Influenza Virus Specific CD8+ T Cells Exacerbate Infection Following High Dose Influenza Challenge of Aged Mice
Identifieur interne : 000978 ( Ncbi/Merge ); précédent : 000977; suivant : 000979Influenza Virus Specific CD8+ T Cells Exacerbate Infection Following High Dose Influenza Challenge of Aged Mice
Auteurs : E. M. Parzych [États-Unis] ; L. J. Dimenna [États-Unis] ; B. P. Latimer [États-Unis] ; J. C. Small [États-Unis] ; S. Kannan [États-Unis] ; B. Manson [États-Unis] ; M. O. Lasaro [États-Unis] ; E. J. Wherry [États-Unis] ; H. C. Ertl [États-Unis]Source :
- BioMed Research International [ 2314-6133 ] ; 2013.
Abstract
Influenza viruses cause severe illnesses and death, mainly in the aged population. Protection afforded by licensed vaccines through subtype-specific neutralizing antibodies is incomplete, especially when the vaccine antigens fail to closely match those of the circulating viral strains. Efforts are underway to generate a so-called universal influenza vaccine expressing conserved viral sequences that induce broad protection to multiple strains of influenza virus through the induction of CD8+ T cells. Here we assess the effect of a potent antiviral CD8+ T cell response on influenza virus infection of young and aged mice. Our results show that CD8+ T cell-inducing vaccines can provide some protection to young mice, but they exacerbate influenza virus-associated disease in aged mice, causing extensive lung pathology and death.
Url:
DOI: 10.1155/2013/876314
PubMed: 24187666
PubMed Central: 3800650
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Manson</name><affiliation wicri:level="2"><nlm:aff id="I1">Wistar Institute, Philadelphia, PA 19104, USA</nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea>Wistar Institute, Philadelphia, PA 19104</wicri:regionArea><placeName><region type="state">Pennsylvanie</region></placeName></affiliation><affiliation wicri:level="2"><nlm:aff id="I5">Cheyney University, Thornbury Township, PA 19139, USA</nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea>Cheyney University, Thornbury Township, PA 19139</wicri:regionArea><placeName><region type="state">Pennsylvanie</region></placeName></affiliation></author><author><name sortKey="Lasaro, M O" sort="Lasaro, M O" uniqKey="Lasaro M" first="M. O." last="Lasaro">M. O. 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Wherry</name><affiliation wicri:level="2"><nlm:aff id="I7">Department of Microbiology and Institute for Immunology, University of Pennsylvania, Philadelphia, PA 19104, USA</nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea>Department of Microbiology and Institute for Immunology, University of Pennsylvania, Philadelphia, PA 19104</wicri:regionArea><placeName><region type="state">Pennsylvanie</region></placeName></affiliation></author><author><name sortKey="Ertl, H C" sort="Ertl, H C" uniqKey="Ertl H" first="H. C." last="Ertl">H. C. Ertl</name><affiliation wicri:level="2"><nlm:aff id="I1">Wistar Institute, Philadelphia, PA 19104, USA</nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea>Wistar Institute, Philadelphia, PA 19104</wicri:regionArea><placeName><region type="state">Pennsylvanie</region></placeName></affiliation><affiliation wicri:level="2"><nlm:aff id="I7">Department of Microbiology and Institute for Immunology, University of Pennsylvania, Philadelphia, PA 19104, USA</nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea>Department of Microbiology and Institute for Immunology, University of Pennsylvania, Philadelphia, PA 19104</wicri:regionArea><placeName><region type="state">Pennsylvanie</region></placeName></affiliation></author></analytic><series><title level="j">BioMed Research International</title><idno type="ISSN">2314-6133</idno><idno type="eISSN">2314-6141</idno><imprint><date when="2013">2013</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Influenza viruses cause severe illnesses and death, mainly in the aged population. Protection afforded by licensed vaccines through subtype-specific neutralizing antibodies is incomplete, especially when the vaccine antigens fail to closely match those of the circulating viral strains. Efforts are underway to generate a so-called universal influenza vaccine expressing conserved viral sequences that induce broad protection to multiple strains of influenza virus through the induction of CD8<sup>+</sup> T cells. Here we assess the effect of a potent antiviral CD8<sup>+</sup> T cell response on influenza virus infection of young and aged mice. Our results show that CD8<sup>+</sup> T cell-inducing vaccines can provide some protection to young mice, but they exacerbate influenza virus-associated disease in aged mice, causing extensive lung pathology and death.