[1] Macaulay Inst, Aberdeen AB15 8QH, Scotland
[2] BioEMCo, Ctr INRA INA PG, CNRS, F-78850 Thiverval Grignon, France
[3] Univ York, Dept Biol, York YO10 5YW, N Yorkshire, England
[4] Biomath & Stat Scotland, Dundee DD2 5DA, Scotland
[5] Scottish Crop Res Inst, Dundee DD2 5DA, Scotland
[6] Univ Aberdeen, Sch Biol Sci, Aberdeen AB24 3UU, Scotland