Cross-Recognition of Avian H5N1 Influenza Virus by Human Cytotoxic T-Lymphocyte Populations Directed to Human Influenza A Virus▿
Identifieur interne : 000360 ( Pmc/Curation ); précédent : 000359; suivant : 000361Cross-Recognition of Avian H5N1 Influenza Virus by Human Cytotoxic T-Lymphocyte Populations Directed to Human Influenza A Virus▿
Auteurs : J. H. C. M. Kreijtz [Pays-Bas] ; G. De Mutsert [Pays-Bas] ; C. A. Van Baalen [Pays-Bas] ; R. A. M. Fouchier [Pays-Bas] ; A. D. M. E. Osterhaus [Pays-Bas] ; G. F. Rimmelzwaan [Pays-Bas]Source :
- Journal of Virology [ 0022-538X ] ; 2008.
Abstract
Since the number of human cases of infection with avian H5N1 influenza viruses is ever increasing, a pandemic outbreak caused by these viruses is feared. Therefore, in addition to virus-specific antibodies, there is considerable interest in immune correlates of protection against these viruses, which could be a target for the development of more universal vaccines. After infection with seasonal influenza A viruses of the H3N2 and H1N1 subtypes, individuals develop virus-specific cytotoxic T-lymphocyte responses, which are mainly directed against the relatively conserved internal proteins of the virus, like the nucleoprotein (NP). Virus-specific cytotoxic T lymphocytes (CTL) are known to contribute to protective immunity against infection, but knowledge about the extent of cross-reactivity with avian H5N1 influenza viruses is sparse. In the present study, we evaluated the cross-reactivity with H5N1 influenza viruses of polyclonal CTL obtained from a group of well-defined HLA-typed study subjects. To this end, the recognition of synthetic peptides representing H5N1 analogues of known CTL epitopes was studied. In addition, the ability of CTL specific for seasonal H3N2 influenza virus to recognize the NP of H5N1 influenza virus or H5N1 virus-infected cells was tested. It was concluded that, apart from some individual epitopes that displayed amino acid variation between H3N2 and H5N1 influenza viruses, considerable cross-reactivity exists with H5N1 viruses. This preexisting cross-reactive T-cell immunity in the human population may dampen the impact of a next pandemic.
Url:
DOI: 10.1128/JVI.02694-07
PubMed: 18353950
PubMed Central: 2395172
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Cross-Recognition of Avian H5N1 Influenza Virus by Human Cytotoxic T-Lymphocyte Populations Directed to Human Influenza A Virus<xref ref-type="fn" rid="fn1">▿</xref> </title><author><name sortKey="Kreijtz, J H C M" sort="Kreijtz, J H C M" uniqKey="Kreijtz J" first="J. H. C. M." last="Kreijtz">J. H. C. M. Kreijtz</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="De Mutsert, G" sort="De Mutsert, G" uniqKey="De Mutsert G" first="G." last="De Mutsert">G. De Mutsert</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Van Baalen, C A" sort="Van Baalen, C A" uniqKey="Van Baalen C" first="C. A." last="Van Baalen">C. A. Van Baalen</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Fouchier, R A M" sort="Fouchier, R A M" uniqKey="Fouchier R" first="R. A. M." last="Fouchier">R. A. M. Fouchier</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Osterhaus, A D M E" sort="Osterhaus, A D M E" uniqKey="Osterhaus A" first="A. D. M. E." last="Osterhaus">A. D. M. E. Osterhaus</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Rimmelzwaan, G F" sort="Rimmelzwaan, G F" uniqKey="Rimmelzwaan G" first="G. F." last="Rimmelzwaan">G. F. Rimmelzwaan</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">18353950</idno><idno type="pmc">2395172</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2395172</idno><idno type="RBID">PMC:2395172</idno><idno type="doi">10.1128/JVI.02694-07</idno><date when="2008">2008</date><idno type="wicri:Area/Pmc/Corpus">000360</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000360</idno><idno type="wicri:Area/Pmc/Curation">000360</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Curation">000360</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Cross-Recognition of Avian H5N1 Influenza Virus by Human Cytotoxic T-Lymphocyte Populations Directed to Human Influenza A Virus<xref ref-type="fn" rid="fn1">▿</xref> </title><author><name sortKey="Kreijtz, J H C M" sort="Kreijtz, J H C M" uniqKey="Kreijtz J" first="J. H. C. M." last="Kreijtz">J. H. C. M. Kreijtz</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="De Mutsert, G" sort="De Mutsert, G" uniqKey="De Mutsert G" first="G." last="De Mutsert">G. De Mutsert</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Van Baalen, C A" sort="Van Baalen, C A" uniqKey="Van Baalen C" first="C. A." last="Van Baalen">C. A. Van Baalen</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Fouchier, R A M" sort="Fouchier, R A M" uniqKey="Fouchier R" first="R. A. M." last="Fouchier">R. A. M. Fouchier</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Osterhaus, A D M E" sort="Osterhaus, A D M E" uniqKey="Osterhaus A" first="A. D. M. E." last="Osterhaus">A. D. M. E. Osterhaus</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author><author><name sortKey="Rimmelzwaan, G F" sort="Rimmelzwaan, G F" uniqKey="Rimmelzwaan G" first="G. F." last="Rimmelzwaan">G. F. Rimmelzwaan</name><affiliation wicri:level="1"><nlm:aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</nlm:aff><country xml:lang="fr">Pays-Bas</country><wicri:regionArea>Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam</wicri:regionArea></affiliation></author></analytic><series><title