[1] Univ Cambridge, Dept Physiol Dev & Neurosci, Downing St, Cambridge CB2 3DY, England
[2] Univ Warwick, Warwick Med Sch, Gibbet Hill Rd, Coventry CV4 7AL, England
[3] Univ Manchester, Core Technol Facil, Fac Biol Med & Hlth, 46 Grafton St, Manchester M13 9NT, England
[4] Univ Cambridge, Gurdon Inst, Tennis Court Rd, Cambridge CB2 1QN, England