About the Authors

Fumio Yoshikawa

Contributed equally to this work with: Fumio Yoshikawa, Yumi Sato

Current address: Yoshikawa Hospital, Izumiotsu, Osaka, 595–0061, Japan

Affiliation Laboratory for Molecular Neurogenesis, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan

Yumi Sato

Contributed equally to this work with: Fumio Yoshikawa, Yumi Sato

Current address: Graduate School of Medical Science, Division of Molecular Virology and Oncology, Cancer Research Institute, Kanazawa University, Kanazawa, Ishikawa, 920–1192, Japan

Affiliation Laboratory for Molecular Neurogenesis, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan

Koujiro Tohyama

Affiliation The Center for Electron Microscopy and Bio-Imaging Research and Department of Physiology, Iwate Medical University, Morioka, Iwate, 020–8505, Japan

Takumi Akagi

Current address: Department of Physiology, Nippon Medical School, Tokyo, 113–8602, Japan

Affiliation Support Unit for Neuromorphological Analysis, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan

Tamio Furuse

Affiliation Technology and Development Team for Mouse Phenotype Analysis, RIKEN BioResource Center, Tsukuba, Ibaraki, 305–0074, Japan

Tetsushi Sadakata

Affiliations Laboratory for Molecular Neurogenesis, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan, Advanced Scientific Research Leaders Development Unit, Gunma University, Maebashi, Gunma, 371–8511, Japan

Mika Tanaka

Affiliation Laboratory for Behavioral Genetics, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan

Yo Shinoda

Affiliations Laboratory for Molecular Neurogenesis, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan, School of Pharmacy, Tokyo University of Pharmacy and Life Sciences, Hachioji, Tokyo, 192–0392, Japan

Tsutomu Hashikawa

Affiliation Support Unit for Neuromorphological Analysis, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan

Shigeyoshi Itohara

Affiliation Laboratory for Behavioral Genetics, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan

Yoshitake Sano

Affiliation Department of Applied Biological Science, Faculty of Science and Technology, Tokyo University of Science, Chiba, 278–8510, Japan

M. Said Ghandour

Affiliation Unite´ Mixte de Recherche 7357, Université de Strasbourg, Strasbourg, 67085, France and Department of Anatomy and Neurobiology, Virginia Commonwealth University, Richmond, Virginia, United States of America

Shigeharu Wakana

Affiliation Technology and Development Team for Mouse Phenotype Analysis, RIKEN BioResource Center, Tsukuba, Ibaraki, 305–0074, Japan

Teiichi Furuichi

tfuruichi@rs.tus.ac.jp

Affiliations Laboratory for Molecular Neurogenesis, RIKEN Brain Science Institute, Wako, Saitama, 351–0198, Japan, Department of Applied Biological Science, Faculty of Science and Technology, Tokyo University of Science, Chiba, 278–8510, Japan

Competing Interests

The authors have declared that no competing interest exist.

Author Contributions

Conceptualization: Y. Sato FY T. Furuichi. Data curation: KT TH SI MSG SW. Formal analysis: Y. Sato Y. Sano T. Furuse. Funding acquisition: KT TH SI MSG SW T. Furuichi. Investigation: Y. Sato FY TA MT T. Furuse. Project administration: KT TH SI MSG SW T. Furuichi. Supervision: KT TH SI MSG SW T. Furuichi. Writing – original draft: Y. Sato T. Furuichi. Writing – review & editing: TS Y. Shinoda Y. Sano MSG.