1. Department of Biological Sciences, Graduate School of Science, University of Tokyo, Hongo, Tokyo 113-0033, Japan
2. Division of Biological Science, Graduate School of Science, Nagoya University, Chikusa-ku, Nagoya 464-8602, Japan
3. Department of Integrated Biosciences, Graduate School of Frontier Sciences, University of Tokyo, Hongo, Tokyo 113-0033, Japan
4. Department of Cell Biology, John Innes Centre, Colney, Norwich NR4 7UH, United Kingdom