Antigenic Change in Human Influenza A(H2N2) Viruses Detected by Using Human Plasma from Aged and Younger Adult Individuals
Identifieur interne : 000C49 ( Pmc/Curation ); précédent : 000C48; suivant : 000C50Antigenic Change in Human Influenza A(H2N2) Viruses Detected by Using Human Plasma from Aged and Younger Adult Individuals
Auteurs : Yukimasa Matsuzawa ; Kiyoko Iwatsuki-Horimoto ; Yoshinori Nishimoto ; Yukiko Abe ; Satoshi Fukuyama ; Taiki Hamabata ; Moe Okuda ; Yui Go ; Tokiko Watanabe ; Masaki Imai ; Yasumichi Arai ; Ron A. M. Fouchier ; Seiya Yamayoshi ; Yoshihiro Kawaoka [Japon, États-Unis]Source :
- Viruses [ 1999-4915 ] ; 2019.
Abstract
Human influenza A(H2N2) viruses emerged in 1957 and were replaced by A(H3N2) viruses in 1968. The antigenicity of human H2N2 viruses has been tested by using ferret antisera or mouse and human monoclonal antibodies. Here, we examined the antigenicity of human H2N2 viruses by using human plasma samples obtained from 50 aged individuals who were born between 1928 and 1933 and from 33 younger adult individuals who were born after 1962. The aged individuals possessed higher neutralization titers against H2N2 viruses isolated in 1957 and 1963 than those against H2N2 viruses isolated in 1968, whereas the younger adults who were born between 1962 and 1968 possessed higher neutralization titers against H2N2 viruses isolated in 1963 than those against other H2N2 viruses. Antigenic cartography revealed the antigenic changes that occurred in human H2N2 viruses during circulation in humans for 11 years, as detected by ferret antisera. These results show that even though aged individuals were likely exposed to more recent H2N2 viruses that are antigenically distinct from the earlier H2N2 viruses, they did not possess high neutralizing antibody titers to the more recent viruses, suggesting immunological imprinting of these individuals with the first H2N2 viruses they encountered and that this immunological imprinting lasts for over 50 years.
Url:
DOI: 10.3390/v11110978
PubMed: 31652870
PubMed Central: 6893718
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Antigenic Change in Human Influenza A(H2N2) Viruses Detected by Using Human Plasma from Aged and Younger Adult Individuals</title><author><name sortKey="Matsuzawa, Yukimasa" sort="Matsuzawa, Yukimasa" uniqKey="Matsuzawa Y" first="Yukimasa" last="Matsuzawa">Yukimasa Matsuzawa</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Iwatsuki Horimoto, Kiyoko" sort="Iwatsuki Horimoto, Kiyoko" uniqKey="Iwatsuki Horimoto K" first="Kiyoko" last="Iwatsuki-Horimoto">Kiyoko Iwatsuki-Horimoto</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Nishimoto, Yoshinori" sort="Nishimoto, Yoshinori" uniqKey="Nishimoto Y" first="Yoshinori" last="Nishimoto">Yoshinori Nishimoto</name><affiliation><nlm:aff id="af2-viruses-11-00978">Center for Supercentenarian Medical Research, Shinjuku-ku, Tokyo 160-8582, Japan;<email>ynishimo@z3.keio.jp</email> (Y.N.);<email>yukiko_abe@keio.jp</email> (Y.A.);<email>yasumich@keio.jp</email> (Y.A.)</nlm:aff></affiliation></author><author><name sortKey="Abe, Yukiko" sort="Abe, Yukiko" uniqKey="Abe Y" first="Yukiko" last="Abe">Yukiko Abe</name><affiliation><nlm:aff id="af2-viruses-11-00978">Center for Supercentenarian Medical Research, Shinjuku-ku, Tokyo 160-8582, Japan;<email>ynishimo@z3.keio.jp</email> (Y.N.);<email>yukiko_abe@keio.jp</email> (Y.A.);<email>yasumich@keio.jp</email> (Y.A.)</nlm:aff></affiliation></author><author><name sortKey="Fukuyama, Satoshi" sort="Fukuyama, Satoshi" uniqKey="Fukuyama S" first="Satoshi" last="Fukuyama">Satoshi Fukuyama</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Hamabata, Taiki" sort="Hamabata, Taiki" uniqKey="Hamabata T" first="Taiki" last="Hamabata">Taiki Hamabata</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Okuda, Moe" sort="Okuda, Moe" uniqKey="Okuda M" first="Moe" last="Okuda">Moe Okuda</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Go, Yui" sort="Go, Yui" uniqKey="Go Y" first="Yui" last="Go">Yui Go</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Watanabe, Tokiko" sort="Watanabe, Tokiko" uniqKey="Watanabe T" first="Tokiko" last="Watanabe">Tokiko Watanabe</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Imai, Masaki" sort="Imai, Masaki" uniqKey="Imai M" first="Masaki" last="Imai">Masaki Imai</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Arai, Yasumichi" sort="Arai, Yasumichi" uniqKey="Arai Y" first="Yasumichi" last="Arai">Yasumichi Arai</name><affiliation><nlm:aff id="af2-viruses-11-00978">Center for Supercentenarian Medical Research, Shinjuku-ku, Tokyo 160-8582, Japan;<email>ynishimo@z3.keio.jp</email> (Y.N.);<email>yukiko_abe@keio.jp</email> (Y.A.);<email>yasumich@keio.jp</email> (Y.A.)