[1] Queen Mary Univ London, William Harvey Res Inst, London EC1M 6BQ, England
[2] Natl Inst Med Res, MRC, Div Mol Neurobiol, London NW7 1AA, England
[3] Nanyang Technol Univ, Lee Kong Chian Sch Med, Singapore 639798, Singapore
[4] Nanyang Technol Univ, Sch Biol Sci, Singapore 639798, Singapore
[5] Karolinska Inst, Dept Microbiol Tumor & Cell Biol, S-17177 Stockholm, Sweden
[6] Hubrecht Inst KNAW, NL-3584 CT Utrecht, Netherlands
[7] Univ Med Ctr Utrecht, NL-3584 CT Utrecht, Netherlands