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THE ROLE of HERPESVIRUSES in MARINE TURTLE DISEASES By THE ROLE OF HERPESVIRUSES IN MARINE TURTLE DISEASES By SADIE SHEA COBERLEY A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2002 Copyright 2002 by Sadie Shea Coberley For the turtles, and Carter and my family for encouraging me to pursue what I love. ACKNOWLEDGEMENTS I would like to thank my mentor, Dr. Paul Klein, for sharing his knowledge and for all of his encouragement and patience throughout my graduate education. He has been a true mentor in every sense of the word, and has done everything possible to prepare me for not only my scientific future, but phases of life outside of the laboratory as well. I would also like to thank my co-mentor, Dr. Rich Condit, first for seeing graduate student potential, and then for taking me in and helping to provide the necessary tools and expertise to cultivate it. In addition, I am indebted to Dr. Larry Herbst, who was not only my predecessor but a pioneer in FP research. His insight into studying such a complex problem has been invaluable. I am grateful for the critical analysis and raised eyebrow of Dr. Daniel Brown and for his assistance with trouble-shooting experiments, evaluating data, and preparing manuscripts. I am also appreciative of the assistance of Dr. Elliott Jacobson for including me in many discussions, necropsies, and analyses of marine turtles with interesting clinical signs of disease, and for sharing his vast knowledge of reptile diseases. I would like to thank Dr. Maureen Goodenow for her valuable input at committee meetings and for her assistance specifically with phylogenetic analyses. I would like to extend my gratitude to Dr. David Bloom for agreeing to join my committee so late in the project and for contributing his fresh enthusiasm and expertise in herpesviral research. I would also like to thank Dr. Wayne McCormack for his input and assistance during the final examination. It has been a blessing to have such a functional iv committee and to have such an enjoyable group of researchers to help me through my scientific struggles. I would like to extend a heartfelt thank you to Diane Duke for her constant moral support, friendship, and laboratory rescues. I would also like to extend a special thank you to all of the members of the Condit laboratory for their invaluable technical guidance, for letting me constantly interrupt them with questions, and for listening to turtle research. This has included Drs. Cari Lackner, Cindy Prins, Susan D’Costa, and Don Latner, fellow IDP graduate student Steve Cresawn, and Jackie Fried. I would also like to say thank you to Dr. Nissin Moussatche for all of his help and for being so much fun to harass, and to Dr. Ed Niles for his timely visit to the Condit lab and for his assistance with bacterial expression that led to the expression of the LETV proteins. I would also like to thank Francesco Origgi and Joyce Merritt for their encouragement and support in the laboratory and their friendship. I would like to thank the members of the University of Florida Interdisciplinary Center for Biotechnology Research (ICBR) Hybridoma core laboratory (Gainesville, FL, USA), Linda Green, Shadi Bootorabi, Jamie Kelso, and Scherwin Henry, for all of their valuable help through the years. I would like to extend my gratitude to Marjorie Chow of the ICBR Molecular Biomarker core laboratory for teaching me the technique of two- dimensional gel electrophoresis and for her encouragement and friendship. I am grateful for the assistance of Drs. Regina Shaw and Bill Farmerie of the ICBR Genomic core laboratory for construction of the LETV genomic library. Finally, I would like to thank Susanna Lamers of Gene Genie Co. (Thibdaux, LA, USA) for her assistance with phylogenetic analysis and the construction of the phylogenetic trees. v I am indebted to all of our collaborators who have been our connection to the sea turtles. I am grateful for all of the hard work and dedication of Richie Moretti, Sue Schaf, and Corinne Rose at the Turtle Hospital (Marathon, FL, USA). I would like to thank all of the members of the University of Central Florida Marine Turtle Research Program headed by Dr. Llewellyn Ehrhart. I would especially like to thank Dean Bagley for all the plasma samples she has collected that we appreciate and rely on daily. I would like to thank Dr. Jeanette Wyneken for the crash course in sea turtle biology and having faith in a lab rat. It is only through these broad collaborations that it has been possible to address such complex problems. I would like to extend my gratitude to all of the members of the turtle community that I have had the privilege to work with and for their constant support and encouragement. Their dedication to the survival of marine turtles is inspiring. Finally, I am grateful for the continued support of the U.S. Fish and Wildlife Service and the assistance of Sandy MacPherson. This research was funded by research grants RWO 180, 194, and 213 from the U.S. Fish and Wildlife Service, Department of the Interior, administered by the Cooperative Fish and Wildlife Unit, University of Florida, Gainesville, FL, USA. vi TABLE OF CONTENTS page ACKNOWLEDGEMENTS ................................................................................................iv LIST OF TABLES .............................................................................................................xii LIST OF FIGURES ..........................................................................................................xiii ABSTRACT......................................................................................................................xvi CHAPTER 1 INTRODUCTION ...........................................................................................................1 Microorganisms Associated with Diseases of Marine Turtles ....................................... 1 Herpesvirus Infections of Marine Turtles ................................................................ 2 GPD-associated herpesvirus ..............................................................................3 LETD-associated herpesvirus ............................................................................4 Fibropapillomatosis-associated herpesvirus ......................................................5 The Herpesvirus Family.................................................................................................. 6 Lytic Infection.......................................................................................................... 8 Human herpesviruses .........................................................................................9 Chelonian herpesviruses ..................................................................................10 Oncogenic Herpesviruses....................................................................................... 11 Human simplex virus .......................................................................................13 Epstein-Barr virus ............................................................................................13 Human herpesvirus 6 .......................................................................................15 Kaposi Sarcoma-associated herpesvirus ..........................................................16 Chelonian herpesviruses and tumors in reptiles...............................................19 Methods for Linking Viruses with Disease Etiology.................................................... 20 Henle-Koch Postulates ........................................................................................... 20 Rivers’ Review....................................................................................................... 21 Hill’s Criteria for Causation................................................................................... 22 Evans Unified Concept........................................................................................... 22 Strategies To Demonstrate Association Between Herpesviral Infection and Onset of Disease by Fulfilling Hill’s Criteria................................................................... 22 Strength of association.....................................................................................23 Consistency of findings....................................................................................24 Biological gradient ...........................................................................................25 Specificity of association.................................................................................27 vii Temporal association.......................................................................................28 Coherence to the biology of the disease...........................................................29 Experimental evidence.....................................................................................30 Biological plausibility and analogy .................................................................32 Strategies to Investigate the Role of Herpesviruses in Marine Turtle Diseases..... 34 2 STUDIES ON THE REPLICATION OF MARINE TURTLE HERPESVIRUSES.....36 Introduction................................................................................................................... 36 Materials and Methods.................................................................................................. 37 FP Filtrate Preparation..........................................................................................
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