</nlm:aff></affiliation></author><author><name sortKey="Fouchier, Ron A M" sort="Fouchier, Ron A M" uniqKey="Fouchier R" first="Ron A. M." last="Fouchier">Ron A. M. Fouchier</name><affiliation><nlm:aff id="af3-viruses-11-00978">Department of Viroscience, Erasmus Medical Center, Doctor Molewaterplein 40, 3015 GD Rotterdam, The Netherlands;<email>r.fouchier@erasmusmc.nl</email></nlm:aff></affiliation></author><author><name sortKey="Yamayoshi, Seiya" sort="Yamayoshi, Seiya" uniqKey="Yamayoshi S" first="Seiya" last="Yamayoshi">Seiya Yamayoshi</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Kawaoka, Yoshihiro" sort="Kawaoka, Yoshihiro" uniqKey="Kawaoka Y" first="Yoshihiro" last="Kawaoka">Yoshihiro Kawaoka</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation><affiliation wicri:level="1"><nlm:aff id="af4-viruses-11-00978">Department of Special Pathogens, International Research Center for Infectious Diseases, Institute Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan</nlm:aff><country xml:lang="fr">Japon</country><wicri:regionArea>Department of Special Pathogens, International Research Center for Infectious Diseases, Institute Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639</wicri:regionArea></affiliation><affiliation wicri:level="1"><nlm:aff id="af5-viruses-11-00978">Influenza Research Institute, Department of Pathobiological Sciences, School of Veterinary Medicine, University of Wisconsin-Madison, Madison, WI 53706, USA</nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea>Influenza Research Institute, Department of Pathobiological Sciences, School of Veterinary Medicine, University of Wisconsin-Madison, Madison, WI 53706</wicri:regionArea></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">31652870</idno><idno type="pmc">6893718</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6893718</idno><idno type="RBID">PMC:6893718</idno><idno type="doi">10.3390/v11110978</idno><date when="2019">2019</date><idno type="wicri:Area/Pmc/Corpus">000C49</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000C49</idno><idno type="wicri:Area/Pmc/Curation">000C49</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Curation">000C49</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Antigenic Change in Human Influenza A(H2N2) Viruses Detected by Using Human Plasma from Aged and Younger Adult Individuals</title><author><name sortKey="Matsuzawa, Yukimasa" sort="Matsuzawa, Yukimasa" uniqKey="Matsuzawa Y" first="Yukimasa" last="Matsuzawa">Yukimasa Matsuzawa</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Iwatsuki Horimoto, Kiyoko" sort="Iwatsuki Horimoto, Kiyoko" uniqKey="Iwatsuki Horimoto K" first="Kiyoko" last="Iwatsuki-Horimoto">Kiyoko Iwatsuki-Horimoto</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Nishimoto, Yoshinori" sort="Nishimoto, Yoshinori" uniqKey="Nishimoto Y" first="Yoshinori" last="Nishimoto">Yoshinori Nishimoto</name><affiliation><nlm:aff id="af2-viruses-11-00978">Center for Supercentenarian Medical Research, Shinjuku-ku, Tokyo 160-8582, Japan;<email>ynishimo@z3.keio.jp</email> (Y.N.);<email>yukiko_abe@keio.jp</email> (Y.A.);<email>yasumich@keio.jp</email> (Y.A.)</nlm:aff></affiliation></author><author><name sortKey="Abe, Yukiko" sort="Abe, Yukiko" uniqKey="Abe Y" first="Yukiko" last="Abe">Yukiko Abe</name><affiliation><nlm:aff id="af2-viruses-11-00978">Center for Supercentenarian Medical Research, Shinjuku-ku, Tokyo 160-8582, Japan;<email>ynishimo@z3.keio.jp</email> (Y.N.);<email>yukiko_abe@keio.jp</email> (Y.A.);<email>yasumich@keio.jp</email> (Y.A.)</nlm:aff></affiliation></author><author><name sortKey="Fukuyama, Satoshi" sort="Fukuyama, Satoshi" uniqKey="Fukuyama S" first="Satoshi" last="Fukuyama">Satoshi Fukuyama</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Hamabata, Taiki" sort="Hamabata, Taiki" uniqKey="Hamabata T" first="Taiki" last="Hamabata">Taiki Hamabata</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Okuda, Moe" sort="Okuda, Moe" uniqKey="Okuda M" first="Moe" last="Okuda">Moe Okuda</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Go, Yui" sort="Go, Yui" uniqKey="Go Y" first="Yui" last="Go">Yui Go</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Watanabe, Tokiko" sort="Watanabe, Tokiko" uniqKey="Watanabe T" first="Tokiko" last="Watanabe">Tokiko Watanabe</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Imai, Masaki" sort="Imai, Masaki" uniqKey="Imai M" first="Masaki" last="Imai">Masaki Imai</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Arai, Yasumichi" sort="Arai, Yasumichi" uniqKey="Arai Y" first="Yasumichi" last="Arai">Yasumichi Arai</name><affiliation><nlm:aff id="af2-viruses-11-00978">Center for Supercentenarian Medical Research, Shinjuku-ku, Tokyo 160-8582, Japan;<email>ynishimo@z3.keio.jp</email> (Y.N.);<email>yukiko_abe@keio.jp</email> (Y.A.);<email>yasumich@keio.jp</email> (Y.A.)