[1] Univ Tokyo, Grad Sch Arts & Sci, Dept Life Sci, Meguro Ku, Tokyo 1538902, Japan
[2] Univ Tokyo, Inst Mol & Cellular Biosci, Bunkyo Ku, Tokyo 1130032, Japan
[3] RIKEN, Brain Sci Inst, Lab Proteolyt Neurosci, Wako, Saitama 3510198, Japan
[4] Tokyo Metropolitan Inst Psychiat, Dept Biol Mol, Setagaya Ku, Tokyo 1568585, Japan
[5] Tsukuba Technopk Oho, Banyu Tsukuba Res Inst, Tsukuba, Ibaraki 3002611, Japan
[6] Tokyo Metropolitan Inst Med Sci, Dept Cell Biol, Bunkyo Ku, Tokyo 1138613, Japan
[7] Yokohama City Univ, Sch Med, Dept Biol Mol, Kanazawa Ku, Kanagawa 2360004, Japan