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Bennett, Ak" uniqKey="Bennett A">AK Bennett</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Thompson, Ww" uniqKey="Thompson W">WW Thompson</name></author><author><name sortKey="Shay, Dk" uniqKey="Shay D">DK Shay</name></author><author><name sortKey="Weintraub, E" uniqKey="Weintraub E">E Weintraub</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Thompson, Ww" uniqKey="Thompson W">WW Thompson</name></author><author><name sortKey="Shay, Dk" uniqKey="Shay D">DK Shay</name></author><author><name sortKey="Weintraub, E" uniqKey="Weintraub E">E 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Biomed Res Int</journal-id><journal-id journal-id-type="iso-abbrev">Biomed Res Int</journal-id><journal-id journal-id-type="publisher-id">BMRI</journal-id><journal-title-group><journal-title>BioMed Research International</journal-title></journal-title-group><issn pub-type="ppub">2314-6133</issn><issn pub-type="epub">2314-6141</issn><publisher><publisher-name>Hindawi Publishing Corporation</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">24187666</article-id><article-id pub-id-type="pmc">3800650</article-id><article-id pub-id-type="doi">10.1155/2013/876314</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title>Influenza Virus Specific CD8<sup>+</sup> T Cells Exacerbate Infection Following High Dose Influenza Challenge of Aged Mice</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Parzych</surname><given-names>E. M.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I2"><sup>2</sup></xref></contrib><contrib contrib-type="author"><name><surname>DiMenna</surname><given-names>L. J.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I3"><sup>3</sup></xref></contrib><contrib contrib-type="author"><name><surname>Latimer</surname><given-names>B. P.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I2"><sup>2</sup></xref></contrib><contrib contrib-type="author"><name><surname>Small</surname><given-names>J. C.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I4"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Kannan</surname><given-names>S.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I4"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Manson</surname><given-names>B.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I5"><sup>5</sup></xref></contrib><contrib contrib-type="author"><name><surname>Lasaro</surname><given-names>M. O.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I6"><sup>6</sup></xref></contrib><contrib contrib-type="author"><name><surname>Wherry</surname><given-names>E. J.</given-names></name><xref ref-type="aff" rid="I7"><sup>7</sup></xref></contrib><contrib contrib-type="author"><name><surname>Ertl</surname><given-names>H. C.</given-names></name><xref ref-type="aff" rid="I1"><sup>1</sup></xref><xref ref-type="aff" rid="I7"><sup>7</sup></xref><xref ref-type="corresp" rid="cor1">*</xref></contrib></contrib-group><aff id="I1"><sup>1</sup>Wistar Institute, Philadelphia, PA 19104, USA</aff><aff id="I2"><sup>2</sup>Drexel University, Philadelphia, PA 19104, USA</aff><aff id="I3"><sup>3</sup>Columbia University, New York, NY 10027, USA</aff><aff id="I4"><sup>4</sup>Graduate Group of the University of Pennsylvania, Philadelphia, PA 19104, USA</aff><aff id="I5"><sup>5</sup>Cheyney University, Thornbury Township, PA 19139, USA</aff><aff id="I6"><sup>6</sup>Boehringer Ingelheim GMBH, Ridgefield, CT D6877, USA</aff><aff id="I7"><sup>7</sup>Department of Microbiology and Institute for Immunology, University of Pennsylvania, Philadelphia, PA 19104, USA</aff><author-notes><corresp