This work was supported by Hunan Provincial Natural Science Foundation of China (2016JJ4038) and 2022NK2050 from the Hunan Provincial Key Research and Development Program. In addition, authors are very grateful for Shanghai Biotree Biotech Co. Ltd for technical support at sequencing services. Professor Yongyao Wu contributed valuable input to the conception of this study. Unfortunately, he passed away from us in 2022 due to cancer, which will be greatly missed. We also thank Dr. Jörg Rinklebe CRediT authorship contribution statement Hongshi Xiao: Methodology, Investigation, Writing – original draft. Jun Tan: Software. Mengjia Li: Data curation. Zhihui Yuan: Supervision, Writing – review. Haiyan Zhou: acquisition, Supervision, Writing – review & editing.