About the Authors

H. Akiko Popiel

Affiliation Department of Degenerative Neurological Diseases, National Institute of Neuroscience, National Center of Neurology and Psychiatry, Kodaira, Tokyo, Japan

Toshihide Takeuchi

Affiliation Department of Degenerative Neurological Diseases, National Institute of Neuroscience, National Center of Neurology and Psychiatry, Kodaira, Tokyo, Japan

Hiromi Fujita

Affiliation Department of Degenerative Neurological Diseases, National Institute of Neuroscience, National Center of Neurology and Psychiatry, Kodaira, Tokyo, Japan

Kazuhiro Yamamoto

Affiliation Division of Laboratory Animals Resources, National Institute of Neuroscience, National Center of Neurology and Psychiatry, Kodaira, Tokyo, Japan

Chiyomi Ito

Affiliation Division of Neurology/Molecular Brain Science, Kobe University Graduate School of Medicine, Kobe, Hyogo, Japan

Hiroshi Yamane

Affiliation Department of Degenerative Neurological Diseases, National Institute of Neuroscience, National Center of Neurology and Psychiatry, Kodaira, Tokyo, Japan

Shin-ichi Muramatsu

Affiliation Division of Neurology, Department of Medicine, Jichi Medical University, Shimotsuke, Tochigi, Japan

Tatsushi Toda

Affiliation Division of Neurology/Molecular Brain Science, Kobe University Graduate School of Medicine, Kobe, Hyogo, Japan

Keiji Wada

Affiliation Department of Degenerative Neurological Diseases, National Institute of Neuroscience, National Center of Neurology and Psychiatry, Kodaira, Tokyo, Japan

Yoshitaka Nagai

nagai@ncnp.go.jp

Affiliations Department of Degenerative Neurological Diseases, National Institute of Neuroscience, National Center of Neurology and Psychiatry, Kodaira, Tokyo, Japan, Core Research for Evolutional Science and Technology (CREST), Japan Science and Technology Agency, Kawaguchi, Saitama, Japan

Competing Interests

Co-author Yoshitaka Nagai is a PLOS ONE Editorial Board member. This does not alter the authors' adherence to all the PLOS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: HAP T. Takeuchi SM T. Toda KW YN. Performed the experiments: HAP T. Takeuchi HF KY CI HY. Analyzed the data: HAP T. Takeuchi T. Toda KW YN. Contributed reagents/materials/analysis tools: SM. Wrote the paper: HAP T. Takeuchi YN.