[1] Univ Bristol, Integrat Epidemiol Unit, Populat Hlth Sci, Bristol BS8 1TH, Avon, England
[2] UCL, Genet Inst UGI, London WC1E 6BT, England
[3] UCL, Ctr Math & Phys Life Sci & Expt Biol CoMPLEX, London WC1E 6BT, England
[4] Univ Bath, Milner Ctr Evolut, Bath BA2 7AY, Avon, England