1. John A. Paulson School of Engineering and Applied Sciences, Harvard University 1 , Cambridge, MA 02138 , USA
2. Institute of Molecular and Cell Biology, A*STAR 2 , Singapore 138673
3. Brigham Women's Hospital and Department of Genetics, Harvard Medical School 3 Department of Pathology , , Boston, MA 02115 , USA
4. Wellcome Trust/CRUK Gurdon Institute, University of Cambridge 4 , Cambridge CB2 1QN , UK