This work was partially supported by grants from the World Premier International Research Center Initiative from MEXT, JST CREST grant #JPMJCR1655, JSPS KAKENHI grants #16K18359, 15F15408, 15H01276, 15K18332, 26115502, 25116530, JP16H06280, and 20H03552, Takeda Science Foundation, Shimadzu Science Foundation, Kanae Foundation, Research Foundation for Opto-Science and Technology, Ichiro Kanehara Foundation, Kato Memorial Bioscience Foundation, Japan Foundation for Applied Enzymology, Senshin Medical Research Foundation, Life Science Foundation of Japan, Uehara Memorial Foundation, Brain Science Foundation, Kowa Life Science Foundation, Inamori Research Grants Program, and GSK Japan to MS.