[1] Univ Toronto, Dept Cell & Syst Biol, Toronto, ON M5S 2B2, Canada
[2] Univ Toronto, CAGEF, Toronto, ON M5S 3B2, Canada
[3] RIKEN Plant Sci Ctr, Yokohama, Kanagawa 2300045, Japan
[4] Natl Inst Basic Biol, Lab Plant Organ Dev, Okazaki 4448585, Japan