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Genetic Characteristics of Field and Attenuated Rabies Viruses And Evira Research Reports 2/2008 Artem Metlin Genetic characteristics of field and attenuated rabies viruses and molecular epidemiology of rabies in Finland and Russia Finnish Food Safety Authority Evira, Helsinki, Finland Department of Basic Veterinary Sciences, Faculty of Veterinary Medicine, University of Helsinki, Helsinki, Finland Genetic characteristics of field and attenuated rabies viruses and molecular epidemiology of rabies in Finland and Russia Artem Metlin ACADEMIC DISSERTATION To be presented with the permission of the Faculty of Veterinary Medicine, University of Helsinki, for public examination in the Walter Hall, EE-building, Viikki on Monday 28th of January 2008 at 12 o’clock HELSINKI 2008 Supervised by Anita Huovilainen, PhD, Docent Virology Unit, Department of Animal Diseases and Food Safety, Finnish Food Safety Authority Evira, Helsinki, Finland Olli Vapalahti, MD, PhD, Professor, Haartman Institute, Faculty of Medicine, University of Helsinki, Faculty of Veterinary Medicine, University of Helsinki, Finland Supervising professor Antti Sukura, DVM, PhD, Professor, Department of Basic Veterinary Sciences, Faculty of Veterinary Medicine, University of Helsinki, Finland Reviewed by Antti Vaheri, MD, PhD, Professor Haartman Institute, University of Helsinki, Finland Thomas Mueller, DVM, PhD, Institute for Epidemiology, WHO Collaborating Centre for Rabies Surveillance and Research, OIE Reference Laboratory for Rabies, Friedrich-Loeffler-Institute – Federal Research Institute for Animal Health, Wusterhausen, Germany Opponent Noel Tordo, DVM, PhD, Pasteur Institute, France 3 Content List of abbreviations ....................................................................................................... 5 Abstract............................................................................................................................ 6 List of original publications............................................................................................ 8 1. Literature review........................................................................................................ 9 1.2. Rabies virus characteristics ................................................................................ 9 1.2.1. Classification ................................................................................................... 9 1.2.2. Structure of virion, genome, and proteins of rabies virus............................... 10 1.3. Virus-host interaction ........................................................................................ 12 1.3.1. Pathogenesis................................................................................................. 12 1.3.2. Clinical signs.................................................................................................. 13 1.3.3. Pathological changes .................................................................................... 15 1.4. Laboratory diagnosis of rabies .......................................................................... 17 1.4.1. Detection of viral antigen ............................................................................... 17 1.4.2. Detection of viral genome.............................................................................. 18 1.4.3. Virus isolation ................................................................................................ 19 1.4.4. Detection of virus neutralizing antibodies ...................................................... 19 1.5. Rabies epidemiology......................................................................................... 21 1.5.1. Rabies situation world-wide........................................................................... 21 1.5.2. Rabies situation in Europe and in Russia ...................................................... 22 1.5.3. Models of rabies epidemics ........................................................................... 24 1.6. Fixed strains of rabies virus .............................................................................. 25 1.6.1. Laboratory strains.......................................................................................... 25 1.6.2. Vaccine strains and anti-rabies vaccines....................................................... 25 1.7. Molecular biology of rabies viruses ................................................................... 30 1.7.1. Antigenic characterization with monoclonal antibodies.................................. 30 1.7.2. Genetic characterization of rabies virus......................................................... 31 1.7.3. Entire genome sequencing of vaccine strains ............................................... 33 2. Aims of the study .................................................................................................... 35 3. Materials and methods............................................................................................ 36 3.1. Field rabies virus isolates.................................................................................. 36 3.2. Vaccine and laboratory strains.......................................................................... 36 3.3. Cell cultures ...................................................................................................... 36 3.4. Monoclonal antibodies ...................................................................................... 37 4 3.5. Laboratory techniques....................................................................................... 37 3.5.1. Fluorescent antibody test .............................................................................. 37 3.5.2. Cell culture inoculation test............................................................................ 37 3.5.3. Reverse-transcriptase polymerase chain reaction......................................... 38 3.5.4. Nucleotide sequencing .................................................................................. 39 3.5.5. Entire genome sequencing of the vaccine strain RV-97 ................................ 39 3.6. Phylogenetic analysis........................................................................................ 40 4. Results ..................................................................................................................... 42 4.1. FAT and cell culture inoculation test ................................................................. 42 4.2. Antigenic characteristics of field rabies viruses (papers I, II and III).................. 42 4.3. Development of RT-PCR tests (papers II and III).............................................. 43 4.4. Phylogenetic analysis and molecular epidemiology (papers III and IV)............. 44 4.5. Entire genome sequencing and characterization of RV-97 strain (paper V)...... 47 5. Discussion ............................................................................................................... 50 6. Concluding remarks................................................................................................ 59 7. Заключение (concluding remarks in Russian) .................................................... 60 8. Acknowledgments................................................................................................... 62 9. References ............................................................................................................... 63 5 List of abbreviations ABLV – Australian bat Lyssavirus BHK-21 – baby hamster kidney cell culture CCIT – cell culture inoculation test CNS – central nervous system CVS – challenge/control virus strain cDNA – complementary DNA synthesized from RNA template DEAE-dextran – diethylaminoethyl-dextran EBLV – European Bat Lyssavirus ELISA – enzyme-linked immuno assay ERA – Evelyn Rokitniki Abelseth strain FAT – fluorescent antibody test FAVN – fluorescent antibody virus neutralization test FITC-globulin – fluorescein isothiocyanate labeled globulin IU – international unit IHCT – immunohistochemical test LAT – latex agglutination test mAb – monoclonal antibody MEM – Eagle’s minimal essential medium MIT – mouse inoculation test MNA – murine neuroblastoma cell culture PV – Pasteur Virus strain PM – Pitman-Moore strain RT-PCR – reverse transcriptase polymerase chain reaction RNP – ribonucleoprotein SAD – Street-Alabama-Dufferin strain TCID – tissue culture infectious dose VNA – virus neutralizing antibody 6 Abstract Rabies is a fatal disease that affects the central nervous system of all warm- blooded mammals. The rabies virus belongs to the order Mononegavirales, family Rhabdoviridae, genus Lyssavirus. This virus has a negative single-stranded RNA genome and the virions are bullet-shaped. Rabies is reported in many countries throughout the world and has been registered in all continents except Australia, where only the bat Lyssaviruses have been found, and in Antarctica where the main vectors of rabies are absent. Russia and most of the bordering countries are affected by rabies. Finland was a rabies-free country from 1959 to 1988, when a sylvatic rabies epidemic appeared with raccoon dogs as the main host and vector of infection. That epidemic was eradicated by the oral vaccination of wild carnivores and the parenteral immunization of dogs and cats; and Finland has been rabies-free since 1991. However, this status is constantly under threat because rabies is endemic in Russia and Estonia. In June 2003, a horse imported to Finland from Estonia was clinically and laboratory diagnosed as rabies positive. The close relationship
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