</nlm:aff></affiliation></author><author><name sortKey="Fouchier, Ron A M" sort="Fouchier, Ron A M" uniqKey="Fouchier R" first="Ron A. M." last="Fouchier">Ron A. M. Fouchier</name><affiliation><nlm:aff id="af3-viruses-11-00978">Department of Viroscience, Erasmus Medical Center, Doctor Molewaterplein 40, 3015 GD Rotterdam, The Netherlands;<email>r.fouchier@erasmusmc.nl</email></nlm:aff></affiliation></author><author><name sortKey="Yamayoshi, Seiya" sort="Yamayoshi, Seiya" uniqKey="Yamayoshi S" first="Seiya" last="Yamayoshi">Seiya Yamayoshi</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation></author><author><name sortKey="Kawaoka, Yoshihiro" sort="Kawaoka, Yoshihiro" uniqKey="Kawaoka Y" first="Yoshihiro" last="Kawaoka">Yoshihiro Kawaoka</name><affiliation><nlm:aff id="af1-viruses-11-00978">Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</nlm:aff></affiliation><affiliation wicri:level="1"><nlm:aff id="af4-viruses-11-00978">Department of Special Pathogens, International Research Center for Infectious Diseases, Institute Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan</nlm:aff><country xml:lang="fr">Japon</country><wicri:regionArea>Department of Special Pathogens, International Research Center for Infectious Diseases, Institute Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639</wicri:regionArea></affiliation><affiliation wicri:level="1"><nlm:aff id="af5-viruses-11-00978">Influenza Research Institute, Department of Pathobiological Sciences, School of Veterinary Medicine, University of Wisconsin-Madison, Madison, WI 53706, USA</nlm:aff><country xml:lang="fr">États-Unis</country><wicri:regionArea>Influenza Research Institute, Department of Pathobiological Sciences, School of Veterinary Medicine, University of Wisconsin-Madison, Madison, WI 53706</wicri:regionArea></affiliation></author></analytic><series><title level="j">Viruses</title><idno type="eISSN">1999-4915</idno><imprint><date when="2019">2019</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Human influenza A(H2N2) viruses emerged in 1957 and were replaced by A(H3N2) viruses in 1968. The antigenicity of human H2N2 viruses has been tested by using ferret antisera or mouse and human monoclonal antibodies. Here, we examined the antigenicity of human H2N2 viruses by using human plasma samples obtained from 50 aged individuals who were born between 1928 and 1933 and from 33 younger adult individuals who were born after 1962. The aged individuals possessed higher neutralization titers against H2N2 viruses isolated in 1957 and 1963 than those against H2N2 viruses isolated in 1968, whereas the younger adults who were born between 1962 and 1968 possessed higher neutralization titers against H2N2 viruses isolated in 1963 than those against other H2N2 viruses. Antigenic cartography revealed the antigenic changes that occurred in human H2N2 viruses during circulation in humans for 11 years, as detected by ferret antisera. These results show that even though aged individuals were likely exposed to more recent H2N2 viruses that are antigenically distinct from the earlier H2N2 viruses, they did not possess high neutralizing antibody titers to the more recent viruses, suggesting immunological imprinting of these individuals with the first H2N2 viruses they encountered and that this immunological imprinting lasts for over 50 years.</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="Noda, T" uniqKey="Noda T">T. Noda</name></author><author><name sortKey="Kawaoka, Y" uniqKey="Kawaoka Y">Y. Kawaoka</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Nabel, G J" uniqKey="Nabel G">G.J. Nabel</name></author><author><name sortKey="Wei, C J" uniqKey="Wei C">C.J. Wei</name></author><author><name sortKey="Ledgerwood, J E" uniqKey="Ledgerwood J">J.E. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Viruses</journal-id><journal-id journal-id-type="iso-abbrev">Viruses</journal-id><journal-id journal-id-type="publisher-id">viruses</journal-id><journal-title-group><journal-title>Viruses</journal-title></journal-title-group><issn pub-type="epub">1999-4915</issn><publisher><publisher-name>MDPI</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">31652870</article-id><article-id pub-id-type="pmc">6893718</article-id><article-id pub-id-type="doi">10.3390/v11110978</article-id><article-id pub-id-type="publisher-id">viruses-11-00978</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>Antigenic Change in Human Influenza A(H2N2) Viruses Detected by Using Human Plasma from Aged and Younger Adult Individuals</article-title></title-group><contrib-group><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-8470-7796</contrib-id><name><surname>Matsuzawa</surname><given-names>Yukimasa</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Iwatsuki-Horimoto</surname><given-names>Kiyoko</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-7062-3057</contrib-id><name><surname>Nishimoto</surname><given-names>Yoshinori</given-names></name><xref