level="j">Journal of Virology</title><idno type="ISSN">0022-538X</idno><idno type="eISSN">1098-5514</idno><imprint><date when="2008">2008</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Since the number of human cases of infection with avian H5N1 influenza viruses is ever increasing, a pandemic outbreak caused by these viruses is feared. Therefore, in addition to virus-specific antibodies, there is considerable interest in immune correlates of protection against these viruses, which could be a target for the development of more universal vaccines. After infection with seasonal influenza A viruses of the H3N2 and H1N1 subtypes, individuals develop virus-specific cytotoxic T-lymphocyte responses, which are mainly directed against the relatively conserved internal proteins of the virus, like the nucleoprotein (NP). Virus-specific cytotoxic T lymphocytes (CTL) are known to contribute to protective immunity against infection, but knowledge about the extent of cross-reactivity with avian H5N1 influenza viruses is sparse. In the present study, we evaluated the cross-reactivity with H5N1 influenza viruses of polyclonal CTL obtained from a group of well-defined HLA-typed study subjects. To this end, the recognition of synthetic peptides representing H5N1 analogues of known CTL epitopes was studied. In addition, the ability of CTL specific for seasonal H3N2 influenza virus to recognize the NP of H5N1 influenza virus or H5N1 virus-infected cells was tested. It was concluded that, apart from some individual epitopes that displayed amino acid variation between H3N2 and H5N1 influenza viruses, considerable cross-reactivity exists with H5N1 viruses. This preexisting cross-reactive T-cell immunity in the human population may dampen the impact of a next pandemic.</p></div></front></TEI><pmc article-type="research-article"><pmc-comment>The publisher of this article does not allow downloading of the full text in XML form.</pmc-comment>
<front><journal-meta><journal-id journal-id-type="nlm-ta">J Virol</journal-id><journal-id journal-id-type="publisher-id">jvi</journal-id><journal-title>Journal of Virology</journal-title><issn pub-type="ppub">0022-538X</issn><issn pub-type="epub">1098-5514</issn><publisher><publisher-name>American Society for Microbiology (ASM)</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">18353950</article-id><article-id pub-id-type="pmc">2395172</article-id><article-id pub-id-type="publisher-id">2694-07</article-id><article-id pub-id-type="doi">10.1128/JVI.02694-07</article-id><article-categories><subj-group subj-group-type="heading"><subject>Pathogenesis and Immunity</subject></subj-group></article-categories><title-group><article-title>Cross-Recognition of Avian H5N1 Influenza Virus by Human Cytotoxic T-Lymphocyte Populations Directed to Human Influenza A Virus<xref ref-type="fn" rid="fn1">▿</xref> </article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Kreijtz</surname><given-names>J. H. C. M.</given-names></name><xref ref-type="aff" rid="aff0"></xref></contrib><contrib contrib-type="author"><name><surname>de Mutsert</surname><given-names>G.</given-names></name><xref ref-type="aff" rid="aff0"></xref></contrib><contrib contrib-type="author"><name><surname>van Baalen</surname><given-names>C. A.</given-names></name><xref ref-type="aff" rid="aff0"></xref></contrib><contrib contrib-type="author"><name><surname>Fouchier</surname><given-names>R. A. M.</given-names></name><xref ref-type="aff" rid="aff0"></xref></contrib><contrib contrib-type="author"><name><surname>Osterhaus</surname><given-names>A. D. M. E.</given-names></name><xref ref-type="aff" rid="aff0"></xref></contrib><contrib contrib-type="author"><name><surname>Rimmelzwaan</surname><given-names>G. F.</given-names></name><xref ref-type="aff" rid="aff0"></xref><xref ref-type="corresp" rid="cor1">*</xref></contrib></contrib-group><aff id="aff0">Department of Virology, Erasmus Medical Center, P.O. Box 2040, Rotterdam, The Netherlands</aff><author-notes><fn id="cor1"><label>*</label><p>Corresponding author. Mailing address: Department of Virology, Erasmus Medical Center, P.O. Box 2040, 3000 CA Rotterdam, The Netherlands. Phone: 31 10 7044243. Fax: 31 10 7044760. E-mail: <email>g.rimmelzwaan@erasmusmc.nl</email></p></fn></author-notes><pub-date pub-type="ppub"><month>6</month><year>2008</year></pub-date><pub-date pub-type="epub"><day>19</day><month>3</month><year>2008</year></pub-date><volume>82</volume><issue>11</issue><fpage>5161</fpage><lpage>5166</lpage><history><date date-type="received"><day>19</day><month>12</month><year>2007</year></date><date date-type="accepted"><day>4</day><month>3</month><year>2008</year></date></history><permissions><copyright-statement>Copyright © 2008, American Society for Microbiology</copyright-statement></permissions><self-uri xlink:title="pdf" xlink:href="zjv01108005161.pdf"></self-uri><abstract><p>Since the number of human cases of infection with avian H5N1 influenza viruses is ever increasing, a pandemic outbreak caused by these viruses is feared. Therefore, in addition to virus-specific antibodies, there is considerable interest in immune correlates of protection against these viruses, which could be a target for the development of more universal vaccines. After infection with seasonal influenza A viruses of the H3N2 and H1N1 subtypes, individuals develop virus-specific cytotoxic T-lymphocyte responses, which are mainly directed against the relatively