id="cor1">*H. C. Ertl: <email>ertl@wistar.upenn.edu</email></corresp><fn fn-type="other"><p>Academic Editor: Kelvin To</p></fn></author-notes><pub-date pub-type="ppub"><year>2013</year></pub-date><pub-date pub-type="epub"><day>26</day><month>9</month><year>2013</year></pub-date><volume>2013</volume><elocation-id>876314</elocation-id><history><date date-type="received"><day>3</day><month>4</month><year>2013</year></date><date date-type="rev-recd"><day>3</day><month>6</month><year>2013</year></date><date date-type="accepted"><day>4</day><month>6</month><year>2013</year></date></history><permissions><copyright-statement>Copyright © 2013 E. M. Parzych et al.</copyright-statement><copyright-year>2013</copyright-year><license xlink:href="https://creativecommons.org/licenses/by/3.0/"><license-p>This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.</license-p></license></permissions><abstract><p>Influenza viruses cause severe illnesses and death, mainly in the aged population. Protection afforded by licensed vaccines through subtype-specific neutralizing antibodies is incomplete, especially when the vaccine antigens fail to closely match those of the circulating viral strains. Efforts are underway to generate a so-called universal influenza vaccine expressing conserved viral sequences that induce broad protection to multiple strains of influenza virus through the induction of CD8<sup>+</sup> T cells. Here we assess the effect of a potent antiviral CD8<sup>+</sup> T cell response on influenza virus infection of young and aged mice. Our results show that CD8<sup>+</sup> T cell-inducing vaccines can provide some protection to young mice, but they exacerbate influenza virus-associated disease in aged mice, causing extensive lung pathology and death.</p></abstract></article-meta></front><floats-group><fig id="fig1" orientation="portrait" position="float"><label>Figure 1</label><caption><p>Vaccine-induced NP-specific CD8<sup>+</sup> T cell responses in young (left) and aged C57Bl/5 female mice (right) in response to vaccination. Groups of young mice (6–8 weeks, <italic>n</italic> = 20) and aged mice (>18 months, <italic>n</italic> = 23) were vaccinated with 10<sup>10</sup> vp of AdC68NP, given IM. They were bled 2, 4, 6, and 8 weeks after vaccination. PBMCs were isolated and stained with an NP-specific tetramer and antibodies to CD8 to identify the frequency of NP-specific CD8<sup>+</sup> T cells. A portion of AdC68NP-primed mice were then boosted with 0.8 × 10<sup>5</sup> TCID<sub>50</sub> of A/X31, along with age-matched groups of previously naïve mice.</p></caption><graphic xlink:href="BMRI2013-876314.001"></graphic></fig><fig id="fig2" orientation="portrait" position="float"><label>Figure 2</label><caption><p>Vaccine-induced numbers of NP-specific CD8<sup>+</sup> T cells in the blood, spleen, and lung. Groups of young and aged mice were vaccinated with 10<sup>10</sup> vp of AdC68NP, given IM. A portion of these primed mice, along with age-matched groups of naïve mice, were subsequently boosted at 2 months after priming with 0.8 × 10<sup>5</sup> TCID<sub>50</sub> of A/X31, given IN. 2 months after the boost, cohorts were euthanized before or after challenge with 3LD<sub>50</sub> A/PR8. Lymphocytes were harvested from the blood, spleen, and lungs of these mice and underwent tetramer and CD8 staining to determine the total number of tetramer<sup>+</sup>CD8<sup>+</sup> cells in each compartment. Graphs show numbers of NP-specific CD8<sup>+</sup> T cells over 10<sup>7</sup> live PBMCs (upper row) or per all live mononuclear cells from spleens (middle rows) or lungs (low rows). The bars reflect group averages ± SD; gray bars represent young mice and black bars represent aged mice. Asterisks indicate significant differences between groups at the <italic>P</italic> ≤ 0.05 level.