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Acknowledgements The authors thank the research resource division (RRD) (RIKEN Center for Brain Science) and RIKEN CBS-EVIDENT Open Collaboration Center (RIKEN Center for Brain Science) for technical assistance, Yukiko Nagai (RIKEN Center for Brain Science) for secretarial work, Kenichi Nagata (Nagoya University) and members of the Laboratory for Proteolytic Neuroscience (RIKEN Center for Brain Science) for beneficial suggestions