenza A Virus (H2N2) from a Domestic Duck in Eastern China</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Ma</surname>
<given-names>Mai-Juan</given-names>
</name>
<xref ref-type="corresp" rid="c2">b</xref>
<xref ref-type="aff" rid="a1">1</xref>
<xref ref-type="author-notes" rid="n1">*</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Yang</surname>
<given-names>Xiao-Xian</given-names>
</name>
<xref ref-type="aff" rid="a1">1</xref>
<xref ref-type="aff" rid="a2">2</xref>
<xref ref-type="author-notes" rid="n1">*</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Qian</surname>
<given-names>Yan-Hua</given-names>
</name>
<xref ref-type="aff" rid="a3">3</xref>
<xref ref-type="author-notes" rid="n1">*</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zhao</surname>
<given-names>Si-Yan</given-names>
</name>
<xref ref-type="aff" rid="a4">4</xref>
<xref ref-type="author-notes" rid="n1">*</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Hua</surname>
<given-names>Sha</given-names>
</name>
<xref ref-type="aff" rid="a5">5</xref>
<xref ref-type="aff" rid="a6">6</xref>
<xref ref-type="author-notes" rid="n1">*</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wang</surname>
<given-names>Tie-Cheng</given-names>
</name>
<xref ref-type="aff" rid="a4">4</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Chen</surname>
<given-names>Shan-Hui</given-names>
</name>
<xref ref-type="aff" rid="a3">3</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Ma</surname>
<given-names>Guang-Yuan</given-names>
</name>
<xref ref-type="aff" rid="a3">3</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Sang</surname>
<given-names>Xiao-Yu</given-names>
</name>
<xref ref-type="aff" rid="a7">7</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Liu</surname>
<given-names>Lin-Na</given-names>
</name>
<xref ref-type="aff" rid="a4">4</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wu</surname>
<given-names>Ai-Ping</given-names>
</name>
<xref ref-type="aff" rid="a5">5</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Jiang</surname>
<given-names>Tai-Jiao</given-names>
</name>
<xref ref-type="aff" rid="a5">5</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Gao</surname>
<given-names>Yu-Wei</given-names>
</name>
<xref ref-type="aff" rid="a4">4</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Gray</surname>
<given-names>Gregory C.</given-names>
</name>
<xref ref-type="aff" rid="a8">8</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zhao</surname>
<given-names>Teng</given-names>
</name>
<xref ref-type="aff" rid="a1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Ling</surname>
<given-names>Xia</given-names>
</name>
<xref ref-type="aff" rid="a3">3</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Wang</surname>
<given-names>Jing-Lin</given-names>
</name>
<xref ref-type="aff" rid="a1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Lu</surname>
<given-names>Bing</given-names>
</name>
<xref ref-type="aff" rid="a3">3</xref>
<xref ref-type="author-notes" rid="n2">†</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Qian</surname>
<given-names>Jun</given-names>
</name>
<xref ref-type="aff" rid="a4">4</xref>
<xref ref-type="author-notes" rid="n2">†</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Cao</surname>
<given-names>Wu-Chun</given-names>
</name>
<xref ref-type="corresp" rid="c1">a</xref>
<xref ref-type="aff" rid="a1">1</xref>
<xref ref-type="aff" rid="a2">2</xref>
<xref ref-type="author-notes" rid="n2">†</xref>
</contrib>
<aff id="a1"><label>1</label>
<institution>State Key Laboratory of Pathogen and Biosecurity, Beijing Institute of Microbiology and Epidemiology</institution>
, Beijing, 100071, P. R. China</aff>
<aff id="a2"><label>2</label>
<institution>School of Public Health, Central South University</institution>
, Changsha, Hunan, 410000 P. R. China</aff>
<aff id="a3"><label>3</label>
<institution>Wuxi Center for Disease Control and Prevention</institution>
, Wuxi, 214023, P. R. China</aff>
<aff id="a4"><label>4</label>
<institution>Military Veterinary Research Institute, Academy of Military Medical Sciences</institution>
, Changchun, 130122, P. R. China</aff>
<aff id="a5"><label>5</label>
<institution>Key Laboratory of Protein and Peptide Pharmaceutical, National Laboratory of Biomacromolecules, Institute of Biophysics</institution>
, Chinese Academy of Sciences, Beijing, 100101,<country>P. R. China</country>
</aff>
<aff id="a6"><label>6</label>
