[1] Univ Southampton, Bioengn Sci Res Grp, Dept Mech Engn, Sch Engn,Fac Engn & Phys Sci, Southampton SO17 1BJ, Hants, England
[2] Univ Aberdeen, Sch Biol Sci, Aberdeen AB24 3UU, Scotland
[3] James Hutton Inst, Ecol Sci Grp, Dundee DD2 5DA, Scotland
[4] Univ Dundee, Sch Sci & Engn, Dundee DD1 4HN, Scotland
[5] Martin Luther Univ Halle Wittenberg, Inst Agr & Nutr Sci, Univ Pl 10, D-06108 Halle, Saale, Germany