1. Oberlin College 1 Biology Department , , Oberlin, OH 44074 USA
2. Cluster of Plant Developmental Biology, Laboratory of Molecular Biology, Wageningen University & Research, Droevendaalsesteeg 1 2 , 6708 PB Wageningen , The Netherlands
3. University of Massachusetts 3 Department of Biology , , Amherst, MA 01003 , USA
4. Sainsbury Laboratory, University of Cambridge 4 , Cambridge, CB2 1LR, UK