</p></caption><graphic xlink:href="BMRI2013-876314.002"></graphic></fig><fig id="fig3" orientation="portrait" position="float"><label>Figure 3</label><caption><p>Cytokine production by vaccine-induced NP-specific CD8<sup>+</sup> T cells. Groups of young and aged mice were either vaccinated with 10<sup>10</sup> vps of AdC68NP only (young and aged: <italic>n</italic> = 4 each) or were then subsequently boosted 8 weeks later with 0.8 × 10<sup>5</sup> TCID<sub>50</sub> of A/X31 (young: <italic>n</italic> = 4, aged not tested as all died following infection). Additional age-matched groups were added that received only A/X31 boost (young and aged: <italic>n</italic> = 4 each). All mice were then challenged with 3LD<sub>50</sub> A/PR8 2 months following the boost, along with age-matched control groups (young: <italic>n</italic> = 5, aged: <italic>n</italic> = 3). At day 20 following the challenge, splenocytes were isolated, and an ICS was conducted to detect production of IFN-<italic>γ</italic> (G), IL-2 (I), and TNF-<italic>α</italic> (T).</p></caption><graphic xlink:href="BMRI2013-876314.003"></graphic></fig><fig id="fig4" orientation="portrait" position="float"><label>Figure 4</label><caption><p>(a) Survival of young and aged mice following A/PR8 challenge. (b) Weight loss following A/PR8 challenge. (c) Lung viral titers in lungs at day 5 after A/PR8 challenge. (d) Lung histology. (e) Lung infiltrates.</p></caption><graphic xlink:href="BMRI2013-876314.004"></graphic></fig><fig id="fig5" orientation="portrait" position="float"><label>Figure 5</label><caption><p>Comparison of CD8<sup>+</sup> T cell frequencies and phenotypes between protected and nonprotected aged mice.</p></caption><graphic xlink:href="BMRI2013-876314.005"></graphic></fig><table-wrap id="tab1" orientation="portrait" position="float"><label>Table 1</label><caption><p>Experimental study design.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">Groups</th><th align="center" rowspan="1" colspan="1">Groups of mice<break></break>(nos. of mice)</th><th align="center" rowspan="1" colspan="1">Prime</th><th align="center" rowspan="1" colspan="1">Analyses after the prime (compartment)</th><th align="center" rowspan="1" colspan="1">Boost (time after prime)</th><th align="center" rowspan="1" colspan="1">Analyses after the boost (compartment)</th><th align="center" rowspan="1" colspan="1">Challenge (time after prime)</th><th align="center" rowspan="1" colspan="1">Time of analyses after challenge (compartment)</th></tr></thead><tbody><tr><td colspan="8" align="center" rowspan="1">Experiment 1 (Figures <xref ref-type="fig" rid="fig1">1(a)</xref> and <xref ref-type="fig" rid="fig1">1(b)</xref>)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 1</td><td align="center" rowspan="1" colspan="1">Young (15)</td><td align="center" rowspan="4" colspan="1">AdC68NP</td><td align="center" rowspan="4" colspan="1">Wks 2, 4, 6, and 8<break></break>(blood)</td><td align="center" rowspan="2" colspan="1"> None</td><td align="center" rowspan="6" colspan="1">Wks 2, 4, 6, 8<break></break>(blood)</td><td align="center" rowspan="8" colspan="1">A/PR8 (mo 4)</td><td align="center" rowspan="8" colspan="1">Day 20<break></break>(blood)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 2</td><td align="center" rowspan="1" colspan="1">Aged (13)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 3</td><td align="center" rowspan="1" colspan="1">Young (4)</td><td align="center" rowspan="4" colspan="1">A/X31 (mo 2)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 4</td><td align="center" rowspan="1" colspan="1">Aged (13)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 5</td><td align="center" rowspan="1" colspan="1">Young (5)</td><td align="center" rowspan="4" colspan="1"> None</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 6</td><td align="center" rowspan="1" colspan="1">Aged (10)</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 7</td><td align="center" rowspan="1" colspan="1">Young (25)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="2" colspan="1"> None</td><td align="center" rowspan="2" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">Group 8</td><td align="center" rowspan="1" colspan="1">Aged (33)</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" colspan="8" rowspan="1"><hr></hr></td></tr><tr><td colspan="8" align="center" rowspan="1">Experiment 2 (Figures <xref ref-type="fig" rid="fig2">2(a)</xref>, <xref ref-type="fig" rid="fig2">2(b)</xref>, and <xref