[1] Univ Cambridge, Sainsbury Lab, 47 Bateman St, Cambridge CB2 1LR, England
[2] Univ Cambridge, Dept Genet, Downing St, Cambridge CB2 3EH, England
[3] Univ Cambridge, Dept Appl Math & Theoret Phys, Cambridge CB3 0WA, England
[4] Lund Univ, Dept Astron & Theoret Phys, Computat Biol & Biol Phys, Lund, Sweden