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Epidemiological and Ecological Studies of Tick-Borne Encephalitis Virus

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Epidemiological and Ecological Studies of Tick-Borne Encephalitis Virus Linköping University Medical Dissertations No. 1399 Epidemiological and Ecological Studies of Tick-borne Encephalitis Virus Pontus Lindblom Division of Medical Microbiology Department of Clinical and Experimental Medicine Faculty of Health Sciences Linköping University Sweden 2014 About the cover The cover displays a nymph of the tick species Ixodes ricinus. The work in this thesis was supported by grants from: The Swedish Research Council, branch of Medicine. The Medical Research Council of South-East Sweden. The County Council of Östergötland. The Wilhelm and Else Stockmann Foundation. The Foundation for Åland Medical Research of the Åland Culture Foundation. The Lions Research Foundation, Linköping. The EU Interreg IV A supported project ScandTick. © Pontus Lindblom 2014 Published articles have been reprinted with permission of respective copyright holders. ISBN: 978-91-7519-381-6 ISSN 0345-0082 Printed by LiU-Tryck, Linköping, Sweden, 2014. Intelligence Not because you think you know everything without questioning, but rather because you question everything you think you know Supervisor Per-Eric Lindgren (Professor) Linköping, Sweden Co-supervisors Pia Forsberg (Professor) Linköping, Sweden Mats Haglund (MD, PhD) Kalmar, Sweden Jan Ernerudh (Professor) Linköping, Sweden Opponent Sarah Randolph (Professor) Oxford, United Kingdom Table of contents Abstract ........................................................................................................................................................ 1 Populärvetenskaplig sammanfattning ........................................................................................................ 3 List of papers ................................................................................................................................................ 5 Introduction .................................................................................................................................................. 7 Evolutionary perspectives on parasites and disease ................................................................................ 7 The Tick ..................................................................................................................................................... 9 Life cycle ................................................................................................................................................ 9 The virus .................................................................................................................................................. 11 Replication .......................................................................................................................................... 13 Subtype distribution ........................................................................................................................... 13 Quasispecies........................................................................................................................................ 15 Vector-host transmission .................................................................................................................... 15 Prevalence in ticks ............................................................................................................................... 17 The disease.............................................................................................................................................. 19 History and present epidemiological situation ................................................................................... 19 Pathogenesis ....................................................................................................................................... 21 Clinical picture ..................................................................................................................................... 22 Diagnosis ............................................................................................................................................. 23 Treatment ........................................................................................................................................... 24 Vaccine ................................................................................................................................................ 24 Aims ............................................................................................................................................................ 26 Materials and methods .............................................................................................................................. 27 Ethics ....................................................................................................................................................... 28 The tick-borne diseases STING study ...................................................................................................... 29 Questionnaires ........................................................................................................................................ 31 Tick photo and measurement ................................................................................................................. 31 Development of RNA extraction from ticks ............................................................................................ 32 cDNA synthesis ........................................................................................................................................ 33 Development of real-time PCR for detection and quantification of TBEV ............................................. 33 Extraction and real-time PCR controls .................................................................................................... 36 Cloning .................................................................................................................................................... 36 In vitro transcription ............................................................................................................................... 37 Sequencing and phylogenetic analysis ................................................................................................... 37 Serological methods ................................................................................................................................ 37 Statistics .................................................................................................................................................. 38 Summary of results .................................................................................................................................... 39 Interactions between ticks and humans ................................................................................................. 39 TBEV in ticks detached from humans ..................................................................................................... 42 Dose response of vaccination in older individuals .................................................................................. 44 Quasispecies of TBEV in blood-feeding ticks .......................................................................................... 47 Concluding remarks .................................................................................................................................... 50 Future perspectives .................................................................................................................................... 52 Acknowledgements .................................................................................................................................... 53 References .................................................................................................................................................. 55 Abstract Ticks are blood-sucking parasites that are an inconvenience for both humans and animals. The tick by itself is normally harmless unless they attack in excessive numbers. The harm from ticks stems from them being excellent vectors for other parasites, in the form of bacteria and virus that via the ticks are provided a bridge to move across the blood streams of different animals, including humans. One of the most pathogenic tick-borne disease for humans is caused by a flavivirus, the tick- borne encephalitis virus (TBEV). Each year approximately 10 000 individuals on the Eurasian continent develop neurological disease, in the form of meningitis, encephalitis, myelitis and radiculitis, following a bite by a TBEV infected tick. To evaluate the risk of TBEV infection after a tick-bite, we have developed a study to investigate ticks that have bitten humans and to follow up the tick-bitten humans to investigate if they get infected, and if they develop symptoms, and further trace the virus back to the tick that is infected with TBEV. Ticks, blood samples, and questionnaires were collected in collaboration with 34 primary health care centers in Sweden and on the Åland Islands during 2008 and 2009. Several demographical and biological factors were investigated regarding the interaction between ticks and humans. The main finding was that men removed the ticks later than women, and that both older men and older women removed the ticks later than younger individuals. This could in part explain why older individuals in general, and men in particular, are at greater risk of
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