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Structures and mechanisms of the Arabidopsis cytokinin transporter AZG1

  • Lingyi Xu,
  • Wei Jia,
  • Xin Tao,
  • Fan Ye,
  • Yan Zhang,
  • Zhong Jie Ding,
  • Shao Jian Zheng,
  • Shuai Qiao,
  • Nannan Su,
  • Yu Zhang,
  • Shan Wu,
  • Jiangtao Guo

摘要

Cytokinins are essential for plant growth and development, and their tissue distributions are regulated by transmembrane transport. Recent studies have revealed that members of the ‘Aza-Guanine Resistant’ (AZG) protein family from Arabidopsis thaliana can mediate cytokinin uptake in roots. Here we present 2.7 to 3.3 Å cryo-electron microscopy structures of Arabidopsis AZG1 in the apo state and in complex with its substrates trans-zeatin (tZ), 6-benzyleaminopurine (6-BAP) or kinetin. AZG1 forms a homodimer and each subunit shares a similar topology and domain arrangement with the proteins of the nucleobase/ascorbate transporter (NAT) family. These structures, along with functional analyses, reveal the molecular basis for cytokinin recognition. Comparison of the AZG1 structures determined in inward-facing conformations and predicted by AlphaFold2 in the occluded conformation allowed us to propose that AZG1 may carry cytokinins across the membrane through an elevator mechanism.