ResearchIn-Press PreviewGastroenterologyHepatology
Open Access | 
10.1172/jci.insight.193805
PNPLA3-I148M genetic variant rewires lipid metabolism to drive programmed cell death in human hepatocytes
Rodrigo M. Florentino,1 Olamide Animasahun,2 Nils Haep,1 Minal Nenwani,3 Kehinde Omoloja,4 Leyla Nurcihan Altay,4 Abhinav Achreja,3 Kazutoyo Morita,1 Takashi Motomura,1 Ricardo Diaz-Aragon,1 Lanuza AP Faccioli,1 Yiyue Sun,1 Zhenghao Liu,1 Zhiping Hu,1 Bo Yang,1 Fulei Wuchu,3 Ajay Shankaran,4 Miya Paserba,3 Annalisa M. Baratta,1 Shohrat Arazov,1 Zehra N. Kocas-Kilicarslan,1 Noah Meurs,3 Jaideep Behari,5 Edgar N. Tafaleng,1 Jonathan Franks,6 Alina Ostrowska,1 Takahiro Tomiyama,7 Kyohei Yugawa,7 Akinari Morinaga,7 Zi Wang,8 Kazuki Takeishi,7 Dillon C. Gavlock,9 Mark Miedel,9 D. Lansing Taylor,10 Ira J. Fox,11 Tomoharu Yoshizumi,7 Deepak Nagrath,2 and Alejandro Soto-Gutierrez1
1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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1Department of Pathology, University of Pittsburgh, Pittsburgh, United States of America
2Center for Transcriptional Medicine, University of Pittsburgh, Pittsburgh, United States of America
3Biointerfaces Institute, University of Michigan, Ann Arbor, United States of America
4Department of Chemical Engineering, University of Michigan, Ann Arbor, United States of America
5Department of Medicine, Division of Gastroenterology, Hepatology, and Nutri, University of Pittsburgh Medical Center, Pittsburgh, United States of America
6Center for Biologic Imaging, University of Pittsburgh, Pittsburgh, United States of America
7Department of Surgery and Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
8Department of Statistic, University of Pittsburgh, Pittsburgh, United States of America
9University of Pittsburgh Drug Discovery Institute, University of Pittsburgh, Pittsburgh, United States of America
10Pittsburgh Liver Research Center, University of Pittsburgh, Pittsburgh, United States of America
11Department of Surgery, Children’s Hospital of Pittsburgh of UPMC, University of Pittsburgh, Pittsburgh, United States of America
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Published October 21, 2025 - More info
Copyright © 2025, Florentino 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/.
