* Corresponding Author; email: [email protected]

1RIKEN Center for Brain Science, Wako, Japan

2Institute for Frontier Life and Medical Sciences, Kyoto University, Shogoin-Kawahara, Sakyo-ku, Kyoto, Japan

3Kyoto University Graduate School of Medicine, Kyoto, Japan

4Institute for Integrated Cell-Material Sciences, Kyoto University, Kyoto, Japan