ref-type="fig" rid="fig2">2(c)</xref>)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 1</td><td align="center" rowspan="1" colspan="1">Young (5)</td><td align="center" rowspan="4" colspan="1">AdC68NP</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="2" colspan="1">None</td><td align="center" rowspan="8" colspan="1">Wk 8<break></break>(blood, spleen, and lung)</td><td align="center" rowspan="8" colspan="1">A/PR8 (mo 4)</td><td align="center" rowspan="8" colspan="1">Days 5, 20<break></break>(blood, spleen, and lung)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 2</td><td align="center" rowspan="1" colspan="1">Aged (8)</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 3</td><td align="center" rowspan="1" colspan="1">Young (5)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="4" colspan="1">A/X31 (mo 2)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 4</td><td align="center" rowspan="1" colspan="1">Aged (5)</td><td align="center" rowspan="1" colspan="1">None</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 5</td><td align="center" rowspan="1" colspan="1">Young (5)</td><td align="center" rowspan="4" colspan="1"> None</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 6</td><td align="center" rowspan="1" colspan="1">Aged (5)</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 7</td><td align="center" rowspan="1" colspan="1">Young (2)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="2" colspan="1">None</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 8</td><td align="center" rowspan="1" colspan="1">Aged (4)</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" colspan="8" rowspan="1"><hr></hr></td></tr><tr><td colspan="8" align="center" rowspan="1">Experiment 3 (Figures <xref ref-type="fig" rid="fig3">3(a)</xref> and <xref ref-type="fig" rid="fig3">3(b)</xref>)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 1</td><td align="center" rowspan="1" colspan="1">Young (4)</td><td align="center" rowspan="3" colspan="1">AdC68NP</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="2" colspan="1">None</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="7" colspan="1">A/PR8 (mo 4)</td><td align="center" rowspan="7" colspan="1">Day 20<break></break>(spleen)</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 2</td><td align="center" rowspan="1" colspan="1">Aged (4)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 3</td><td align="center" rowspan="1" colspan="1">Young (4)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="3" colspan="1">A/X31 (mo 2)</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 4</td><td align="center" rowspan="1" colspan="1">Young (5)</td><td align="center" rowspan="4" colspan="1"> None</td><td align="center" rowspan="1" colspan="1">None</td><td align="center" rowspan="1" colspan="1">None</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 5</td><td align="center" rowspan="1" colspan="1">Aged (4)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 6</td><td align="center" rowspan="1" colspan="1">Young (5)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="2" colspan="1">None</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 7</td><td align="center" rowspan="1" colspan="1">Aged (3)</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" colspan="8" rowspan="1"><hr></hr></td></tr><tr><td colspan="8" align="center" rowspan="1">Experiment 4 (Figures <xref ref-type="fig" rid="fig4">4(a)</xref>, <xref ref-type="fig" rid="fig4">4(b)</xref>, <xref ref-type="fig" rid="fig4">4(c)</xref>, and <xref ref-type="fig" rid="fig4">4(e)</xref>)<sup>1</sup></td></tr><tr><td align="left" rowspan="1" colspan="1">Group 1</td><td align="center" rowspan="1" colspan="1">Young (15) [5]</td><td align="center" rowspan="4" colspan="1">AdC68NP</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="2" colspan="1">None</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="8" colspan="1">A/PR8 (mo 4)</td><td align="center" rowspan="8" colspan="1">Checked daily for weight loss and survival;<break></break>viral titers in lungs tested on day 5;<break></break>lung infiltrates tested on day 5.