[1] Natl Inst Med Res, Div Dev Biol, London NW7 1AA, England
[2] Univ Cambridge, Wellcome Trust Canc Res UK Inst, Cambridge CB2 1QR, England
[3] Univ Cambridge, Dept Zool, Cambridge CB2 1QR, England
[4] Univ Illinois, Dept Mol Genet, Chicago, IL 60607 USA