[1] Hokkaido Univ, Grad Sch Med, Inst Anim Expt, Kita Ku, Sapporo, Hokkaido 0608638, Japan
[2] Natl Inst Infect Dis, Dept Virol 1, Musashimurayama 2080011, Japan
[3] Iwate Univ, Fac Agr, Vet Physiol Lab, Morioka, Iwate 0208550, Japan
[4] Hokkaido Univ, Grad Sch Vet Med, Dept Environm Vet Sci, Publ Hlth Lab, Sapporo, Hokkaido 0600818, Japan