1. College of Health, Medicine and Life Sciences, Department of Life Sciences, Brunel University London 1 , London UB8 3PH , UK
2. Wellcome Centre for Cell Biology, Institute of Cell Biology, University of Edinburgh 2 , Edinburgh EH16 4SB , UK
3. Institute for Genetic Medicine, Hokkaido University 3 , Kita-15, Nishi-7, Kita-Ku, Sapporo 060-0815 , Japan