[1] Harvard Univ, Dept Phys, Cambridge, MA 02138 USA
[2] Harvard Univ, FAS Ctr Syst Biol, Cambridge, MA 02138 USA
[3] Simons Fdn, Ctr Computat Biol, New York, NY USA
[4] Harvard Univ, John A Paulson Sch Engn & Appl Sci, Cambridge, MA 02138 USA
[5] NYU, Courant Inst Math Sci, New York, NY USA
[6] Harvard Univ, Dept Mol & Cellular Biol, Cambridge, MA 02138 USA