1. Department of Physics and Astronomy, Vrije Universiteit Amsterdam 1 , 1081HV Amsterdam, The Netherlands
2. Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital 2 , Toronto, Ontario M5G 1X5, Canada
3. Wellcome/CRUK Gurdon Institute, University of Cambridge 3 , Cambridge CB2 1QN, United Kingdom
4. Biological Physics Theory Unit, OIST Graduate University 4 , Onna-son, Okinawa 904-0495, Japan