1. Department of Biophysics and Biochemistry, Graduate School of Science, University of Tokyo, Bunkyo, 113-0032, Japan
2. Neuroscience Research Institute, University of California-Santa Barbara, Santa Barbara, CA 93106, USA
3. PRESTO, Japan Science and Technology Corporation, Kawaguchi, 332-0012, Japan
4. Laboratory for Developmental Genomics, RIKEN Center for Developmental Biology, Kobe, 650-0047 Japan