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ABSTRACT Title of Document: INTERFERENCE of HOST ABSTRACT Title of Document: INTERFERENCE OF HOST INNATE IMMUNE RESPONSE BY HEPATITIS E VIRUS Yuchen Nan, Ph.D., 2014 Directed By: Associate Professor Yanjin Zhang, Department of Veterinary Medicine The host antiviral innate immunity mainly relies on host pattern recognition receptors (PRR) and downstream interferon (IFN) signaling. Host PRR for RNA viruses include Toll-like receptors (TLR) and Retinoic acid-inducible gene I (RIG-I) like receptors (RLR). Activation of both TLR and RLR pathways can eventually lead to the secretion of type I IFNs, which can modulate both innate and adaptive immune responses against viral pathogens, including hepatitis E virus (HEV). HEV causes acute hepatitis in humans and has been responsible for several outbreaks of hepatitis across the world. Currently, no commercial vaccine is available for the prevention of HEV infection in any country except China. HEV biology and pathogenesis as well as its responses to host innate immunity are poorly understood, though other hepatitis viruses, including the hepatitis A, B and C viruses, have been much better studied. In this study, how HEV interferes with IFN induction and IFN-activated signaling had been examined. Results showed that the protein encoded by HEV ORF1 can inhibit type I IFN synthesis and downstream JAK/STAT signaling pathway. However, the HEV ORF3 product is able to enhance RIG-I-mediated signaling to a certain extent. These data suggest that HEV proteins interfere with the host innate immune response and may exert the diverse roles depending on the stage and/or context of infection. These studies contribute to a better understanding of HEV pathogenesis and may facilitate a strategy development for the prevention and control of HEV infection. INTERFERENCE OF HOST INNATE IMMUNE RESPONSE BY HEPATITIS E VIRUS By Yuchen Nan Dissertation submitted to the Faculty of the Graduate School of the University of Maryland, College Park, in partial fulfillment of the requirements for the degree of Doctor of Philosophy 2014 Advisory Committee: Associate Professor Yanjin Zhang, Chair Professor Xiaoping Zhu Associate Professor Wenxia Song Associate Professor Utpal Pal Assistant Professor Georgiy Belov © Copyright by Yuchen Nan 2014 Preface My past five years as a graduate student in the Department of Veterinary Medicine, University of Maryland, College Park had been the most exceptional part of my life. One reason is that I had incorrect conceptions and expectation about life as a Ph.D. student or a research scientist before I joined the department as I grew up in Northwest China, a remote and less developed region. Life and study as a foreign graduate student in this country for five years not only open my eyes, but also broaden my mind. I appreciate the training received here because it not only teaches me how to solve academic research problems and think logically, but also makes me be strong when facing difficulties and obstacles. No matter what kind of life will be for my next step, I am sure that I am ready for any challenge or obstacle. ii Dedication This body of work is dedicated to the memory of my grandfather Xiangqian Nan and grandmother Jinnv Cai. I wish they could see all this, and I know they would be proud for my achievement today. I also want to dedicate this work to my father Jiangzhen Nan and mother Yan Dong for raising me up. I know my achievement finally realizing your dream and expectation. You will be proud of me as well. Lastly, but the most importantly, I want to dedicate this work to my wife Chunyan Wu and our upcoming daughter Olivia. Chunyan, reunion with you after our graduation from Northwest A&F University is the happiest time in my memory. Our love is so exciting for all of us and surprising for anyone who knows us. With your graduation with Ph.D. degree from University of Munich, you could have lived a different life by taking a job in China. However, you choose to stay with me and support me without reservation. I am highly grateful for the sacrifice you have made for me. Olivia, your will be the first member from our family who is born in the United States. I wish you grow up with a happy childhood and have the freedom to choose the lifestyle in the way you want. You will no longer suffer the hardship I experienced, just like I did not have the pain your grandfather had in the past. iii Acknowledgements First of all, I thank my academic advisor Dr. Yanjin Zhang for his offering me the opportunity to study in the United States. His patience in guiding and supporting my study in the last five years is indispensable for my achievement today. His scientific advices made me avoid wasting time in some dead end ideas and get out quickly via trouble shooting. His wise advices and suggestions not only give me courage but also make me be optimistic for life and future. His sharing of the common experience as a foreigner makes me adapt to the new life here in the shortest time possible. Secondly, I thank my uncle Dr. Bin He from the University of Illinois at Chicago. With his encouragement, I came to this country as a graduate student. He helped me a lot both in academic field and in personal life during the past few years here. Thirdly, I thank my committee members Drs. Belov, Pal, Song, and Zhu for their kind help during my studies. Their kind advices and suggestions contributed to my research and study. Fourthly, I thank all the faculty members and staff I met in the Department of Veterinary Medicine. Drs. Zhu, Pal. Samal, Belov, Bossis and Shi are exemplary scientists for me. I also thank Drs. Wenxia Song, Jeffery DeStefano, Brenda Fredericksen, David Mosser and Vincent Lee for teaching me and sharing knowledge with me in the courses I took. Furthermore, I thank Drs. Suzanne Emerson in NIH, and X-J Meng at Virginia Tech University for sharing the plasmids and antibodies used in this study. iv Lastly, I thank my current colleagues Zexu Ma, Liping Yang, and Yonglin Yang for their help in daily work. I also thank my former colleagues, Meiyan Shen, Krit Rithipichai, Deendayal Patel, Rong Wang and Ying Yu for their help to me in the past. Finally, I give my special thanks to my best friend Fan Yang for his friendship, which is a strong support for me to live in this country during the past four years. v Table of Contents Preface........................................................................................................................... ii Dedication .................................................................................................................... iii Acknowledgements ...................................................................................................... iv Table of Contents ......................................................................................................... vi List of Tables ............................................................................................................... ix List of Figures ............................................................................................................... x Chapter 1:Hepatitis E Virus .......................................................................................... 1 Introduction ............................................................................................................... 1 HEV Virology ........................................................................................................... 1 Genotypes of Hepatitis E viruses .............................................................................. 3 Viral Proteins of HEV ............................................................................................... 6 ORF1 polyprotein ................................................................................................. 6 The capsid protein encoded by ORF2 ................................................................. 16 ORF3-encoded protein ........................................................................................ 19 Replication cycle of Hepatitis E Virus.................................................................... 24 Cell culture and propagation of Hepatitis E virus ................................................... 25 Transmission and epidemiology of HEV ................................................................ 27 Pathogenesis, clinical signs and diagnosis of HEV ................................................ 29 Treatment and Prevention of HEV ......................................................................... 31 Summary ................................................................................................................. 32 Chapter 2:Interferons and Induction ........................................................................... 34 Introduction ............................................................................................................. 34 Interferons and Their Classification........................................................................ 34 Toll-like Receptor Pathway .................................................................................... 37 Discovery of Toll-like receptors in mammalian hosts ........................................ 38 TLR ligands ........................................................................................................ 40 TLR signaling ..................................................................................................... 43 RIG-I like Receptor Pathway ................................................................................
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