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ResearchIn-Press PreviewHepatologyMetabolism
Open Access | 
10.1172/JCI196374
Iron overload in steatotic hepatocytes drives systemic metabolic dysfunction via alterations in hepatokine production
Hye Jin Jo,1 Ayoung Kim,2 Hyunsoo Rho,3 Ae Kyung Park,1 Gil-Hwan Kim,1 Seo Jeong Jo,1 Hao Yuxin,1 You-Jung Hong,1 Ji Min Yeon,1 Hwang Chan Yu,4 Mi-Young Song,5 Jeongwoo Park,6 Yeon Hee Jeong,7 Sung Eun Hong,8 Hyo Jin Yeon,8 Da Young Oh,9 Philipp E. Scherer,9 Cheol Soo Choi,10 Dong Hyeon Lee,7 Sung Hwan Ki,11 Keon Wook Kang,6 Murim Choi,12 Byung-Hyun Park,4 Eun Ju Bae,1 Sang Geon Kim,14 Won Kim,7 and Chang Yeob Han1
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Jo, H. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Kim, A. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Rho, H. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Park, A. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Kim, G. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Jo, S. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Yuxin, H. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Hong, Y. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Yeon, J. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Yu, H. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Song, M. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Park, J. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Jeong, Y. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
Find articles by Hong, S. in: PubMed | Google Scholar
1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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Scherer, P.
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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Kim, W.
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1School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonbuk, Korea, Republic of
2Department of Pathology, Washington University School of Medicine, St. Louis, United States of America
3College of Pharmacy, Graduate School of Pharmaceutical Sciences, Ewha Womans University, Seoul, Korea, Republic of
4Graduate School of Medical Science and Engineering, Korea Advanced Institute of Science and Technology, Daejeon, Korea, Republic of
5Department of Biochemistry and Molecular Biology, Jeonbuk National University Medical School, Jeonbuk, Korea, Republic of
6College of Pharmacy and Research Institute of Pharmaceutical Sciences, Seoul National University, Seoul, Korea, Republic of
7Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea, Republic of
8Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea, Republic of
9Touchstone Diabetes Center, Department of Internal Medicine, University of Texas Southwestern Medical Center, Dallas, United States of America
10Department of Internal Medicine, Gil Medical Center, Gachon University College of Medicine, Incheon, Korea, Republic of
11MRC-OSTRC, Research Institute of Pharmaceutical Sciences, College of Pharmacy, Chosun University, Gwangju, Korea, Republic of
12Department of Biomedical Sciences, College of Medicine, Seoul National University, Seoul, Korea, Republic of
13School of Pharmacy and Institute of New Drug Development, Jeonbuk National University, Jeonju, Korea, Republic of
14College of Pharmacy and Integrated Research Institute for Drug Development, Dongguk University, Goyang, Korea, Republic of
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Published April 28, 2026 - More info
Copyright © 2026, Jo et al. This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.
Copyright © 2026 American Society for Clinical Investigation
ISSN: 0021-9738 (print), 1558-8238 (online)