ref-type="aff" rid="af2-viruses-11-00978">2</xref></contrib><contrib contrib-type="author"><name><surname>Abe</surname><given-names>Yukiko</given-names></name><xref ref-type="aff" rid="af2-viruses-11-00978">2</xref></contrib><contrib contrib-type="author"><name><surname>Fukuyama</surname><given-names>Satoshi</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Hamabata</surname><given-names>Taiki</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Okuda</surname><given-names>Moe</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Go</surname><given-names>Yui</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Watanabe</surname><given-names>Tokiko</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Imai</surname><given-names>Masaki</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Arai</surname><given-names>Yasumichi</given-names></name><xref ref-type="aff" rid="af2-viruses-11-00978">2</xref></contrib><contrib contrib-type="author"><name><surname>Fouchier</surname><given-names>Ron A.M.</given-names></name><xref ref-type="aff" rid="af3-viruses-11-00978">3</xref></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0001-7768-5157</contrib-id><name><surname>Yamayoshi</surname><given-names>Seiya</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref></contrib><contrib contrib-type="author"><name><surname>Kawaoka</surname><given-names>Yoshihiro</given-names></name><xref ref-type="aff" rid="af1-viruses-11-00978">1</xref><xref ref-type="aff" rid="af4-viruses-11-00978">4</xref><xref ref-type="aff" rid="af5-viruses-11-00978">5</xref><xref rid="c1-viruses-11-00978" ref-type="corresp">*</xref></contrib></contrib-group><aff id="af1-viruses-11-00978"><label>1</label>Division of Virology, Department of Microbiology and Immunology, Institute of Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan;<email>ymatsuza@ims.u-tokyo.ac.jp</email> (Y.M.);<email>fukuyamarry@gmail.com</email> (S.F.);<email>kk176356@stu-cbms.k.u-tokyo.ac.jp</email> (M.O.);<email>yuigo555@gmail.com</email> (Y.G.);<email>tokiko.watanabe@wisc.edu</email> (T.W.);<email>mimai@ims.u-tokyo.ac.jp</email> (M.I.);<email>yamayo@ims.u-tokyo.ac.jp</email> (S.Y.)</aff><aff id="af2-viruses-11-00978"><label>2</label>Center for Supercentenarian Medical Research, Shinjuku-ku, Tokyo 160-8582, Japan;<email>ynishimo@z3.keio.jp</email> (Y.N.);<email>yukiko_abe@keio.jp</email> (Y.A.);<email>yasumich@keio.jp</email> (Y.A.)</aff><aff id="af3-viruses-11-00978"><label>3</label>Department of Viroscience, Erasmus Medical Center, Doctor Molewaterplein 40, 3015 GD Rotterdam, The Netherlands;<email>r.fouchier@erasmusmc.nl</email></aff><aff id="af4-viruses-11-00978"><label>4</label>Department of Special Pathogens, International Research Center for Infectious Diseases, Institute Medical Science, University of Tokyo, Minato-ku, Tokyo 108-8639, Japan</aff><aff id="af5-viruses-11-00978"><label>5</label>Influenza Research Institute, Department of Pathobiological Sciences, School of Veterinary Medicine, University of Wisconsin-Madison, Madison, WI 53706, USA</aff><author-notes><corresp id="c1-viruses-11-00978"><label>*</label>Correspondence: <email>yoshihiro.kawaoka@wisc.edu</email></corresp></author-notes><pub-date pub-type="epub"><day>23</day><month>10</month><year>2019</year></pub-date><pub-date pub-type="collection"><month>11</month><year>2019</year></pub-date><volume>11</volume><issue>11</issue><elocation-id>978</elocation-id><history><date date-type="received"><day>30</day><month>9</month><year>2019</year></date><date date-type="accepted"><day>21</day><month>10</month><year>2019</year></date></history><permissions><copyright-statement>© 2019 by the authors.</copyright-statement><copyright-year>2019</copyright-year><license license-type="open-access"><license-p>Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (<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>Human influenza A(H2N2) viruses emerged in 1957 and were replaced by A(H3N2) viruses in 1968. The antigenicity of human H2N2 viruses has been tested by using ferret antisera or mouse and human monoclonal antibodies. Here, we examined the antigenicity of human H2N2 viruses by using human plasma samples obtained from 50 aged individuals who were born between 1928 and 1933 and from 33 younger adult individuals who were born after 1962. The aged individuals possessed higher neutralization titers against H2N2 viruses isolated in 1957 and 1963 than those against H2N2 viruses isolated in 1968, whereas the younger adults who were born between 1962 and 1968 possessed higher neutralization titers against H2N2 viruses isolated in 1963 than those against other H2N2 viruses. Antigenic cartography revealed the antigenic changes that occurred in human H2N2 viruses during circulation in humans for 11 years, as detected by ferret antisera. These results show that even though aged individuals were likely exposed to more recent H2N2 viruses that are antigenically distinct from the earlier H2N2 viruses, they did not possess high neutralizing antibody titers to the more recent viruses, suggesting immunological imprinting of these individuals with the first H2N2 viruses they encountered and that this immunological imprinting lasts for over 50 years.