conserved internal proteins of the virus, like the nucleoprotein (NP). Virus-specific cytotoxic T lymphocytes (CTL) are known to contribute to protective immunity against infection, but knowledge about the extent of cross-reactivity with avian H5N1 influenza viruses is sparse. In the present study, we evaluated the cross-reactivity with H5N1 influenza viruses of polyclonal CTL obtained from a group of well-defined HLA-typed study subjects. To this end, the recognition of synthetic peptides representing H5N1 analogues of known CTL epitopes was studied. In addition, the ability of CTL specific for seasonal H3N2 influenza virus to recognize the NP of H5N1 influenza virus or H5N1 virus-infected cells was tested. It was concluded that, apart from some individual epitopes that displayed amino acid variation between H3N2 and H5N1 influenza viruses, considerable cross-reactivity exists with H5N1 viruses. This preexisting cross-reactive T-cell immunity in the human population may dampen the impact of a next pandemic.</p></abstract></article-meta></front><floats-wrap><fig position="float" id="f1"><label>FIG. 1.</label><caption><p>The presence of known CTL epitopes in H5N1 strains. The percentage of H5N1 viruses with an epitope sequence identical to human influenza viruses (white bars) is shown. The black bars indicate the percentage of H5N1 viruses with one or more amino acid substitutions in the epitope sequence. The absolute numbers of each variant of an epitope are shown in panel B, where each color represents a single variant (sequences can be found in Table <xref ref-type="table" rid="t1">1</xref>). For this analysis almost 900 H5N1 viruses for which sequence information was available in the influenza sequence database (<xref ref-type="bibr" rid="r12">12</xref>) were analyzed.</p></caption><graphic xlink:href="zjv0110805890001"></graphic></fig><fig position="float" id="f2"><label>FIG. 2.</label><caption><p>Epitope-specific IFN-γ production by CTLs after stimulation with peptide-pulsed BLCL. The number of IFN-γ-producing cells per 10,000 CD8<sup>+</sup> T cells (5,000 cells for subject 3) from subjects from HLA groups I (A), II (B), and III (C) were measured by ELISPOT assay. CD8<sup>+</sup> T cells were isolated from PBMC populations expanded in vitro with influenza virus A/NL/18/94 and subsequently stimulated with peptide variants as indicated. (*, peptide sequence of the known human influenza virus CTL epitopes).</p></caption><graphic xlink:href="zjv0110805890002"></graphic></fig><fig position="float" id="f3"><label>FIG. 3.</label><caption><p>Recognition of NP derived from H3N2 and H5N1 influenza virus by in vitro expanded PBMC specific for influenza virus A/NL/18/94 (H3N2). The lytic activity of in vitro expanded PBMC was tested with MHC-I matched BLCL nucleofected with NP-GFP coding plasmid (NP of either influenza virus A/NL/18/94 [black circles] or A/VN/1194/04 [open circles]) or empty control GFP plasmid (gray circles). The NP-specific lytic activity was tested with PBMC of the subjects from group I (subjects 1 to 4), II (subjects 5 to 9), and III (subjects 10 to 14 and 16). E:T ratios were 0, 3.125, 6.25, 12.5, 25, and 50 for all subjects except numbers 13 and 14, for whom E:T ratios were 0, 0.3, 1, 3, 10 and 30. The lytic activities against control plasmid-transfected target cells are not visible for subjects 13 and 14 as a result of negative values for the percentages of specific lysis, which were caused by slight increases in the number of GFP-positive viable cells. Standard deviation of the means was <10%.</p></caption><graphic xlink:href="zjv0110805890003"></graphic></fig><fig position="float" id="f4"><label>FIG. 4.</label><caption><p>Recognition of influenza virus-infected BLCL by CTLs. The number of IFN-γ-producing cells per 10,000 CD8<sup>+</sup> T cells was measured by ELISPOT assay after stimulation with BLCL infected with influenza virus A/NL/18/94 or A/VN/1194/04. Each symbol represents an individual subject from group I (A), II (B), or III (C). Uninfected BLCL were used as negative controls. The horizontal bars represent the average responses of all study subjects in groups I, II, and III.</p></caption><graphic xlink:href="zjv0110805890004"></graphic></fig><table-wrap position="float" id="t1"><label>TABLE 1.</label><caption><p>Variant sequences of known CTL epitopes in H5N1 viruses</p></caption><table frame="hsides" rules="groups"><thead><tr><th colspan="1" rowspan="2" align="center" valign="middle">Sequence name</th><th colspan="9" rowspan="1" align="center" valign="bottom">Variant sequence of the CTL epitope at the indicated HLA allele<xref ref-type="table-fn" rid="t1fn1"><italic>a</italic></xref><hr></hr></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th><th colspan="1" rowspan="1" align="center" valign="bottom"></th></tr><tr><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A1 PB1 (591-599)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A3 M1 (13-21)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A*0201 M1 (58-66)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*3501 M1 (128-135)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A*0201 NS1 (122-130)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A*0201 NS1 (123-132)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*44 NS1 (158-166)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A1 NP (44-52)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A*6801 NP (91-99)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*1402 NP (146-154)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*2705 NP (174-184)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A3 NP (188-198)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-A3 NP (265-273)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*3701 NP (339-347)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*08 NP (380-388)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*2702 NP (381-388)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*2705 NP (383-391)</th><th colspan="1" rowspan="1" align="center" valign="bottom">HLA-B*3501 NP (418-426)</th></tr></thead><tbody><tr><td colspan="1" rowspan="1" align="left" valign="top">Epitope</td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>VS<underline>D</underline>GGPNL<underline>Y</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>SIIPSGPL<underline>K</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>GIL<underline>G</underline>FVFT<underline>L</underline></monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>A<underline>S</underline>CMGLI<underline>Y</underline></monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>AIM<underline>D</underline>KNI<underline>IL</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>I<underline>MD</underline>KNI<underline>IL</underline>KA</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>G<underline>E</underline>ISPLPSL</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>CT<underline>E</underline>LKLSD<underline>Y</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>K<underline>T</underline>GGPIYK<underline>R</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>TT<underline>Y</underline>Q<underline>R</underline>TRA<underline>L</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>R<underline>R</underline>SGAAGAAVK</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>T<underline>M</underline>VMELVRMI<underline>K</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>I<underline>L</underline>RGSVAH</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>E<underline>D</underline>LRVLS<underline>FI</underline></monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>ELRS<underline>R</underline>YWAI</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>L<underline>R</underline>SRYWAI</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>S<underline>R</underline>YWAIRTR</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>L<underline>P</underline>FEKSTV<underline>M</underline></monospace></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top">Variant</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 1</td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>I--------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-V-------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-M-------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>------S-</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----T---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----T-----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>------H--</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--------H</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-------R-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>S--------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>---------I-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>------I----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----I---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----S---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----K----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>---K----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-K-------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----RA-I-</monospace></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 2</td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-------P-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--V------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--W------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----V---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----V-----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-------F-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--------Q</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----V---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>A--------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-I------L--</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--A--------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----M---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----IS---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>D-LGK---K</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-LGK---K</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>------K--</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----R--I-</monospace></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 