I.persulcatus BHK. Ixodidae 30% C-1 A-1 Sofjin KH TBE. cdna

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1 55 pp ,,,, BHK 1 cdna 1 Ns Ixodidae 30% ,000 TBE cdna TEL FAX takasima@vetmed.hokudai.ac.jp E- 4, 7, 14 Oshima 5-11 I C-1 A-1 Sofjin KH I.persulcatus

2 Strain Year of isolation Geographical origin Source Accession no. Envelope 3'-NCR VL99-m11 Vladiostok AB AB KH99-m9 Khabarovsk AB D Khabarovsk Human brain AB IR99-1m1 Irkutsk (1)* AB AB IR99-1m4 Irkutsk (1) AB AB IR99-2m3 Irkutsk (2) AB IR99-2m7 Irkutsk (2) AB AB IR99-2f7 Irkutsk (2) AB IR99-2f13 Irkutsk (2) AB AB Oshima Oshima Dog blood AB AB Oshima I Oshima I. ovatus AB AB Oshima A Oshima A. speciosus AB AB KH Khabarovsk AB AB KH Khabarovsk AB AB KH Khabarovsk AB AB Sofjin-HO 1937 Primorsky Human brain AB AB ( )* : Virus isolation point number 4 3 louping ill VL99-m11 KH99-m9 D1283 Oshima Sofjin Vasilchenko Aina IR99-2f7-2f13VL99-m11 D1283 KH98-5 Ohima5-10

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pp.35 44 2005 39 1,000FFU 28 IR99-2f7 0%,IR99-2f13 10% VL99-m11 20% D1283 Oshima5 10 40% KH98-5 50% 3 5

5 pp ,000FFU 28 IR99-2f7 0%,IR99-2f13 10% VL99-m11 20% D1283 Oshima % KH % FFU IR99-2f7 IR99-2f13 VL99-m11 Oshima5-10 IR99-2f7 IR99-2f13 VL99-m11 Oshima D1283 KH % 20% 3. Oshima5-10 BHK 5 BHK Oshima Cl-1 Oshima 5-10 Oshima Cl-1 BHK Oshima Cl-1 9 Oshima Oshima Cl-1 1/100 Oshima Cl-1 Oshima ,000FFU 2 Oshima Cl-1 80% Oshima % 2 10FFU Oshima Cl-1 Oshima5-10

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7 pp Oshima Cl-1 Oshima FFU/ml FFU/g FFU/g Oshima Cl FFU/ml FFU/g 9 11 Oshima Cl-1 Oshima 5-10 Oshima Oshima Cl-1 Oshima Cl-1 Oshima NS5 20 E Oshima Cl-1 E E II 15 3, 10 GAGs 1, 11, 13 2 GAGs GAGs Oshima 5-10 BHK-21 GAGs Oshima Cl-1 GAGs E E

8 42 55

9 pp GAGs 4. cdna cdna Neudoerfl Hypr Vasilchenko 6, 12 Oshima5-10 cdna Oshima 5-10 cdna long high-fiedelity RT-PCR one-step cloning 9 cdna 0-1C 4 3 UTR 2 E- 1 Ns5 3 O-IC-pt BHK Oshima IC-pt 12 quasi-species 4 E- P 40 A 378 K 674 T 794 O-IC E-NS5-mt 4 50FFU Oshima 5-10 O-IC-pt % O-IC-pt Ns5 2 S 40 A 378 K 674 T 994 O-ICNS5-mt % S 40 A 378 K 674 T 994 E- 1 P 40 A 378 K 674 T 994 O-ICE-NS5-mt 25% E S40 P Hs5-V 378 A Ns5-R 674 K 5. 10, Bernard KA, Klimstra WB, Johnston RE.: Mutations in the E2 glycoprotein of Venezuelan equine encephalitis virus confer heparan sulfate interaction, low morbidity, and rapid clearance from flood of mice. Viriligy 276: , Calisher C, Karabatsos N, Dalrymple JM, Shope RE, Porterfield JS, Westaway EG, Brandt WE.: Antigenic relationships between flaviviruses as determimed by cross-neutralization tests with polyclonal antisera. J Gen Virol 30: 37-43, Chambers TJ, Hahn CS, Galler R, Rice CM.: Flavivirus genome organization, expression, and replication. Ann Rev Microbiol 44: , Gojobori T, Moriyama EN, Kimura M.: Molecular clock of viral evolution, and the neutral theory. Pro Nat Acad Sci 87: , Goto A, Hayasaka D, Yoshii K, Mizutani T, Kariwa H, Takashima I.: A BHK-21 cell culture-adapted tickborne encephalitis virus mutant is attenuated for neuroinvasiveness. Vaccine 21: , Gritsun TS, Gould EA.: Development and analysis of a tick-borne encephalitis virus infectious clone using a novel and rapid strategy. J Virol Methods 76: , Hayasaka D, Suzuki K, Kariwa H, Ivanov L, Volkov V, Demenev V, Mizutani T, Gojobori T, Takashima I.: Phylogenetic and virulence analysis of tick-borne encephalitis viruses from Japan and far-eastern Russia. J Gen Virol 80: ,. 8 Hayasaka D, Ivanov L, Leonova GN, Goto A, Yoshii K, Mizutani T, Kariwa H, Takashima I.: Distribution and characterization of tick-borne encephalitis viruses from Siberia and far-eastern Asia. J Gen Virol 82: , Hayasaka D, Gritsun TS Yoshii K, Ueki T, Goto A, Mizutani T, Kariwa H, Iwasaki T, Gould EA, Takashima I.: Amino acid changes responsible for attenuation of virus neurovirulence in an infectious cdna clone of the Oshima strain of tick-borne encephalitis virus. J Gen Virol 85: , Holzmann H, Stiasny K, Echer M, Kung C, Heinz FX.: Characterization of monoclonal antibody-escape mutants of tick-borne encephalitis virus with reduced neuroinvasiveness in mice. J Gen Virol 78: 31-37, Lee E, Lobigs M.: Substitutions at the putative receptor-binding site of an encephalitic flavivirus alter virulence and host cell tropism and reveal a role for glycosamaminoglycans in entry. J Virol 74: , Mandl CW, Ecker M, Holzmann H, Kunz C, Heinz F X.: Infectious cdna clones of tick-borne encephalitis virus European subtype prototypic strain Neudoerfl