<institution>University of Chinese Academy of Sciences</institution>
, Beijing, 100049,<country>China</country>
</aff>
<aff id="a7"><label>7</label>
<institution>Harbin Veterinary Research Institute, Chinese Academy of Agricultural Sciences</institution>
, Harbin, 150001,<country>China</country>
</aff>
<aff id="a8"><label>8</label>
<institution>Duke Infectious Disease & Duke Health Institute, Duke University Medical Center</institution>
, Durham, NC, 27710,<country>USA</country>
</aff>
</contrib-group>
<author-notes><corresp id="c1"><label>a</label>
<email>caowc@bmi.ac.cn</email>
</corresp>
<corresp id="c2"><label>b</label>
<email>mjma@163.com</email>
</corresp>
<fn id="n1"><label>*</label>
<p>These authors contributed equally to this work.</p>
</fn>
<fn id="n2"><label>†</label>
<p>These authors jointly supervised this work.</p>
</fn>
</author-notes>
<pub-date pub-type="epub"><day>23</day>
<month>12</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="collection"><year>2014</year>
</pub-date>
<volume>4</volume>
<elocation-id>7588</elocation-id>
<history><date date-type="received"><day>08</day>
<month>08</month>
<year>2014</year>
</date>
<date date-type="accepted"><day>28</day>
<month>11</month>
<year>2014</year>
</date>
</history>
<permissions><copyright-statement>Copyright © 2014, Macmillan Publishers Limited. All rights reserved</copyright-statement>
<copyright-year>2014</copyright-year>
<copyright-holder>Macmillan Publishers Limited. All rights reserved</copyright-holder>
<license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by-nc-nd/4.0/"><pmc-comment>author-paid</pmc-comment>
<license-p>This work is licensed under a Creative Commons Attribution-NonCommercial-NoDerivs 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 in order 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-nc-nd/4.0/">http://creativecommons.org/licenses/by-nc-nd/4.0/</ext-link>
</license-p>
</license>
</permissions>
<abstract><p>While H2N2 viruses have been sporadically isolated from wild and domestic birds, H2N2 viruses have not been detected among human populations since 1968. Should H2N2 viruses adapt to domestic poultry they may pose a risk of infection to people, as most anyone born after 1968 would likely be susceptible to their infection. We report the isolation of a novel influenza A virus (H2N2) cultured in 2013 from a healthy domestic duck at a live poultry market in Wuxi City, China. Sequence data revealed that the novel H2N2 virus was similar to Eurasian avian lineage avian influenza viruses, the virus had been circulating for ≥ two years among poultry, had an increase in α2,6 binding affinity, and was not highly pathogenic. Approximately 9% of 100 healthy chickens sampled from the same area had elevated antibodies against the H2 antigen. Fortunately, there was sparse serological evidence that the virus was infecting poultry workers or had adapted to infect other mammals. These findings suggest that a novel H2N2 virus has been circulating among domestic poultry in Wuxi City, China and has some has increased human receptor affinity. It seems wise to conduct better surveillance for novel influenza viruses at Chinese live bird markets.</p>
</abstract>
</article-meta>
</front>
<floats-group><fig id="f1"><label>Figure 1</label>
<caption><title>Phylogenetic relationships of each gene of novel H2N2 strain and closely related viruses.</title>
<p>All branch lengths are drawn to a scale of nucleotide substitutions per site. The novel H2N2 virus is highlighted in red. Detailed phylogenetic trees with virus designations can be found in <xref ref-type="supplementary-material" rid="s1">Supplemental Figure 1</xref>
.</p>
</caption>
<graphic xlink:href="srep07588-f1"></graphic>
</fig>
<fig id="f2"><label>Figure 2</label>
<caption><title>Temporal reconstruction of the phylogenetic relationships of each gene segment.</title>
<p>The novel H2N2 strain is highlighted by red. Internal nodes are reconstructed common ancestors with 95% highest posterior density (HPD) intervals on their dates given by the light red bars. Detailed temporal reconstruction of phylogenetic trees with virus designations can be found in <xref ref-type="supplementary-material" rid="s1">Supplemental Figure 1</xref>
.</p>
</caption>
<graphic xlink:href="srep07588-f2"></graphic>
</fig>
<fig id="f3"><label>Figure 3</label>
<caption><title>Reassortment model for the origins of the novel H2N2.</title>
<p>The colors of the novel H2N2 gene segments indicate their origins. The picture of duck was drawn by the author Mai-Juan Ma.</p>