</p></abstract><kwd-group><kwd>influenza A virus</kwd><kwd>aged individuals</kwd><kwd>H2N2</kwd><kwd>antigenic drift</kwd><kwd>antigenic change</kwd></kwd-group></article-meta></front><floats-group><fig id="viruses-11-00978-f001" orientation="portrait" position="float"><label>Figure 1</label><caption><p>Phylogenetic tree based on the amino acid sequences of HA1 derived from human H2N2 viruses. This tree was constructed by using the Maximum Likelihood method and JTT matrix-based model. Virus isolates used for antigenic analysis are highlighted in magenta.</p></caption><graphic xlink:href="viruses-11-00978-g001"></graphic></fig><fig id="viruses-11-00978-f002" orientation="portrait" position="float"><label>Figure 2</label><caption><p>Neutralization titers of aged individuals (<bold>A</bold>) and younger adult individuals (<bold>B</bold>) against human H2N2 viruses. The neutralization titers against the viruses indicated in <xref rid="viruses-11-00978-t002" ref-type="table">Table 2</xref> were plotted. A non-parametric Friedman’s test followed by a two-stage linear step-up procedure of Benjamini, Krieger, and Yekutieli was performed. * and *** mean <italic>p</italic> < 0.05 and <italic>p</italic> < 0.0001, respectively. (<bold>C</bold>) Bubble chart of the neutralization titers of all of the individuals. The bigger the circle, the higher the neutralization titer.</p></caption><graphic xlink:href="viruses-11-00978-g002"></graphic></fig><fig id="viruses-11-00978-f003" orientation="portrait" position="float"><label>Figure 3</label><caption><p>An antigenic map of human H2N2 viruses. An antigenic map was generated from the neutralization data shown in <xref rid="viruses-11-00978-t002" ref-type="table">Table 2</xref>. Viruses are represented as circles and plasma samples obtained from aged or younger adults are represented as black or grey squares, respectively. Sera with no or only 1 numerical antibody titer are not shown as they cannot be placed properly in an antigenic map. The grid indicates one unit of antigenic distance, a two-fold dilution in neutralization titer.</p></caption><graphic xlink:href="viruses-11-00978-g003"></graphic></fig><table-wrap id="viruses-11-00978-t001" orientation="portrait" position="float"><object-id pub-id-type="pii">viruses-11-00978-t001_Table 1</object-id><label>Table 1</label><caption><p>Characteristics of the participants in this study.</p></caption><table frame="hsides" rules="groups"><thead><tr><th rowspan="4" colspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin"></th><th colspan="10" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">No. of Individuals</th></tr><tr><th colspan="10" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1">Birth Year</th></tr><tr><th colspan="6" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1">Aged Individuals</th><th colspan="4" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1">Younger Adult Individuals</th></tr><tr><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1928</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1929</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1930</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1931</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1932</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1933</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1960s</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1970s</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1980s</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1990s</th></tr></thead><tbody><tr><td colspan="2" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1">Males</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">10</td><td align="center" valign="middle" rowspan="1" colspan="1">5</td><td align="center" valign="middle" rowspan="1" colspan="1">1</td><td align="center" valign="middle" rowspan="1" colspan="1">2</td></tr><tr><td colspan="2" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1">Females</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">5</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">5</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1</td></tr><tr><td colspan="2" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1">Total</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">19</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3</td></tr><tr><td rowspan="3" align="center" valign="middle" style="border-bottom:solid