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Acknowledgements
We thank Mariko Kurakata for her assistance with cell culture. We thank Ikue Hiraga
and the Laboratory of Radioisotope Research, Research Support Center, Juntendo University Graduate School of Medicine, for technical assistance. We also thank the staff
scientists at the University of Tokyo’s cryo-EM facility. This study was partly supported
by the Japan Society for the Promotion of Sciences KAKENHI (grant numbers 19K07105
and 22K06652 to N.K., 19H03404 and 22H02805 to T.M. and JP16H04748 and
21H02411 to H.O.); the Platform Project for Supporting Drug Discovery and Life Science
Research (Basis for Supporting Innovative Drug Discovery and Life Science Research
[BINDS]; grant number JP20am0101080 to H.O. and T.M. and JP19am0101115 (support
number 0064)); the Practical Research Project for Rare/Intractable Diseases from the
Japan Agency for Medical Research and Development (AMED; grant number
19ek0109202) to N.K.); an Intramural Research Grant (2–5) for Neurological and Psychiatric Disorders of NCNP (to T.M.); and the Vehicle Racing Commemorative Foundation (6114, 6237, and 6303) to T.M. and H.O.
Author contributions
T.K., N.K., T.M., and H.O. conceived and designed the project. M. Kodama and H.O.
performed cell culture. T.K. performed protein purification. K.S. performed negativestaining EM studies. A.T. and T.K. prepared the grid for cryo-EM. A.T. and M. Kikkawa
processed the images. H.O. performed model building and refinement. T.M. and N.K.
performed the functional analysis. T.K., A.T., N.K., K.S., M. Kodama, T.S., M. Kikkawa,
T.M., and H.O. interpreted the data. T.M. and H.O. supervised the project. H.O. and
T.M. wrote the manuscript with input from all authors. All authors reviewed the results
and approved the final version of the manuscript.
Competing interests
The authors declare no competing interests.
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Additional information
Supplementary information The online version contains supplementary material
available at https://doi.org/10.1038/s41467-022-30429-x.
Correspondence and requests for materials should be addressed to Takashi Murayama
or Haruo Ogawa.
Peer review information Nature Communications thanks Huan Li and Montserrat
Samso for their contribution to the peer review of this work. Peer reviewer reports are
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