</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 2</td><td align="center" rowspan="1" colspan="1">Aged (13) [15] [<bold><italic>7</italic></bold>]</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 3</td><td align="center" rowspan="1" colspan="1">Young (4) [5]</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="4" colspan="1">A/X31 (mo 2)<sup>2</sup></td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 4</td><td align="center" rowspan="1" colspan="1">Aged (8) [5] [<bold><italic>5</italic></bold>]</td><td align="center" rowspan="1" colspan="1">None</td><td align="center" rowspan="1" colspan="1">None</td></tr><tr><td align="left" rowspan="1" colspan="1">Group 5</td><td align="center" rowspan="1" colspan="1">Young (5) [5]</td><td align="center" rowspan="4" colspan="1"> None</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 6</td><td align="center" rowspan="1" colspan="1">Aged (10) [7] [<bold><italic>4</italic></bold>]</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 7</td><td align="center" rowspan="1" colspan="1">Young (25) [9]</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="2" colspan="1">None</td><td align="center" rowspan="1" colspan="1"> </td></tr><tr><td align="left" rowspan="1" colspan="1">Group 8</td><td align="center" rowspan="1" colspan="1">Aged (38) [11] [<bold><italic>7</italic></bold>]</td><td align="center" rowspan="1" colspan="1"> </td><td align="center" rowspan="1" colspan="1"> </td></tr></tbody></table><table-wrap-foot><fn><p><sup>1</sup>Number of mice tested for survival are shown in (), numbers of mice tested for viral loads are shown in regular font in [], and numbers of mice used to determine subsets of lung infiltrating cells are shown on bold italic in []. </p></fn><fn><p><sup>2</sup>Some of the mice were primed at a young age. They were boosted ~18 months later.</p></fn></table-wrap-foot></table-wrap></floats-group></pmc><affiliations><list><country><li>États-Unis</li></country><region><li>Connecticut</li><li>Pennsylvanie</li><li>État de New York</li></region><settlement><li>New York</li></settlement><orgName><li>Université Columbia</li></orgName></list><tree><country name="États-Unis"><region name="Pennsylvanie"><name sortKey="Parzych, E M" sort="Parzych, E M" uniqKey="Parzych E" first="E. M." last="Parzych">E. M. Parzych</name></region><name sortKey="Dimenna, L J" sort="Dimenna, L J" uniqKey="Dimenna L" first="L. J." last="Dimenna">L. J. Dimenna</name><name sortKey="Dimenna, L J" sort="Dimenna, L J" uniqKey="Dimenna L" first="L. J." last="Dimenna">L. J. Dimenna</name><name sortKey="Ertl, H C" sort="Ertl, H C" uniqKey="Ertl H" first="H. C." last="Ertl">H. C. Ertl</name><name sortKey="Ertl, H C" sort="Ertl, H C" uniqKey="Ertl H" first="H. C." last="Ertl">H. C. Ertl</name><name sortKey="Kannan, S" sort="Kannan, S" uniqKey="Kannan S" first="S." last="Kannan">S. Kannan</name><name sortKey="Kannan, S" sort="Kannan, S" uniqKey="Kannan S" first="S." last="Kannan">S. Kannan</name><name sortKey="Lasaro, M O" sort="Lasaro, M O" uniqKey="Lasaro M" first="M. O." last="Lasaro">M. O. Lasaro</name><name sortKey="Lasaro, M O" sort="Lasaro, M O" uniqKey="Lasaro M" first="M. O." last="Lasaro">M. O. Lasaro</name><name sortKey="Latimer, B P" sort="Latimer, B P" uniqKey="Latimer B" first="B. P." last="Latimer">B. P. Latimer</name><name sortKey="Latimer, B P" sort="Latimer, B P" uniqKey="Latimer B" first="B. P." last="Latimer">B. P. Latimer</name><name sortKey="Manson, B" sort="Manson, B" uniqKey="Manson B" first="B." last="Manson">B. Manson</name><name sortKey="Manson, B" sort="Manson, B" uniqKey="Manson B" first="B." last="Manson">B. Manson</name><name sortKey="Parzych, E M" sort="Parzych, E M" uniqKey="Parzych E" first="E. M." last="Parzych">E. M. Parzych</name><name sortKey="Small, J C" sort="Small, J C" uniqKey="Small J" first="J. C." last="Small">J. C. Small</name><name sortKey="Small, J C" sort="Small, J C" uniqKey="Small J" first="J. C." last="Small">J. C. Small</name><name sortKey="Wherry, E J" sort="Wherry, E J" uniqKey="Wherry E" first="E. J." last="Wherry">E. J. Wherry</name></country></tree></affiliations></record>Pour manipuler ce document sous Unix (Dilib)
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