1. State Key Laboratory of Plant Genomics, Institute of Genetics and Developmental Biology, The Innovative Academy of Seed Design, Chinese Academy of Sciences, Beijing 100101, China; 2. University of Chinese Academy of Sciences, Beijing 100039, China; 3. Center for Microbes, Development and Health, Institut Pasteur of Shanghai, Chinese Academy of Sciences, Shanghai 200031, China †Yihua Ma and Qingwen Chen contributed equally to this work. *Correspondence: Guodong Wang (gdwang@genetics.ac.cn)