New lyssavirus genotype from the Lesser Mouse-eared Bat (Myotis blythi), Kyrghyzstan. (Research).The Aravan virus was isolated from a Lesser Mouse-eared Bat The Lesser Mouse-Eared Bat (Myotis blythii) is a species of vesper bat in the Vespertilionidae family. It can be found in the following countries: Afghanistan, Albania, Austria, Bangladesh, Bhutan, Bulgaria, China, France, Georgia, Germany, Greece, Hungary, Iran, (Myotis Myotis genus of bats. Includes M. thysanodes (fringed myotis bat), M. myotis (European common mouse-eared bat), M. lucifugus (little brown bat). blythi) in the Osh region of Kyrghyzstan, central Asia, in 1991. We determined the complete sequence of the nucleoprotein nucleoprotein Macromolecular complex consisting of a protein linked to a nucleic acid, either DNA or RNA. The proteins that combine with DNA are generally of characteristic types called histones and protamines. (N) gene and compared it with those of 26 representative lyssaviruses obtained from databases. The Aravan virus was distinguished from seven distinct genotypes on the basis of nucleotide and amino acid amino acid (əmē`nō), any one of a class of simple organic compounds containing carbon, hydrogen, oxygen, nitrogen, and in certain cases sulfur. These compounds are the building blocks of proteins. identity. Phylogenetic phy·lo·ge·net·ic adj. 1. Of or relating to phylogeny or phylogenetics. 2. Relating to or based on evolutionary development or history. analysis based on both nucleotide and amino acid sequences showed that the Aravan virus was more closely related to genotypes 4, 5, and--to a lesser extent--6, which circulates among insectivorus bats in Europe and Africa. The Aravan virus does not belong to any of the seven known genotypes of lyssaviruses, namely, rabies rabies (rā`bēz, ră`–) or hydrophobia (hī'drəfō`bēə), acute viral infection of the central nervous system in dogs, foxes, raccoons, skunks, bats, and other animals, and in , Lagos bat, Mokola, and Duvenhage viruses and European bat lyssavirus 1, European bat lyssavirus 2, and Australian bat lyssavirus
********** The Lyssavirus genus includes seven genotypes: rabies virus rabies virus n. A rather large, bullet-shaped virus of the genus Lyssavirus that causes rabies. (RABV, genotype 1), Lagos bat virus Lagos bat virus is a lyssavirus that causes a rabies-like illness in mammals in southern and central Africa. It was first isolated from a fruit bat (Eidolon helvum) from Lagos Island, Nigeria in 1956. (genotype 2), Mokola virus Mo·ko·la virus n. A rabies-related virus of the genus Lyssavirus found most commonly in Africa and causing a fatal neurological disease in humans and cats. (genotype 3), Duvenhage virus (genotype 4), European bat lyssavirus 1 (EBLV-1, genotype 5), European bat lyssavirus 2 (EBLV-2, genotype 6), and Australian bat lyssavirus (ABLV ABLV Air-Breathing Launch Vehicle ABLV American Bank of the Lehigh Valley (Allentown, Pennsylvania) ABLV Australian Bat Lyssa Virus , genotype 7) (1,2). Lagos bat virus was isolated from frugivorous frugivorous fruit-eating. bats (Eidolon ei·do·lon n. pl. ei·do·lons or ei·do·la 1. A phantom; an apparition. 2. An image of an ideal. [Greek eid helvum) in Nigeria in 1956 (3) and in 1974 from another bat (Micropterus pusillus) in the Central Africa Republic (4). Mokola virus was isolated from shrews (Crocidura sp.) and a child in Nigeria in 1968 (5,6), a girl in Nigeria in 1971 (7), and cats in Zimbabwe (8). Duvenhage virus was originally isolated from a human who died after being bitten by a bat in South Africa South Africa, Afrikaans Suid-Afrika, officially Republic of South Africa, republic (2005 est. pop. 44,344,000), 471,442 sq mi (1,221,037 sq km), S Africa. in 1970 (9) and from Miniopterus sp. bats in 1981 (10). EBLV-1 was isolated from bats (Eptesicus serotinus) in Germany in 1968 (11), in Poland in 1985 (12), in Denmark, Holland, and Spain in 1987, and in France in 1989 (13). Some