1 State Key Laboratory of Plant Physiology and Biochemistry, Department of Plant Sciences, College of Biological Sciences, China Agricultural University, Beijing 100193, China 2 China Agricultural University (CAU)-University of California, Riverside (UCR) Joint Center for Biological Sciences and Biotechnology, College of Biological Sciences, China Agricultural University, Beijing 100193, China 3 Center for Plant Cell Biology and Department of Botany and Plant Sciences, University of California, Riverside, CA 92521, USA