1. MRC Cell Biology Unit, Laboratory of Molecular Cell Biology, University College London, Gower Street, London, WC1E6BT, UK
2. Present address - Cell Biology Laboratories, School of Biochemistry, Medical Sciences Building, University of Bristol, Bristol, BS81TD, UK
3. William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, Charterhouse Square, London EC1M6BQ, UK