isolates of EBLV-1 were obtained from bats in Ukraine and from one human case of bat origin in Russia in 1985 (14,15). EBLV-2 was isolated from a human in Finland in 1985 (16), and from bats in Holland, the Netherlands, Switzerland, and the U.K. EBLV-2 is mainly carried by bats of the Myotis genus (Myotis dasycneme and M. daubentonii) (17). ABLV was isolated from five species of flying fox bats, one species of an insectivorous insectivorous eating insects to the extent that they are significant as a contributor to the patient's diet. bat, and two infected humans in 1996 (1,18,19). Rabies viruses have been reported in Kazakhstan, central Asia (20). Terrestrial rabies viruses have been enzootic en·zo·ot·ic adj. Prevalent among or restricted to animals of a specific geographic area. Used of a disease. n. An enzootic disease. enzootic peculiar to or present constantly in a location. See also endemic. in all Central Asian countries and are mainly carried by dogs. Field rabies viruses have been isolated and characterized in Asia, specifically Pakistan, China, Indonesia, Thailand, the Philippines, Malaysia, India, and Sri Lanka Sri Lanka (srē läng`kə) [Sinhalese,=resplendent land], formerly Ceylon, ancient Taprobane, officially Democratic Socialist Republic of Sri Lanka, island republic (2005 est. pop. (21-26). Isolation of lyssaviruses from bats has been reported only in India and Thailand; however, these viruses were reported as RABV (27,28). Recently, Arguin et al. detected neutralizing antibodies against ABLV in the serum of six bat species (Mineopterus schreibersi, Taphozous melanopogan, Philetor brachypteus, Scotophilus kuhli, Pteropus hypomelanus, and Rousettus amplexicaudatus) in the Philippines (29). Aravan virus was originally isolated from the brain of a lesser mouse-eared bat (Myotis blythi) in Kyrghyzstan in 1991. The antigenic profile of the virus was analyzed by using two panels of antinucleocapsid (N) gene monoclonal antibodies developed at the Wistar Institute of Anatomy and Biology (USA) and the Central Veterinary Laboratory of Great Britain (Weybridge, U.K.) (30-32). These results demonstrated that the virus differed from rabies and serotypes 2 (Lagos bat virus), 3 (Mokola virus), 4 (Duvenhage virus), 5 (EBLV-1), and 6 (EBLV-2). Furthermore, 386 nucleotides (nt) of the N gene were determined from reverse transcription-polymerase chain reaction (RT-PCR RT-PCR reverse transcriptase-polymerase chain reaction. See PCR1. ) product. Phylogenetic analysis suggested that the Aravan virus did not belong to the rabies virus group (33). In the present study, we determined the entire coding region of the N protein of Aravan virus and evaluated the phylogenetic relationships with other members of the Lyssavirus genus. Materials and Methods Viruses Aravan virus was isolated from the brain of one lesser mouse-eared bat (Myotis blythi) during a survey of 269 bats collected in the Osh region of Kyrghyzstan from 1988 to 1992 (30,32). A direct fluorescent antibody Direct fluorescent antibody (DFA or dFA) is a laboratory test that uses antibodies tagged with fluorescent dye to detect the presence of microorganisms. This is the main test used to detect rabies in animals and requires the examination of brain tissue. test was conducted. Aravan virus-infected mouse brains were impressed on glass slides, air-dried, and fixed with acetone acetone (ăs`ĭtōn), dimethyl ketone (dīmĕth`əl kē`tōn), or 2-propanone (prō`pənōn), CH3COCH3 . To detect the lyssavirus antigen, specimens were stained with fluorescein isothiocyanate (FITC FITC fluorescein isothiocyanate; used as a fluorescent label for proteins, especially antibodies. )-labeled anti-rabies globulin globulin, any of a large family of proteins of a spherical or globular shape that are widely distributed throughout the plant and animal kingdoms. Many of them have been prepared in pure crystalline form. (BBL "Be back later." See digispeak. (chat) BBL - (I will) be back later. , Cockeysville, MD) or FITC-labeled anti-rabies monoclonal globulin (Centocor Inc., Malvern, PA). FITC-labeled anti-nucleoprotein monoclonal antibodies (NC-MAbs, W502) cross- reactive to lyssaviruses were also used (19). Amplification of Nucleoprotein cDNA and Direct Sequencing Total RNA RNA: see nucleic acid. RNA in full ribonucleic acid One of the two main types of nucleic acid (the other being DNA), which functions in cellular protein synthesis in all living cells and replaces DNA as the carrier of genetic was extracted from virus-infected mouse brain emulsions with a commercial reagent (RNeasy Mini Kit, QIAGEN, Germany). cDNA was obtained with a T-Primed First-Strand kit (Amersharm Biosciences Corporation, Piscataway, NJ). PCR PCR polymerase chain reaction. PCR abbr. polymerase chain reaction Polymerase chain reaction (PCR) amplification and sequencing of the N gene were performed by using the sense primer AraN-S01 (5'-ATGTACCACCTCTACAATGG-3', nt 55-74) and an antisense antisense, DNA or RNA manipulated in a laboratory so that its components (nucleotides) form a complementary copy of normal, or "sense," messenger RNA (mRNA; see nucleic acid). primer AraNC-1400 (5'-TC ATGCTCAATTGTAAAAC-3', nt 1456-1474). The cDNA template (2 [micro]L) was amplified by using primers (AraN-S01 and AraNC-1400), according to the manufacturer's instruction (Super Taq Premix premix a finite mixture of nutritional supplements such as minerals and vitamins, usually combined with a carrier and ready for mixing with a total ration. Kit, Sawady Technology, Tokyo, Japan). PCR reactions were incubated at 94[degrees]C for 2 min, subjected to 40 cycles of 94[degrees]C for 30 s, 48[degrees] for 20 s, and 68[degrees]C for 2 min, and a final extension at 68[degrees]C for 7 min in a DNA DNA: see nucleic acid. DNA or deoxyribonucleic acid One of two types of nucleic acid (the other is RNA); a complex organic compound found in all living cells and many viruses. It is the chemical substance of genes. thermal cycler (GeneAmpPCR System 9700 Applied ,Biosystems, Perkin-Elmer Corporation, Japan) (24,25). PCR products were purified by using a commercial kit (QIAquick PCR Purification Kit, QIAGEN). The sequences of the purified DNA products were determined on an automated sequencer See MIDI sequencer. (music) sequencer - Any system for recording and/or playback of music via a programmable memory which stores music not as audio data, but as some representation of notes. (ABI Abi (ā`bī) [short for Abijah], in the Bible, King Hezekiah's mother. (Application Binary Interface) A specification for a specific hardware platform combined with the operating system. model 310, Applied Biosystems, Foster City, CA) by using a PRIMS Ready Reaction Dyedeoxy Terminator Cycle Sequencing Kit (Applied Biosystems). Phylogenetic Analysis The 1350-nt and the deduced 450 amino acid (aa) sequences of the N gene of the Aravan virus were aligned with 26 lyssaviruses by using Clustal W program (34). A phylogenetic tree was constructed with the computer software MEGA 2 (35). Pairwise evolutionary nucleotide distances, including both transitions and transversions, were estimated according to Kimura's two-parameter method. Phylogenetic trees were constructed by the neighbor-joining method with 1,000 replicates to generate bootstrap See boot. (operating system, compiler) bootstrap - To load and initialise the operating system on a computer. Normally abbreviated to "boot". From the curious expression "to pull oneself up by one's bootstraps", one of the legendary feats of Baron von Munchhausen. probabilities at each node (36). Results and Discussion Direct Fluorescent Antibody Assay The three stains used in this study reacted against the Aravan virus infected mouse brain impressions. Fluorescence showed more scattered inclusions than those of the challenge virus standard in the acetone-fixed mouse brain smear (data not shown). The