1. Graduate School of Medicine, The University of Tokyo 1 Department of Neuropathology , , Tokyo 113-0033 , Japan
2. Graduate School of Life Sciences, Tohoku University 2 Laboratory of Membrane Trafficking Mechanisms, Department of Integrative Life Sciences , , Sendai 980-8578 , Japan
3. Graduate School and Faculty of Medicine, The University of Tokyo 3 Department of Biochemistry and Molecular Biology , , Tokyo 113-0033 , Japan