10 44 55 pp and high virulence strain Hypr. J Gen Virol 78: , Mandl CW, Kroschewshi H, Allisan SL, Kofler R, Holzmann H, Meixner T.: Adaptation of tick-borne encephalitis virus to BHK-21 cells results in the formation of multiple heparan subfate binding sites in the envelope protein and attenuation in vivo. J Virol 75: , Nei M, Gojobori T.: Simple methods for estimating the numbers of synonymous and nonsynonymous nucleotide substitutions. Mol Biol Evol 3: , Rey FA, Heinz FX, Mandl C, Kunz C, Harrison SC.: The envelope glycoprotein from tick-borne encephalitis virus at 2 resolution. Nature 375: , Shope RE.: Medical significance of togaviruses: an overview of diseases caused by togaviruses in man and domestic and wild vertebrate animals. PP [Schlesinger, R.W. ed] In the togaviruses. Academic Press, New York, Takashima I, Morita K, Chiba M, Hayasaka D, Sato T, Takezawa C, Igarashi A, Kariwa H, Yoshimatsu K, Arikawa J, Hashimoto N.: A case of tick-borne encephalitis in Japan and isolation of the virus. J Clin Microbiol 35: , Takeda T, Ito T, Osada M, Takahashi K, Takashima I.: Isolation of tick-borne encephalitis virus from wild rodents and a seroepidemiological survey in Hokkaido, Japan. Am. J Trop Med Hyg, 60: Takeda T, Ito T, Chiba M, Takahashi K, Niioka T, Takashima I.: Isolation of tick-borne enxephalitis virus from Ixodes ovatus Acari:Ixodidae in Japan. J Med Entomol 35: , Wallner G, Mandl CW, Ecker M, Holzmann H, Stiasny K, Kunz C.: Characterization and complete genome sequences of high- and low-virulence variants of tickborne encephalitis viris. J Gen Virol 77: , Phylogenetic analysis and pathogenicity of tick-borne encephalitis virus from Japan and far-east Russia Ikuo Takashima, Daisuke Hayasaka, Akiko Goto, Kentarou Yoshii, Hiroaki Kariwa. Laboratory of Public Health, Department of Environmental Veterinary Sciences, Graduate School of Veterinary Medicine, Hokkaido University, Kita-18, Nishi-9, Kitaku, Sapporo , Japan Phylogenetic analysis of tick-borne encephalitis(tbe) virus revaled that Hokkaido strain of TBE virus evolved several hundreds years ago in far-east Russia. TBE virus strains in Irkutsk area were identified as Siberian subtype of TBE virus. BHK-cell adapted mutant of TBE virus showed lower neuro-invasive virulence in mice than parent virus. The mutant carried one amino acid substitution in envelope protein which resulted in increase of positive charge of the protein. The mutant-infected mice showed lower virus titers in bloods and spleens than the parent-infected mice. Infectious c-dna clone of TBE virus Hokkaido strain was successfully generated and was applied to examine the neurovirulence in mice. One amino acid change in envelope protein and 2 amino acid changes in Ns5 protein showed a synergistic effect on reduced neurovirulence in mice.

Distribution and characterization of tick-borne encephalitis viruses from Siberia and far-eastern Asia

Journal of General Virology (2001), 82, 1319 1328. Printed in Great Britain... Distribution and characterization of tick-borne encephalitis viruses from Siberia and far-eastern Asia Daisuke Hayasaka, 1