The Neuraminidase Stalk Deletion Serves as Major Virulence Determinant of H5N1 Highly Pathogenic Avian Influenza Viruses in Chicken
Identifieur interne : 000100 ( Pmc/Curation ); précédent : 000099; suivant : 000101The Neuraminidase Stalk Deletion Serves as Major Virulence Determinant of H5N1 Highly Pathogenic Avian Influenza Viruses in Chicken
Auteurs : Olga Stech [Allemagne] ; Jutta Veits [Allemagne] ; El-Sayed M. Abdelwhab [Allemagne] ; Ute Wessels [Allemagne] ; Thomas C. Mettenleiter [Allemagne] ; Jürgen Stech [Allemagne]Source :
- Scientific Reports [ 2045-2322 ] ; 2015.
Abstract
Highly pathogenic avian influenza viruses (HPAIV) cause devastating losses in gallinaceous poultry world-wide and raised concerns of a novel pandemic. HPAIV develop from low-pathogenic precursors by acquisition of a polybasic HA cleavage site (HACS), the prime virulence determinant. Beside that HACS, other adaptive changes accumulate in those precursors prior to transformation into an HPAIV. Here, we aimed to unravel such virulence determinants in addition to the HA gene. Stepwise reduction of HPAIV genes revealed that the HPAIV HA and NA form a minimum set of virulence determinants, sufficient for a lethal phenotype in chicken. Abolishing the NA stalk deletion considerably reduced lethality and prevented transmission. Conversely, the analogous stalk deletion reconstructed in the NA of an LPAIV reassortant carrying only the HPAIV HA resulted in 100% lethality both after primary and contact infection. Remarkably, the unmodified LPAIV NA with its long stalk, when exclusively introduced into the H5N1 HPAIV, still enabled high virulence and efficient transmission. Therefore, irrespective of an NA stalk deletion, minor virulence determinants in addition to the essential polybasic HACS contribute to high virulence, whereas the NA stalk deletion alone may serve as major virulence determinant.
Url:
DOI: 10.1038/srep13493
PubMed: 26306544
PubMed Central: 4549673
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">The Neuraminidase Stalk Deletion Serves as Major Virulence Determinant of H5N1 Highly Pathogenic Avian Influenza Viruses in Chicken</title><author><name sortKey="Stech, Olga" sort="Stech, Olga" uniqKey="Stech O" first="Olga" last="Stech">Olga Stech</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Veits, Jutta" sort="Veits, Jutta" uniqKey="Veits J" first="Jutta" last="Veits">Jutta Veits</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Abdelwhab, El Sayed M" sort="Abdelwhab, El Sayed M" uniqKey="Abdelwhab E" first="El-Sayed M." last="Abdelwhab">El-Sayed M. Abdelwhab</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Wessels, Ute" sort="Wessels, Ute" uniqKey="Wessels U" first="Ute" last="Wessels">Ute Wessels</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Mettenleiter, Thomas C" sort="Mettenleiter, Thomas C" uniqKey="Mettenleiter T" first="Thomas C." last="Mettenleiter">Thomas C. Mettenleiter</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Stech, Jurgen" sort="Stech, Jurgen" uniqKey="Stech J" first="Jürgen" last="Stech">Jürgen Stech</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">26306544</idno><idno type="pmc">4549673</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4549673</idno><idno type="RBID">PMC:4549673</idno><idno type="doi">10.1038/srep13493</idno><date when="2015">2015</date><idno type="wicri:Area/Pmc/Corpus">000100</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000100</idno><idno type="wicri:Area/Pmc/Curation">000100</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Curation">000100</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">The Neuraminidase Stalk Deletion Serves as Major Virulence Determinant of H5N1 Highly Pathogenic Avian Influenza Viruses in Chicken</title><author><name sortKey="Stech, Olga" sort="Stech, Olga" uniqKey="Stech O" first="Olga" last="Stech">Olga Stech</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Veits, Jutta" sort="Veits, Jutta" uniqKey="Veits J" first="Jutta" last="Veits">Jutta Veits</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Abdelwhab, El Sayed M" sort="Abdelwhab, El Sayed M" uniqKey="Abdelwhab E" first="El-Sayed M." last="Abdelwhab">El-Sayed M. Abdelwhab</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Wessels, Ute" sort="Wessels, Ute" uniqKey="Wessels U" first="Ute" last="Wessels">Ute Wessels</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Mettenleiter, Thomas C" sort="Mettenleiter, Thomas C" uniqKey="Mettenleiter T" first="Thomas C." last="Mettenleiter">Thomas