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Acknowledgements

We would like to thank all the lab members of the Molecular Neuroscience Research Center (MNRC) for the

helpful discussions and sincere cooperation. This research was supported by the Central Research Laboratory

at Shiga University of medical science (SUMS). M.M. is supported by research grants from the Kato Memorial

Bioscience Foundation, the Japan Epilepsy Research Foundation (JERF), the Hoansha Foundation, MSD Life

Science Foundation, the Ichiro Kanehara Foundation for the Promotion of Medical Sciences and Medical Care,

Japan Brain Foundation, Takeda Science Foundation and the Japan Spina Bifida and Hydrocephalus Research

Foundation. This study was supported Grants-in-Aid for Scientific Research for Young Scientists from the Japan

Intractable diseases (Nanbyo) Research Foundation. This study was supported by MEXT KAKENHI Grant

Number 15K15387, JSPS KAKENHI Grant Numbers 15H01486, 18K07788, and 19H04774, the Leading Initiative

for Excellent Young Researchers (LEADER) 5013323 and Initiative for Rare and Undiagnosed Diseases (IRUD)

of AMED.

Author contributions

F.A.J and M.M. conceived the project and designed experiments. F.A.J, T.M. and A.Ts. conducted most of the

experiments. A.Ta. and Y.T. conducted gene expression analysis and assisted cell culture experiments. Y.M.

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performed flow cytometer experiments. T.Y. assisted confocal imaging and time-lapse video recordings. M.N.

and I.T. assisted in the data analysis and discussed the results with M.M., F.A.J. and M.M. conceived the experiment. F.A.J. and M.M. and wrote the manuscript. All authors reviewed the manuscript.

Competing interests The authors declare no competing interests.

Additional information

Supplementary information is available for this paper at https​://doi.org/10.1038/s4159​8-020-74874​-4.

Correspondence and requests for materials should be addressed to M.M.

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