[1] Univ Durham, Sch Biol & Biomed Sci, Integrat Cell Biol Lab, Durham DH1 3LE, England
[2] Swedish Univ Agr Sci, Umea Plant Sci Ctr, S-090183 Umea, Sweden
[3] Rothamsted Res, Harpenden AL5 2JQ, Herts, England
[4] Salk Inst Biol Studies, Carlsbad, CA 92009 USA