1. University of Oxford, Weatherall Institute of Molecular Medicine 1 , Radcliffe Department of Medicine, Oxford OX3 9DS , UK
2. University of Oxford, Institute of Developmental and Regenerative Medicine 2 , Department of Physiology, Anatomy and Genetics, Oxford OX3 7DQ , UK
3. University College London, Department of Cell & Developmental Biology 3 , London WC1E 6BT , UK
4. Stowers Institute for Medical Research 4 , Kansas City, MO 64110 , USA