1. MRC-National Institute for Medical Research, The Ridgeway, Mill Hill, London NW7 1AA, UK
2. Department of Mathematics and CoMPLEX, University College London, Gower Street, London WC1E 6BT, UK
3. Department of Cell and Developmental Biology and Department of Genetics, Evolution and Environment, University College London, Gower Street, London WC1E 6BT, UK