results confirmed that the Aravan virus is a lyssavirus. Nucleotide and Deduced Amino Acid Sequence Identities among the Aravan Virus and Other Lyssaviruses The 1350-nt and the deduced 450 aa sequences of the Aravan virus were compared with 26 representative lyssaviruses belonging to seven genotypes (Table 1). We selected 16 representative rabies variants from the eight diverse groups, including rabies variants from geographic areas of Asia near Kyrghyzstan and from bats and raccoons in North and South America (25,37). The nucleotide and amino acid sequence identities among all 27 lyssaviruses, including Aravan virus, were calculated. Then genotype 1 was represented by seven rabies viruses (SRL 1. SRL - Bharat Jayaraman. ["Towards a Broader Basis for Logic Programming", B. Jayaraman, TR CS Dept, SUNY Buffalo, 1990]. 2. SRL - Schema Representation language. 3. SRL - Structured Robot Language. C. Blume & W. Jacob, U Karlsruhe. 1032, 86118BRE (Business Rules Engine) Software that automates policies and procedures within an organization, whether legal, internal or operational. The use of a rules engine (BRE) requires placing the company rules in an external repository that can be easily reviewed rather than , 1500AFS A distributed file system for large, widely dispersed Unix and Windows networks from Transarc Corporation, now part of IBM. It is noted for its ease of administration and expandability and stems from Carnegie-Mellon's Andrew File System. AFS - Andrew File System , 9218TCH TCH Trans-Canada Highway TCH Texas Children's Hospital (Houston, TX) TCH The Children's Hospital TCH Traffic Channel TCH Threshold Crossing Height TCH Toyota Camry Hybrid TCH Taurocholic Acid , 8738THA THA Total hip arthroplasty. See Total hip replacement. , insectivorous bat/Chile, and PA R89), and genotypes 2, 3, 4, 5, 6, and 7 were represented by Lagos bat virus (8619NGA Noun 1. NGA - a combat support agency that provides geographic intelligence in support of national security National Geospatial-Intelligence Agency ), Mokola virus (MOK/U22843), Duvenhage virus (86132AS), EBLV-1 (8918FRA Fra: see Angelico, Fra; Bartolommeo di Pagholo del Fattorino, Fra; Fra Filippo Lippi under Lippi. ), EBLV-2 (9007FIN), and ABLV (Balina/AF006497), respectively (Table 2). The nucleotide sequence identity of Aravan virus with the genotypes 4, 5, 6, and 7 was 77% to 78%; with genotype 1, 75% to 77%; and with genotypes 2 and 3, 72% to 74%. The most extensive nucleotide sequence differences among isolates of genotype 1 were between the raccoon raccoon, nocturnal New World mammal of the genus Procyon. The common raccoon of North America, Procyon lotor, also called coon, is found from S Canada to South America, except in parts of the Rocky Mts. and in deserts. isolate (PA R89) and the African and Asian isolates (82.8% to 82.9% identity). The Aravan virus demonstrated 92% aa sequence identity with genotypes 4, 5, and 7; 89% with genotype 6; and 81% to 85% with genotypes 2 and 3. The maximum variation of amino acid sequences within genotype 1 was exhibited between a vampire bat isolate from Brazil and an African isolate (93.1% to 93.3% identity). Genotype 4 (Duvenhage virus) was most closely related to genotype 5 (EBLV-1) with nucleotide and amino acid sequence identities of 79.8% and 93.3 %, respectively. ABLV (genotype 7) was closely related to SRL1032 (genotype 1, Sri Lankan rabies virus) with a 93.1% aa sequence identity. These values were almost same as maximum variation of genotype 1. Based on our present data, we determined that isolates sharing <79.8% nt and 93.1% to 93.3% aa sequence identities belonged to different genotypes. In several studies, thresholds of <80% nt and 92% or 93% aa sequence identities warranted the proposal of a new genotype (1,23,38). Hence, the nucleotide and amino acid percentage identity values demonstrated that Aravan virus should be regarded as a new lyssavirus genotype. Phylogenetic Analysis A phylogenetic tree of 27 lyssaviruses, including the Aravan virus, based on the 1350-nt sequence of the N gene was constructed by using the vesicular vesicular /ve·sic·u·lar/ (ve-sik´u-ler) 1. composed of or relating to small, saclike bodies. 