C. Mettenleiter</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Stech, Jurgen" sort="Stech, Jurgen" uniqKey="Stech J" first="Jürgen" last="Stech">Jürgen Stech</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></nlm:aff><country xml:lang="fr">Allemagne</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author></analytic><series><title level="j">Scientific Reports</title><idno type="eISSN">2045-2322</idno><imprint><date when="2015">2015</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Highly pathogenic avian influenza viruses (HPAIV) cause devastating losses in gallinaceous poultry world-wide and raised concerns of a novel pandemic. HPAIV develop from low-pathogenic precursors by acquisition of a polybasic HA cleavage site (HACS), the prime virulence determinant. Beside that HACS, other adaptive changes accumulate in those precursors prior to transformation into an HPAIV. Here, we aimed to unravel such virulence determinants in addition to the HA gene. Stepwise reduction of HPAIV genes revealed that the HPAIV HA and NA form a minimum set of virulence determinants, sufficient for a lethal phenotype in chicken. Abolishing the NA stalk deletion considerably reduced lethality and prevented transmission. Conversely, the analogous stalk deletion reconstructed in the NA of an LPAIV reassortant carrying only the HPAIV HA resulted in 100% lethality both after primary and contact infection. Remarkably, the unmodified LPAIV NA with its long stalk, when exclusively introduced into the H5N1 HPAIV, still enabled high virulence and efficient transmission. Therefore, irrespective of an NA stalk deletion, minor virulence determinants in addition to the essential polybasic HACS contribute to high virulence, whereas the NA stalk deletion alone may serve as major virulence determinant.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Neumann, G" uniqKey="Neumann G">G. Neumann</name></author><author><name sortKey="Chen, H" uniqKey="Chen H">H. Chen</name></author><author><name sortKey="Gao, G F" uniqKey="Gao G">G. F. Gao</name></author><author><name sortKey="Shu, Y" uniqKey="Shu Y">Y. Shu</name></author><author><name sortKey="Kawaoka, Y" uniqKey="Kawaoka Y">Y. Kawaoka</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Capua, I" uniqKey="Capua I">I. Capua</name></author><author><name sortKey="Marangon, S" uniqKey="Marangon S">S. 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<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">26306544</article-id><article-id pub-id-type="pmc">4549673</article-id><article-id pub-id-type="pii">srep13493</article-id><article-id pub-id-type="doi">10.1038/srep13493</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>The Neuraminidase Stalk Deletion Serves as Major Virulence Determinant of H5N1 Highly Pathogenic Avian Influenza Viruses in Chicken</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Stech</surname><given-names>Olga</given-names></name><xref ref-type="aff" rid="a1">1</xref><xref ref-type="author-notes" rid="n1">*</xref></contrib><contrib contrib-type="author"><name><surname>Veits</surname><given-names>Jutta</given-names></name><xref ref-type="aff" rid="a1">1</xref><xref ref-type="author-notes" rid="n1">*</xref></contrib><contrib contrib-type="author"><name><surname>Abdelwhab</surname><given-names>El-Sayed M.</given-names></name><xref ref-type="aff" rid="a1">1</xref><xref ref-type="author-notes" rid="n1">*</xref></contrib><contrib contrib-type="author"><name><surname>Wessels</surname><given-names>Ute</given-names></name><xref ref-type="aff" rid="a1">1</xref></contrib><contrib contrib-type="author"><name><surname>Mettenleiter</surname><given-names>Thomas C.</given-names></name><xref ref-type="aff" rid="a1">1</xref></contrib><contrib contrib-type="author"><name><surname>Stech</surname><given-names>Jürgen</given-names></name><xref ref-type="corresp" rid="c1">a</xref><xref ref-type="aff" rid="a1">1</xref></contrib><aff id="a1"><label>1</label><institution>Institute of Molecular Virology and Cell Biology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health</institution>, Südufer 10, 17493 Greifswald-Insel Riems,<country>Germany</country></aff></contrib-group><author-notes><corresp id="c1"><label>a</label><email>juergen.stech@fli.bund.de</email></corresp><fn id="n1"><label>*</label><p>These authors contributed equally to this work.