About the Authors

Shengping Li

Contributed equally to this work with: Shengping Li, Ji Qian, Yuan Yang

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Ji Qian

Contributed equally to this work with: Shengping Li, Ji Qian, Yuan Yang

Affiliation State Key Laboratory of Genetic Engineering, Center for Fudan–VARI Genetic Epidemiology and MOE Key Laboratory of Contemporary Anthropology, School of Life Sciences, Fudan University, Shanghai, China

Yuan Yang

Contributed equally to this work with: Shengping Li, Ji Qian, Yuan Yang

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Wanting Zhao

Affiliation Human Genetics, Genome Institute of Singapore, A*STAR, Singapore, Singapore

Juncheng Dai

Affiliation MOE Key Laboratory of Modern Toxicology, Jiangsu Key Laboratory of Cancer Biomarkers, Prevention, and Treatment, and State Key Laboratory of Reproductive Medicine, Nanjing Medical University, Nanjing, China

Jin-Xin Bei

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Jia Nee Foo

Affiliation Division of Genetics, Brigham and Women's Hospital, Harvard Medical School Boston and Broad Institute of Harvard and MIT, Cambridge, Massachusetts, United States of America

Paul J. McLaren

Affiliation Division of Genetics, Brigham and Women's Hospital, Harvard Medical School Boston and Broad Institute of Harvard and MIT, Cambridge, Massachusetts, United States of America

Zhiqiang Li

Affiliation Bio-X Institutes, Key Laboratory for the Genetics of Developmental and Neuropsychiatric Disorders (Ministry of Education), Shanghai Jiao Tong University, Shanghai, China

Jingmin Yang

Affiliation State Key Laboratory of Genetic Engineering, Center for Fudan–VARI Genetic Epidemiology and MOE Key Laboratory of Contemporary Anthropology, School of Life Sciences, Fudan University, Shanghai, China

Feng Shen

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Li Liu

Affiliation Department of Hepatobiliary Surgery, Nantong Tumor Hospital, Nantong, China

Jiamei Yang

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Shuhong Li

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Shandong Pan

Affiliation MOE Key Laboratory of Modern Toxicology, Jiangsu Key Laboratory of Cancer Biomarkers, Prevention, and Treatment, and State Key Laboratory of Reproductive Medicine, Nanjing Medical University, Nanjing, China

Yi Wang

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Wenjin Li

Affiliation Bio-X Institutes, Key Laboratory for the Genetics of Developmental and Neuropsychiatric Disorders (Ministry of Education), Shanghai Jiao Tong University, Shanghai, China

Xiangjun Zhai

Affiliation Department of Infection Diseases, Jiangsu Province Center for Disease Prevention and Control, Nanjing, China

Boping Zhou

Affiliation The Third People's Municipal Hospital of Shenzhen, Shenzhen, China

Lehua Shi

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Xinchun Chen

Affiliation The Third People's Municipal Hospital of Shenzhen, Shenzhen, China

Minjie Chu

Affiliation MOE Key Laboratory of Modern Toxicology, Jiangsu Key Laboratory of Cancer Biomarkers, Prevention, and Treatment, and State Key Laboratory of Reproductive Medicine, Nanjing Medical University, Nanjing, China

Yiqun Yan

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Jun Wang

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Shuqun Cheng

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Jiawei Shen

Affiliation Bio-X Institutes, Key Laboratory for the Genetics of Developmental and Neuropsychiatric Disorders (Ministry of Education), Shanghai Jiao Tong University, Shanghai, China

Weihua Jia

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Jibin Liu

Affiliation Department of Hepatobiliary Surgery, Nantong Tumor Hospital, Nantong, China

Jiahe Yang

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Zujia Wen

Affiliation Bio-X Institutes, Key Laboratory for the Genetics of Developmental and Neuropsychiatric Disorders (Ministry of Education), Shanghai Jiao Tong University, Shanghai, China

Aijun Li

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Ying Zhang

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Guoliang Zhang

Affiliation The Third People's Municipal Hospital of Shenzhen, Shenzhen, China

Xianrong Luo

Affiliation No. 458 Hospital of Chinese People's Liberation Army, Guangzhou, China

Hongbo Qin

Affiliation The Eighth Municipal People's Hospital of Guangzhou, Guangzhou, China

Minshan Chen

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Hua Wang

Affiliation Department of Infection Diseases, Jiangsu Province Center for Disease Prevention and Control, Nanjing, China

Li Jin

Affiliation State Key Laboratory of Genetic Engineering, Center for Fudan–VARI Genetic Epidemiology and MOE Key Laboratory of Contemporary Anthropology, School of Life Sciences, Fudan University, Shanghai, China

Dongxin Lin

Affiliation State Key Laboratory of Molecular Oncology and Department of Etiology and Carcinogenesis, Cancer Institute and Hospital, Chinese Academy of Medical Sciences and Peking Union Medical College, Beijing, China