thin" colspan="1">Medical history <sup>b</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">Heart disease</td><td align="center" valign="middle" rowspan="1" colspan="1">3/9</td><td align="center" valign="middle" rowspan="1" colspan="1">1/9</td><td align="center" valign="middle" rowspan="1" colspan="1">3/8</td><td align="center" valign="middle" rowspan="1" colspan="1">1/8</td><td align="center" valign="middle" rowspan="1" colspan="1">1/8</td><td align="center" valign="middle" rowspan="1" colspan="1">1/8</td><td align="center" valign="middle" rowspan="1" colspan="1">NA <sup>a</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" rowspan="1" colspan="1">NA</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">Diabetes</td><td align="center" valign="middle" rowspan="1" colspan="1">0/9</td><td align="center" valign="middle" rowspan="1" colspan="1">0/9</td><td align="center" valign="middle" rowspan="1" colspan="1">1/8</td><td align="center" valign="middle" rowspan="1" colspan="1">2/8</td><td align="center" valign="middle" rowspan="1" colspan="1">1/8</td><td align="center" valign="middle" rowspan="1" colspan="1">1/8</td><td align="center" valign="middle" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" rowspan="1" colspan="1">NA</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Cancer</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3/9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2/9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2/8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">4/8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2/8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2/8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NA</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NA</td></tr></tbody></table><table-wrap-foot><fn><p><sup>a</sup> Not available. <sup>b</sup> Summary of available medical history of each individual.</p></fn></table-wrap-foot></table-wrap><table-wrap id="viruses-11-00978-t002" orientation="portrait" position="float"><object-id pub-id-type="pii">viruses-11-00978-t002_Table 2</object-id><label>Table 2</label><caption><p>Neutralization titers of aged and younger adult individuals.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">ID</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Birth Year (age)</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">M1/57 <sup>a</sup></th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">K1/63 <sup>b</sup></th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">B1/68 <sup>c</sup></th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">B2/68 <sup>d</sup></th></tr></thead><tbody><tr><td align="center" valign="middle" rowspan="1" colspan="1">1</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">2</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">3</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">5</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">6</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">7</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">9</td><td align="center" valign="middle" rowspan="1" colspan="1">1928 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">10</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">11</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">12</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">13</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">14</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">15</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (89)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">17</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (88)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">18</td><td align="center" valign="middle" rowspan="1" colspan="1">1929 (88)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">19</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (88)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">20</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (88)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">21</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (88)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">22</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (88)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">23</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">24</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">25</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">26</td><td align="center" valign="middle" rowspan="1" colspan="1">1930 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">27</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">28</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">29</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">30</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (87)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">31</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">33</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">34</td><td