</p></fn></author-notes><pub-date pub-type="epub"><day>26</day><month>08</month><year>2015</year></pub-date><pub-date pub-type="collection"><year>2015</year></pub-date><volume>5</volume><elocation-id>13493</elocation-id><history><date date-type="received"><day>03</day><month>02</month><year>2015</year></date><date date-type="accepted"><day>22</day><month>07</month><year>2015</year></date></history><permissions><copyright-statement>Copyright © 2015, Macmillan Publishers Limited</copyright-statement><copyright-year>2015</copyright-year><copyright-holder>Macmillan Publishers Limited</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>Highly pathogenic avian influenza viruses (HPAIV) cause devastating losses in gallinaceous poultry world-wide and raised concerns of a novel pandemic. HPAIV develop from low-pathogenic precursors by acquisition of a polybasic HA cleavage site (HACS), the prime virulence determinant. Beside that HACS, other adaptive changes accumulate in those precursors prior to transformation into an HPAIV. Here, we aimed to unravel such virulence determinants in addition to the HA gene. Stepwise reduction of HPAIV genes revealed that the HPAIV HA and NA form a minimum set of virulence determinants, sufficient for a lethal phenotype in chicken. Abolishing the NA stalk deletion considerably reduced lethality and prevented transmission. Conversely, the analogous stalk deletion reconstructed in the NA of an LPAIV reassortant carrying only the HPAIV HA resulted in 100% lethality both after primary and contact infection. Remarkably, the unmodified LPAIV NA with its long stalk, when exclusively introduced into the H5N1 HPAIV, still enabled high virulence and efficient transmission. Therefore, irrespective of an NA stalk deletion, minor virulence determinants in addition to the essential polybasic HACS contribute to high virulence, whereas the NA stalk deletion alone may serve as major virulence determinant.</p></abstract></article-meta></front><floats-group><fig id="f1"><label>Figure 1</label><caption><title>Simultaneous presence of TG05 and R65 genes determined by Sanger sequencing of supernatants following plasmid co-transfection to obtain mixtures of random reassortants.</title><p>(<bold>A</bold>) The TG05 polymerase gene plasmids (PB2, PB1, PA, NP) and the R65 HA plasmid were co-transfected together with mixtures of the NA, M, and NS plasmids originating both from TG05 and R65. (<bold>B</bold>) The TG05 plasmids except the HA plasmid were co-transfected together with all eight R65 plasmids.</p></caption><graphic xlink:href="srep13493-f1"></graphic></fig><fig id="f2"><label>Figure 2</label><caption><title>High lethality following oculonasal infection of chickens with 600 μl co-transfection supernatants.</title><p>Daily clinical score: 0: healthy, 1: ill, 2: severely ill or 3: dead. (<bold>A</bold>) Birds infected with supernatant from co-transfection of the TG05 polymerase gene plasmids (PB2, PB1, PA, NP), the R65 HA plasmid and mixtures of the NA, M, and NS plasmids originating both from TG05 and R65. (<bold>B</bold>) Birds infected with supernatant from co-transfection of the TG05 plasmids except the HA plasmid and all eight R65 plasmids.</p></caption><graphic xlink:href="srep13493-f2"></graphic></fig><fig id="f3"><label>Figure 3</label><caption><title>Minimal R65 gene constellations sufficient for a lethal phenotype and transmission in chicken.</title><p>We infected the birds oculonasally with 10<sup>5</sup> pfu virus and added bystander animals on day 1 p. i. Daily clinical score: 0: healthy, 1: ill, 2: severely ill or 3: dead.</p></caption><graphic xlink:href="srep13493-f3"></graphic></fig><fig id="f4"><label>Figure 4</label><caption><title>NA stalk deletion is crucial for transmission among chickens.</title><p>We infected chickens oculonasally with 10<sup>5</sup> pfu virus and added bystander animals on day 1 p. i. Daily clinical score: 0: healthy, 1: ill, 2: severely ill or 3: dead.</p></caption><graphic xlink:href="srep13493-f4"></graphic></fig><table-wrap position="float" id="t1"><label>Table 1</label><caption><title>Virulence of parent viruses, TG05/R65 reassortants, and NA stalk variants in chicken.