3</td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>------IP-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>F--------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-------T-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>------T---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>---L-----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>------N--</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----F-RG</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>AA-------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--F--------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----QI----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>V--------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----H---</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>----H---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-------P-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----RAAI-</monospace></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 4</td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>G--------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-T-------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----TV--</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----TV----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--------I</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>------T--</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--------G</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>A--H-----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>Q----VI----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--------K</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----RV-I-</monospace></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 5</td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-A-------</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-L-------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----A---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----A-----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----S---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>---F-----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>AP-------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--------P</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----RA---</monospace></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 6</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>ATS------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-T-----T-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>T-----T---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-------Y-</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>A-------V</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>GK---KWMI</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 7</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--VR----Q</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----D---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----D-----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>A--------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>A-SQ-----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--H------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 8</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--VL-----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>--V--T---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-V--T-----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-----I---</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--------G</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 9</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>S----T---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----T-S---</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 10</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----T-S-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>--N-------</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 11</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>---N-----</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-V--T-T---</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 12</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>--V--T-T-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-V----T---</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 13</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>--V----T-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----T-L---</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 14</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----T-L-</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----I-----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 15</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----I---</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>-V--TV----</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 16</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>--V--TV--</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----T--W--</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr><tr><td colspan="1" rowspan="1" align="left" valign="top"> 17</td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"><monospace>-----T--W</monospace></td><td colspan="1" rowspan="1" align="left" valign="top"><monospace>----T----G</monospace></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td><td colspan="1" rowspan="1" align="center" valign="top"></td></tr></tbody></table><table-wrap-foot><fn id="t1fn1"><label>a</label><p>Variant sequences of known CTL epitopes were ranked according to their relative prevalence. Anchor residues of the epitopes are underlined. Amino acid residue positions are given in parentheses.</p></fn></table-wrap-foot></table-wrap></floats-wrap></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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