[1] Univ Cambridge, Dept Physiol Dev & Neurosci, Cambridge CB2 3EG, England
[2] Univ Cambridge, Dept Med, Cambridge CB2 3EG, England
[3] Natl Hlth Serv Fdn Trust, Papworth Hosp, Cambridge CB23 3RE, England
[4] Univ Calif San Diego, Dept Pediat, Div Biol Sci, La Jolla, CA 92093 USA
[5] Univ Calif San Diego, Skaggs Sch Pharm & Pharmaceut Sci, La Jolla, CA 92093 USA