</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 align="center" valign="top" charoff="50">Morbidity</th><th align="center" valign="top" charoff="50">Mortality</th><th align="center" valign="top" charoff="50">MDT (d) and Range</th><th align="center" valign="top" charoff="50">Clinical Score</th></tr></thead><tbody valign="top"><tr><td colspan="5" align="left" valign="top" charoff="50"><italic>Parent Viruses</italic></td></tr><tr><td align="left" valign="top" charoff="50">TG05<xref ref-type="fn" rid="t1-fn1">1</xref></td><td align="center" valign="top" charoff="50">0/10</td><td align="center" valign="top" charoff="50">0/10</td><td align="center" valign="top" charoff="50">n/a</td><td align="center" valign="top" charoff="50">0.00</td></tr><tr><td align="left" valign="top" charoff="50">TG05-HA<sub>R65</sub><xref ref-type="fn" rid="t1-fn1">1</xref></td><td align="center" valign="top" charoff="50">8/10</td><td align="center" valign="top" charoff="50">3/10</td><td align="center" valign="top" charoff="50">n/a</td><td align="center" valign="top" charoff="50">0.90</td></tr><tr><td align="left" valign="top" charoff="50">R65<xref ref-type="fn" rid="t1-fn2">2</xref></td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">3.8 (3–5)</td><td align="center" valign="top" charoff="50">2.20</td></tr><tr><td colspan="5" align="left" valign="top" charoff="50"><italic>Selected Reassortant</italic></td></tr><tr><td align="left" valign="top" charoff="50">TG05-HA/PB2/NP/M/NA<sub>R65</sub><xref ref-type="fn" rid="t1-fn2">2</xref></td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">3.5 (3–4)</td><td align="center" valign="top" charoff="50">2.26</td></tr><tr><td colspan="5" align="left" valign="top" charoff="50"><italic>Tailored Reassortants</italic></td></tr><tr><td align="left" valign="top" charoff="50">TG05-HA/PB2/PB1/PA/NP<sub>R65</sub></td><td align="center" valign="top" charoff="50">4/8</td><td align="center" valign="top" charoff="50">3/8</td><td align="center" valign="top" charoff="50">7.3 (6–8)</td><td align="center" valign="top" charoff="50">0.62</td></tr><tr><td align="left" valign="top" charoff="50">TG05/HA/NA/M/NS<sub>R65</sub></td><td align="center" valign="top" charoff="50">10/10</td><td align="center" valign="top" charoff="50">10/10</td><td align="center" valign="top" charoff="50">2.9 (2–3)</td><td align="center" valign="top" charoff="50">2.54</td></tr><tr><td align="left" valign="top" charoff="50">TG05/HA/NA/M<sub>R65</sub></td><td align="center" valign="top" charoff="50">10/10</td><td align="center" valign="top" charoff="50">10/10</td><td align="center" valign="top" charoff="50">3.6 (3–4)</td><td align="center" valign="top" charoff="50">2.43</td></tr><tr><td align="left" valign="top" charoff="50">TG05/HA/NA/NS<sub>R65</sub></td><td align="center" valign="top" charoff="50">10/10</td><td align="center" valign="top" charoff="50">10/10</td><td align="center" valign="top" charoff="50">5.2 (4–6)</td><td align="center" valign="top" charoff="50">1.80</td></tr><tr><td align="left" valign="top" charoff="50">TG05/HA<sub>R65</sub>NA<sub>R65</sub></td><td align="center" valign="top" charoff="50">8/8</td><td align="center" valign="top" charoff="50">8/8</td><td align="center" valign="top" charoff="50">5.9 (5–7)</td><td align="center" valign="top" charoff="50">1.67</td></tr><tr><td align="left" valign="top" charoff="50">TG05/HA/M/NS<sub>R65</sub></td><td align="center" valign="top" charoff="50">4/8</td><td align="center" valign="top" charoff="50">3/8</td><td align="center" valign="top" charoff="50">7.0 (6–9)</td><td align="center" valign="top" charoff="50">0.51</td></tr><tr><td align="left" valign="top" charoff="50">R65-NA<sub>TG05</sub><xref ref-type="fn" rid="t1-fn2">2</xref></td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">6.2 (4–8)</td><td align="center" valign="top" charoff="50">1.73</td></tr><tr><td colspan="5" align="left" valign="top" charoff="50"><italic>NA Stalk Variants</italic></td></tr><tr><td align="left" valign="top" charoff="50">TG05-HA<sub>R65</sub>NA<sub>long-R65</sub><xref ref-type="fn" rid="t1-fn3">3</xref></td><td align="center" valign="top" charoff="50">5/6</td><td align="center" valign="top" charoff="50">4/6</td><td align="center" valign="top" charoff="50">5.5 (5–6)</td><td align="center" valign="top" charoff="50">1.32</td></tr><tr><td align="left" valign="top" charoff="50">TG05-HA<sub>R65</sub>NA<sub>short-TG05</sub><xref ref-type="fn" rid="t1-fn3">3</xref></td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">6/6</td><td align="center" valign="top" charoff="50">4.2 (3–5)</td><td align="center" valign="top" charoff="50">2.22</td></tr></tbody></table><table-wrap-foot><fn id="t1-fn1"><p>Morbidity, mortality and overall clinical score after oculonasal infection with 10<sup>5</sup> pfu.</p></fn><fn id="t1-fn2"><p><sup>1</sup>Data of this group from previous study<xref ref-type="bibr" rid="b12">12</xref>.</p></fn><fn id="t1-fn3"><p><sup>2</sup>Identical group as in <xref ref-type="fig" rid="f3">Fig. 3</xref>.</p></fn><fn id="t1-fn4"><p><sup>3</sup>Identical group as in <xref ref-type="fig" rid="f4">Fig. 4</xref>.</p></fn></table-wrap-foot></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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