</caption>
<graphic xlink:href="srep07588-f3"></graphic>
</fig>
<fig id="f4"><label>Figure 4</label>
<caption><title>Characterization of the receptor-binding properties of isolated H2N2 virus.</title>
<p>Binding of virus to a2,3-linked (3′SL-PAA) or a2,6-linked (6′SL-PAA) sialylglycan receptors was determined by solid-phase binding assays. (a) DK13-H2N2 (A/duck/Wuxi/2/13 (H2N2)); (b) XD09-H1N1 (A/Changchun/XD/2009 (H1N1)); (c) CK06-H9N2 (A/Chicken/Jilin/Hu-3/2006(H9N2)). Blue line, binding to (3′SL-PAA); red line, binding to (6′SL-PAA).</p>
</caption>
<graphic xlink:href="srep07588-f4"></graphic>
</fig>
<fig id="f5"><label>Figure 5</label>
<caption><title>Hemagglutination inhibition (HI) Antibody against H2N2 virus among human population and live poultry (mean antibody counts shown with 95% confidence intervals).</title>
<p>(a) HI antibody of H2N2 virus among poultry. (b) HI antibody of H2N2 virus among general health population, poultry workers, and swine workers. (c) HI antibody of H2N2 virus among different age population.</p>
</caption>
<graphic xlink:href="srep07588-f5"></graphic>
</fig>
<table-wrap position="float" id="t1"><label>Table 1</label>
<caption><title>Time of most recent common ancestors (tMRCA) for the novel H2N2 virus. Observed closest strains for the H2N2 segments, time of divergence, and the mean evolutionary rate are listed</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="justify" valign="top" charoff="50">Gene</th>
<th align="center" valign="top" charoff="50">Divergence time</th>
<th align="center" valign="top" charoff="50">Observed closest strain</th>
<th align="center" valign="top" charoff="50">Mean evolutionary rate (10<sup>−3</sup>
)</th>
</tr>
</thead>
<tbody valign="top"><tr><td align="center" valign="top" charoff="50">PB2</td>
<td align="char" valign="top" char=" " charoff="50">May 2010 (Dec 2009, Oct 2010)</td>
<td align="char" valign="top" char="/" charoff="50">A/mallard/Korea/1203/2010_H10N8</td>
<td align="center" valign="top" charoff="50">2.133 (1.849, 2.421)</td>
</tr>
<tr><td align="center" valign="top" charoff="50">PB1</td>
<td align="char" valign="top" char=" " charoff="50">Jul 2010 (Sep 2009, Mar 2011)</td>
<td align="char" valign="top" char="/" charoff="50">A/duck/Hunan/S11200/2012_H4N6</td>
<td align="center" valign="top" charoff="50">2.591 (2.344, 2.846)</td>
</tr>
<tr><td align="center" valign="top" charoff="50">PA</td>
<td align="char" valign="top" char=" " charoff="50">Jun 2010 (Apr 2009, Jul 2011)</td>
<td align="char" valign="top" char="/" charoff="50">A/duck/Vietnam/LBM185/2012_H6N6</td>
<td align="center" valign="top" charoff="50">1.864 (1.405, 2.351)</td>
</tr>
<tr><td align="center" valign="top" charoff="50">HA</td>
<td align="char" valign="top" char=" " charoff="50">May 2006 (Jan 2005, Aug 2007)</td>
<td align="char" valign="top" char="/" charoff="50">A/environment/Guangdong/2/2009_H2N3</td>
<td align="center" valign="top" charoff="50">3.859 (3.462, 4.278)</td>
</tr>
<tr><td align="center" valign="top" charoff="50">NP</td>
<td align="char" valign="top" char=" " charoff="50">Jun 2010 (Oct 2008, Sep 2011)</td>
<td align="char" valign="top" char="/" charoff="50">A/duck/Vietnam/LBM81/2012_H11N9</td>
<td align="center" valign="top" charoff="50">1.562 (1.305, 1.816)</td>
</tr>
<tr><td align="center" valign="top" charoff="50">NA</td>
<td align="char" valign="top" char=" " charoff="50">Jun 2011 (Jul 2010, Jan 2012)</td>
<td align="char" valign="top" char="/" charoff="50">A/duck/Jiangsu/m234/2012_H5N2</td>
<td align="center" valign="top" charoff="50">3.049 (2.697, 3.398)</td>
</tr>
<tr><td align="center" valign="top" charoff="50">M</td>
<td align="char" valign="top" char=" " charoff="50">Oct 2003 (May 2002, Sep 2004)</td>
<td align="char" valign="top" char="/" charoff="50">A/duck/Eastern-China/29/2004_H6N2</td>
<td align="center" valign="top" charoff="50">1.696 (1.400, 2.011)</td>
</tr>
<tr><td align="center" valign="top" charoff="50">NS</td>
<td align="char" valign="top" char=" " charoff="50">Sep 2009 (Nov 2008, Jun 2010)</td>
<td align="char" valign="top" char="/" charoff="50">A/duck/Nanjing/1102/2010_H4N8</td>
<td align="center" valign="top" charoff="50">2.357 (1.977, 2.773)</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="t1-fn1"><p>The mean evolutionary rate is represented as the number of substitutions per site per year. The values in parentheses represent the 95% highest posterior density (HPD) intervals.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap position="float" id="t2"><label>Table 2</label>