align="center" valign="middle" rowspan="1" colspan="1">1931 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">35</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">36</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">37</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">38</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">39</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (86)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">40</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">41</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">42</td><td align="center" valign="middle" rowspan="1" colspan="1">1932 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">43</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">44</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">45</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">46</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">47</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">48</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">>512</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">49</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">50</td><td align="center" valign="middle" rowspan="1" colspan="1">1933 (85)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">51</td><td align="center" valign="middle" rowspan="1" colspan="1">1962 (56)</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">52</td><td align="center" valign="middle" rowspan="1" colspan="1">1962 (55)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">53</td><td align="center" valign="middle" rowspan="1" colspan="1">1962 (55)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">54</td><td align="center" valign="middle" rowspan="1" colspan="1">1962 (55)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">55</td><td align="center" valign="middle" rowspan="1" colspan="1">1963 (55)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">>512</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">56</td><td align="center" valign="middle" rowspan="1" colspan="1">1963 (55)</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">>512</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">57</td><td align="center" valign="middle" rowspan="1" colspan="1">1963 (55)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">58</td><td align="center" valign="middle" rowspan="1" colspan="1">1963 (55)</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">256</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">59</td><td align="center" valign="middle" rowspan="1" colspan="1">1964 (54)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">60</td><td align="center" valign="middle" rowspan="1" colspan="1">1965 (53)</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">61</td><td align="center" valign="middle" rowspan="1" colspan="1">1966 (52)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">62</td><td align="center" valign="middle" rowspan="1" colspan="1">1966 (51)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">128</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">63</td><td align="center" valign="middle" rowspan="1" colspan="1">1967 (51)</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">64</td><td align="center" valign="middle" rowspan="1" colspan="1">1967 (50)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">65</td><td align="center" valign="middle" rowspan="1" colspan="1">1968 (50)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">66</td><td align="center" valign="middle" rowspan="1" colspan="1">1968 (49)</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">67</td><td align="center" valign="middle" rowspan="1" colspan="1">1969 (49)</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">68</td><td align="center" valign="middle" rowspan="1" colspan="1">1969 (49)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">69</td><td align="center" valign="middle" rowspan="1" colspan="1">1969 (48)</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">70</td><td align="center" valign="middle" rowspan="1" colspan="1">1970 (48)</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">71</td><td align="center" valign="middle" rowspan="1" colspan="1">1970 (47)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1">32</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">72</td><td align="center" valign="middle" rowspan="1" colspan="1">1970 (47)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">73</td><td align="center" valign="middle" rowspan="1" colspan="1">1970 (47)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">16</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">74</td><td