1. Department of Organismic and Evolutionary Biology, Harvard University , Cambridge, MA 02138, USA
2. Department of Physics, Harvard University , Cambridge, MA 02138, USA
3. Quantitative Biology Initiative, Harvard University , Cambridge, MA 02138, USA
4. NSF-Simons Center for Mathematical and Statistical Analysis of Biology, Harvard University , Cambridge, MA 02138, USA
5. Department of Applied Physics, Stanford University , Stanford, CA 94305, USA
6. Department of Bioengineering, Stanford University , Stanford, CA 94305, USA