<caption><title>Selected characteristic amino acids of the novel influenza A(H2N2) 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="justify" valign="top" charoff="50">Viral protein</th>
<th align="center" valign="top" charoff="50">Amino acid position</th>
<th align="center" valign="top" charoff="50">A/duck/WX/2/13</th>
<th align="center" valign="top" charoff="50">Comments</th>
</tr>
</thead>
<tbody valign="top"><tr><td align="center" valign="top" charoff="50">PB2</td>
<td align="char" valign="top" char="." charoff="50">627</td>
<td align="center" valign="top" charoff="50">E</td>
<td align="left" valign="top" charoff="50">E627K: Mammalian host adaptation<xref ref-type="bibr" rid="b42">42</xref>
<xref ref-type="bibr" rid="b43">43</xref>
</td>
</tr>
<tr><td align="center" valign="top" charoff="50">HA</td>
<td align="char" valign="top" char="." charoff="50">138</td>
<td align="center" valign="top" charoff="50">A</td>
<td align="left" valign="top" charoff="50">S138A: Increased virus binding to human-type receptors<xref ref-type="bibr" rid="b44">44</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">186</td>
<td align="center" valign="top" charoff="50">N</td>
<td align="left" valign="top" charoff="50">G186V: Increased virus binding to human-type receptors<xref ref-type="bibr" rid="b45">45</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">190</td>
<td align="center" valign="top" charoff="50">E</td>
<td align="left" valign="top" charoff="50">E190D: Increased virus binding to human-type receptors<xref ref-type="bibr" rid="b46">46</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">225</td>
<td align="center" valign="top" charoff="50">G</td>
<td align="left" valign="top" charoff="50">G225D: Increased virus binding to human-type receptors<xref ref-type="bibr" rid="b46">46</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">226</td>
<td align="center" valign="top" charoff="50">Q</td>
<td align="left" valign="top" charoff="50">Q226L: Increased virus binding to human-type receptors<xref ref-type="bibr" rid="b26">26</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">228</td>
<td align="center" valign="top" charoff="50">G</td>
<td align="left" valign="top" charoff="50">H2N2 Human: S<xref ref-type="bibr" rid="b47">47</xref>
</td>
</tr>
<tr><td align="center" valign="top" charoff="50">NA</td>
<td align="center" valign="top" charoff="50">65-71</td>
<td align="center" valign="top" charoff="50">No deletion</td>
<td align="left" valign="top" charoff="50">Deletion of 65–71 increased virulence in mice<xref ref-type="bibr" rid="b48">48</xref>
</td>
</tr>
<tr><td align="center" valign="top" charoff="50">M1</td>
<td align="char" valign="top" char="." charoff="50">30</td>
<td align="center" valign="top" charoff="50">D</td>
<td align="left" valign="top" charoff="50">N30D: Increased virulence in mice (most influenza A viruses encode 30D)<xref ref-type="bibr" rid="b49">49</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">174</td>
<td align="center" valign="top" charoff="50">R</td>
<td align="left" valign="top" charoff="50">K174K(R): Host marker of influenza A<xref ref-type="bibr" rid="b44">44</xref>
</td>
</tr>
<tr><td align="center" valign="top" charoff="50">NS1</td>
<td align="char" valign="top" char="." charoff="50">42</td>
<td align="center" valign="top" charoff="50">S</td>
<td align="left" valign="top" charoff="50">P42S: Increased virulence in mice (most avian influenza A viruses encode 42S)<xref ref-type="bibr" rid="b50">50</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">97</td>
<td align="center" valign="top" charoff="50">E</td>
<td align="left" valign="top" charoff="50">D97E: Increased avian H5N1 viruses' virulence in mammals<xref ref-type="bibr" rid="b47">47</xref>
</td>
</tr>
<tr><td align="left" valign="top" charoff="50"> </td>
<td align="char" valign="top" char="." charoff="50">127</td>
<td align="center" valign="top" charoff="50">N</td>
<td align="left" valign="top" charoff="50">TDRVA127N: Association with H3N2 virulence<xref ref-type="bibr" rid="b47">47</xref>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="t2-fn1"><p>HA amino acid residues have H3 numbering; NA amino acid residues have N2 numbering.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</floats-group>
</pmc>
</record>
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