align="center" valign="middle" rowspan="1" colspan="1">1970 (47)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">75</td><td align="center" valign="middle" rowspan="1" colspan="1">1972 (45)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">76</td><td align="center" valign="middle" rowspan="1" colspan="1">1974 (44)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">77</td><td align="center" valign="middle" rowspan="1" colspan="1">1978 (39)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">78</td><td align="center" valign="middle" rowspan="1" colspan="1">1980 (37)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">79</td><td align="center" valign="middle" rowspan="1" colspan="1">1982 (36)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">80</td><td align="center" valign="middle" rowspan="1" colspan="1">1985 (33)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">81</td><td align="center" valign="middle" rowspan="1" colspan="1">1990 (27)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">82</td><td align="center" valign="middle" rowspan="1" colspan="1">1992 (26)</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" rowspan="1" colspan="1">4</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">83</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1994 (23)</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">4</td></tr></tbody></table><table-wrap-foot><fn><p>Neutralization titers of plasma samples of aged individuals (ID 1–50) and younger adult individuals (ID 51–83) against A/Netherland/M1/57 <sup>a</sup>, A/Netherland/K1/63 <sup>b</sup>, A/Netherland/B1/68 <sup>c</sup>, or A/Netherland/B2/68 <sup>d</sup> were measured.</p></fn></table-wrap-foot></table-wrap><table-wrap id="viruses-11-00978-t003" orientation="portrait" position="float"><object-id pub-id-type="pii">viruses-11-00978-t003_Table 3</object-id><label>Table 3</label><caption><p>Comparison of amino acids that are important for HA antigenicity <sup>a.</sup></p></caption><table frame="hsides" rules="groups"><thead><tr><th rowspan="2" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" colspan="1">Isolate</th><th colspan="7" align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1">Amino Acid Position at</th></tr><tr><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">126</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">128</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">132</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">139</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">154</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">184</th><th align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">188</th></tr></thead><tbody><tr><td align="center" valign="middle" rowspan="1" colspan="1">A/Netherland/M1/57</td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">R</td><td align="center" valign="middle" rowspan="1" colspan="1">N</td><td align="center" valign="middle" rowspan="1" colspan="1">S</td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">T</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">A/Netherland/K1/63</td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">R</td><td align="center" valign="middle" rowspan="1" colspan="1">N</td><td align="center" valign="middle" rowspan="1" colspan="1">P</td><td align="center" valign="middle" rowspan="1" colspan="1">A</td><td align="center" valign="middle" rowspan="1" colspan="1">A</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">A/Netherland/B1/68</td><td align="center" valign="middle" rowspan="1" colspan="1">E</td><td align="center" valign="middle" rowspan="1" colspan="1">D</td><td align="center" valign="middle" rowspan="1" colspan="1">K</td><td align="center" valign="middle" rowspan="1" colspan="1">K</td><td align="center" valign="middle" rowspan="1" colspan="1">P</td><td align="center" valign="middle" rowspan="1" colspan="1">A</td><td align="center" valign="middle" rowspan="1" colspan="1">A</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">A/Netherland/B2/68</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">K</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">D</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">K</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">K</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Q</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">E</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">A</td></tr></tbody></table><table-wrap-foot><fn><p><sup>a</sup> Amino acids that are different among the HAs of the four isolates at previously identified antigenic determinants [<xref rid="B11-viruses-11-00978" ref-type="bibr">11</xref>] are listed.</p></fn></table-wrap-foot></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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