[1] Hokkaido Univ, Grad Sch Med, Dept Ophthalmol & Visual Sci, Kita Ku, Sapporo, Hokkaido 0608638, Japan
[2] Hokkaido Univ, Grad Sch Med, Dept Biochem, Kita Ku, Sapporo, Hokkaido 0608638, Japan
[3] Tokyo Metropolitan Inst Neurosci, Dept Mol Neurobiol, Fuchu, Tokyo 1838526, Japan