[1] Teikyo Univ, Dept Biosci, Utsunomiya, Tochigi 3208551, Japan
[2] Aberystwyth Univ, Inst Biol Sci, Aberystwyth SY23 3DA, Dyfed, Wales
[3] Ehime Univ, Fac Sci, Dept Biol & Earth Sci, Matsuyama, Ehime 7908577, Japan
[4] Univ Tasmania, Sch Plant Sci, Hobart, Tas 7001, Australia
[5] Salk Inst Biol Studies, Plant Biol Lab, La Jolla, CA 92037 USA