[1] Washington State Univ, Dept Bot, Pullman, WA 99164 USA
[2] Australian Natl Univ, Div Plant Ind, Canberra, ACT 2601, Australia
[3] Univ Lancaster, Dept Biol Sci, Lancaster LA1 4YQ, England
[4] Univ Sheffield, Dept Anim & Plant Sci, Robert Hill Inst, Sheffield S10 2TN, S Yorkshire, England