Hongbing Shen

Affiliation MOE Key Laboratory of Modern Toxicology, Jiangsu Key Laboratory of Cancer Biomarkers, Prevention, and Treatment, and State Key Laboratory of Reproductive Medicine, Nanjing Medical University, Nanjing, China

Lin He

Affiliations Bio-X Institutes, Key Laboratory for the Genetics of Developmental and Neuropsychiatric Disorders (Ministry of Education), Shanghai Jiao Tong University, Shanghai, China, Institutes for Nutritional Sciences, Shanghai Institute of Biological Sciences, Chinese Academy of Sciences, Shanghai, China, Institutes of Biomedical Sciences, Fudan University, Shanghai, China

Paul I. W. de Bakker

Affiliation Division of Genetics, Brigham and Women's Hospital, Harvard Medical School Boston and Broad Institute of Harvard and MIT, Cambridge, Massachusetts, United States of America

Hongyang Wang

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Yi-Xin Zeng

Affiliation Department of Hepatobiliary Oncology, State Key Laboratory of Oncology in South China, Sun Yat-Sen University Cancer Center, Guangzhou, China

Mengchao Wu

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Zhibin Hu

ehphwp@126.com (W Zhou); liuj3@gis.a-star.edu.sg (J Liu); shiyongyong@gmail.com (Y Shi); zhibin_hu@njmu.edu.cn (Z Hu)

Affiliation MOE Key Laboratory of Modern Toxicology, Jiangsu Key Laboratory of Cancer Biomarkers, Prevention, and Treatment, and State Key Laboratory of Reproductive Medicine, Nanjing Medical University, Nanjing, China

Yongyong Shi

ehphwp@126.com (W Zhou); liuj3@gis.a-star.edu.sg (J Liu); shiyongyong@gmail.com (Y Shi); zhibin_hu@njmu.edu.cn (Z Hu)

Affiliation Bio-X Institutes, Key Laboratory for the Genetics of Developmental and Neuropsychiatric Disorders (Ministry of Education), Shanghai Jiao Tong University, Shanghai, China

Jianjun Liu

ehphwp@126.com (W Zhou); liuj3@gis.a-star.edu.sg (J Liu); shiyongyong@gmail.com (Y Shi); zhibin_hu@njmu.edu.cn (Z Hu)

Affiliation Human Genetics, Genome Institute of Singapore, A*STAR, Singapore, Singapore

Weiping Zhou

ehphwp@126.com (W Zhou); liuj3@gis.a-star.edu.sg (J Liu); shiyongyong@gmail.com (Y Shi); zhibin_hu@njmu.edu.cn (Z Hu)

Affiliation Eastern Hepatobiliary Surgery Hospital, Second Military Medical University, Shanghai, China

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Study concept and design: Weiping Zhou, Shengping Li, Jianjun Liu, Zhibin Hu, Yongyong Shi, Ji Qian, Yuan Yang. Subject recruitment and sample preparation: Weiping Zhou, Yuan Yang, Ji Qian, Jingmin Yang, Feng Shen, Jiamei Yang, Yi Wang, Lehua Shi, Yiqun Yan, Shuqun Cheng, Jiahe Yang, Aijun Li, Li Jin (Shanghai samples); Shengping Li, Shuhong Li, Jun Wang, Boping Zhou, Weihua Jia, Ying Zhang, Xinchun Chen, Guoliang Zhang, Xianrong Luo, Hongbo Qin, Minshan Chen (Guangzhou samples); Zhibin Hu, Li Liu, Shandong Pan, Minjie Chu, Xiangjun Zhai, Jibin Liu, Hua Wang (Jiangsu samples). GWAS data processing: Zhiqiang Li, Jiawei Shen, Lehua Shi. Sample genotyping: Wenjin Li, Zujia Wen (GWAS Scan); Li Liu, Shandong Pan, Minjie Chu (Validation stages). Analysis and interpretation of data: Weiping Zhou, Shengping Li, Jianjun Liu, Zhibin Hu, Yongyong Shi, Ji Qian, Yuan Yang, Juncheng Dai, Wanting Zhao. Drafting of the manuscript: Jianjun Liu, Zhibin Hu. Critical revision of the manuscript for important intellectual content: Weiping Zhou, Shengping Li, Jianjun Liu, Zhibin Hu, Yongyong Shi, Jin-Xin Bei, Dongxin Lin, Hongbing Shen, Lin He, Hongyang Wang, Yi-Xin Zeng, Mengchao Wu. Statistical analysis: Jungheng Dai, Wanting Zhao, Jia Nee Foo, Paul J McLaren, Paul IW de Bakker, Zhibin Hu, Jianjun Liu. Obtained funding: Weiping Zhou, Shengping Li, Jianjun Liu, Zhibin Hu, Yongyong Shi, Ji Qian. Administrative, technical, or material support: Hongbing Shen, Lin He, Hongyang Wang, Yi-Xin Zeng, Mengchao Wu. Study supervision: Weiping Zhou, Shengping Li, Jianjun Liu, Zhibin Hu, Yongyong Shi, Ji Qian.