1. College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China; 2. State Key Laboratory of Crop Genetics & Germplasm Enhancement and Utilization, Nanjing Agricultural University, Nanjing 210095, China; 3. Sanya Institute of Nanjing Agricultural University, Nanjing Agricultural University, Sanya 572025, China †These authors contributed equally to this work. *Correspondence: Aying Zhang (ayzhang@njau.edu.cn)