[1] Kyoto Univ, Inst Virus Res, Lab Viral Pathogenesis, Kyoto 6068507, Japan
[2] Kyoto Univ, Inst Virus Res, Lab Viral Control, Kyoto 6068507, Japan
[3] Toyama Med & Pharmaceut Univ, Lab Anim Res Ctr, Toyama 9300152, Japan
[4] Kyoto Univ, Coll Med Technol, Kyoto 6068507, Japan
[5] Nippon Med Coll, Dept Microbiol & Immunol, Tokyo 1138602, Japan