1. Tokyo Gakugei University 1 Department of Biology , , Koganei-shi, Tokyo 184-8501 , Japan
2. Université de Lyon, ENS de Lyon, UCBL, INRAE, CNRS 2 Laboratoire de Reproduction et Développement des Plantes , , 46 Allée d'Italie, 69364 Lyon Cedex 07 , France
3. Bioproduction Research Institute, National Institute of Advanced Industrial Science and Technology (AIST) 3 Plant Gene Regulation Research Group , , Higashi 1-1-1, Tsukuba, Ibaraki 305-8566 , Japan
4. Graduate School of Science, The University of Tokyo 4 Department of Biological Sciences , , Tokyo 113-0033 , Japan
5. International Research Center for Agriculture and Environmental Biology, Kumamoto University 5 , Kumamoto 860-8555 , Japan