Structural insights into the substrate uptake and inhibition of the human creatine transporter (hCRT)

X Xinyi Yuan (Department of Biochemistry and Molecular Biology, School of Basic Medical Sciences, Hangzhou Normal University) J Jian Yin (Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University) C Chang Liu X Xudong Chen (Ministry of Education Key Laboratory of Protein Science, Tsinghua-Peking Center for Life Sciences, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University) M Meiying Chen (Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University) Y Yixue Wang (Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University) Z Zi Yang (Beijing National Laboratory for Molecular Science, State Key Laboratory of Rare Earth Materials Chemistry and Applications, College of Chemistry and Molecular Engineering) Y Yue Wang L Li Jiang (Department of Radiation Oncology The First Affiliated Hospital of Guangxi Medical University Nanning China) N Niyun Zhou (Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University) X Xiaojuan Wang (Department of Endocrinology, Genetics and Metabolism, National Center for Children’s Health, Beijing Children’s Hospital Capital Medical University) B Botong Liu (Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University) Z Zhaoqi Ma (College of Biological Science, China Agricultural University) K Kaiyan Wang (National Clinical Research Center for Oral Diseases & Shaanxi Key Laboratory of Stomatology, School of Stomatology, The Fourth Military Medical University) H Hongen Li (Department of Ophthalmology, The Chinese People’s Liberation Army General Hospital & The Chinese People’s Liberation Army Medical School) S Sensen Zhang Y Yongfeng Shang (Department of Biochemistry and Molecular Biology, School of Basic Medical Sciences, Hangzhou Normal University) M Maojun Yang (Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University)

Abstract

Creatine plays a vital role in cellular energy production and adenosine triphosphate (ATP) homeostasis and has also been identified as a neurotransmitter in the mammalian brain. Creatine is transported into cells by the human creatine transporter (hCRT) (SLC6A8), an Na + /Cl − -dependent symporter encoded on the X chromosome. Mutations in hCRT cause cerebral creatine deficiency syndrome 1, a neurological disorder marked by intellectual disability, speech delay, and seizures. Beyond its role in the brain and muscle, hCRT is highly expressed in metabolically active tumors. Many cancer cells, including colorectal cancer and glioblastoma, upregulate hCRT to sustain intracellular creatine levels and buffer ATP under energy stress. Pharmacological blockade of hCRT by RGX202 has been shown to impair tumor growth by disrupting energy homeostasis. Here, we report the high-resolution cryo-Electron Microscopy (cryo-EM) structures of human hCRT in three states: apo, creatine-bound, and RGX202-bound. hCRT adopts a canonical LeuT-fold with 12 transmembrane helices and two pseudosymmetric inverted repeats. Creatine is coordinated in the central substrate-binding site through interactions with transmembrane helices TM1, TM3, TM6, and TM8, while the inhibitor RGX202 occupies the same binding pocket, engaging in overlapping contacts that competitively block creatine access. Our structural and mechanistic findings clarify substrate recognition and inhibitory binding of hCRT, providing a molecular rationale for targeting hCRT in both inherited metabolic diseases and cancer therapy.

Article Details

Volume / Issue Vol. 122, Issue 36
Published September 09, 2025
ISSN 0027-8424
Publisher National Academy of Sciences

Authors (18)

X

Xinyi Yuan

Department of Biochemistry and Molecular Biology, School of Basic Medical Sciences, Hangzhou Normal University

J

Jian Yin

Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University

C

Chang Liu

X

Xudong Chen

Ministry of Education Key Laboratory of Protein Science, Tsinghua-Peking Center for Life Sciences, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University

M

Meiying Chen

Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University

Y

Yixue Wang

Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University

Z

Zi Yang

Beijing National Laboratory for Molecular Science, State Key Laboratory of Rare Earth Materials Chemistry and Applications, College of Chemistry and Molecular Engineering

Y

Yue Wang

L

Li Jiang

Department of Radiation Oncology The First Affiliated Hospital of Guangxi Medical University Nanning China

N

Niyun Zhou

Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University

X

Xiaojuan Wang

Department of Endocrinology, Genetics and Metabolism, National Center for Children’s Health, Beijing Children’s Hospital Capital Medical University

B

Botong Liu

Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University

Z

Zhaoqi Ma

College of Biological Science, China Agricultural University

K

Kaiyan Wang

National Clinical Research Center for Oral Diseases & Shaanxi Key Laboratory of Stomatology, School of Stomatology, The Fourth Military Medical University

H

Hongen Li

Department of Ophthalmology, The Chinese People’s Liberation Army General Hospital & The Chinese People’s Liberation Army Medical School

S

Sensen Zhang

Y

Yongfeng Shang

Department of Biochemistry and Molecular Biology, School of Basic Medical Sciences, Hangzhou Normal University

M

Maojun Yang

Ministry of Education Key Laboratory of Protein Science, Beijing Advanced Innovation Center for Structural Biology, School of Life Sciences, Tsinghua University