[1] Univ Tokyo, Inst Med Sci, Dept Microbiol & Immunol, Div Mucosal Immunol, Tokyo 1088639, Japan
[2] Japan Sci & Technol Agcy JST, CREST, Tokyo, Japan
[3] Univ Tokyo, Inst Med Sci, Dept Microbiol & Immunol, Div Infect Genet, Tokyo 1088639, Japan
[4] Columbia Univ, Med Ctr, Dept Microbiol & Immunol, New York, NY USA
[5] Univ Tokyo, Inst Med Sci, Int Res & Dev Ctr Mucosal Vaccine, Tokyo 1088639, Japan