[1] Univ Tokyo, Inst Med Sci, Dept Microbiol & Immunol, Div Virol,Minato Ku, Tokyo 1088639, Japan
[2] Univ Tokyo, Inst Med Sci, Fine Morphol Lab, Dept Basic Med Sci,Minato Ku, Tokyo 1088639, Japan
[3] Hokkaido Univ, Grad Sch Vet Med, Dept Dis Control, Microbiol Lab, Sapporo, Hokkaido 0600818, Japan
[4] Univ Wisconsin, Sch Vet Med, Dept Pathobiol Sci, Madison, WI 53706 USA