2. pertaining to or made up of vesicles on the skin. 3. stomatitis Stomatitis Definition Inflammation of the mucous lining of any of the structures in the mouth, which may involve the cheeks, gums, tongue, lips, and roof or floor of the mouth. Indiana virus (VSIV, tsW16B/U B/U Backup B/U Business Unit B/U Broken Up 13898) as an outgroup (Figure, a). The lyssaviruses divided into two groups: one group consisted of genotypes 2 and 3, and the other consisted of genotypes 1, 4, 5, 6, 7, and the Aravan virus. The latter group was divided into six distinct clusters corresponding to genotypes 1, 7, 6, and 5 (high bootstrap values of 98%, 99%, 100%, and 100%, respectively), then Aravan virus and genotype 4. Moreover, the Aravan virus clustered with genotypes 4, 5, and 6 (low bootstrap value of 59%). Duvenhage virus (genotype 4) and EBLV-1 (genotype 5) formed the same cluster (high bootstrap value of 91%), and are therefore closely related. The Aravan virus occupied the phylogenetic position between genotype 6 and the cluster of genotypes 4 and 5. We also constructed a phylogenetic tree based on the deduced 450-aa sequences of the N gene (Figure,b). Similar to the nucleotide data, the amino acid sequences divided into two large groups and further subdivided into eight groups. One group consisted of genotypes 2 and 3 (bootstrap value of 89%), and the other group consisted of genotypes 1, 7, 6, 4, and 5, and the Aravan virus (high bootstrap value of 100%). The latter group had three distinct clusters corresponding to genotypes 1, 7, and 6 (high bootstrap values of 100%, 99%, and 100%, respectively), genotypes 4 and 5 (same cluster with a high bootstrap value of 98%), and the Aravan virus. The Aravan virus did not group with any other genotypes and is located at a position close to the cluster of genotypes 4 and 5 (bootstrap value of 66%). [FIGURE OMITTED] These results, along with those in Table 2 and the Figure, suggest that the Aravan virus does not belong to any of the seven Lyssavirus genotypes (rabies, Lagos bat, Mokola, Duvenhage, EBLV-1, EBLV-2, and ABLV). Thus, we propose that the Aravan virus forms an independent cluster and is a new member of the Lyssavirus genus. In this article, we have reported the first lyssavirus distinct from rabies virus originating on the Asian continent. The Aravan virus was more closely related to genotypes 4, 5, and, to a lesser extent, 6, which circulates among insectivorus bats in Europe and Africa. The lesser mouse-eared bat, from which the Aravan virus was isolated, is widely distributed in northern Africa, the Mediterranean, southern Europe, Crimea, Caucasus, Palestine, southwest Asia, and parts of central and eastern Asia. This information should be considered in the discussion of lyssavirus classification and evolution, as it suggests the possibility of a broader geographic distribution of the Aravan virus. We have no information about human rabies caused by bat exposure from central Asia, and rabies surveillance in this area is not known well. Based on this information and the virus' misdiagnoses as rabies, we consider that transmission of Aravan virus to humans is possible. Indeed, this finding stimulates interest in new genotypes of lyssaviruses and is important from the viewpoint of public health, necessitating further lyssavirus surveillance of bats on the Asian continent.
Table 1. Lyssavirus isolates used in this study
Yr
Genotype (a) isolated Virus (strain) Country of isolation
1 (Rabies) ? CTN China
1 (Rabies) 1983 8738THA Thailand
1 (Rabies) ? ? India
1 (Rabies) 1996 SRL1032 Sri Lanka
1 (Rabies) 1992 9218TCH Chad
1 (Rabies) 1988 9141RUS Russia
1 (Rabies) ? 9196FX Canada
1 (Rabies) 1987 1500AFS Rep. South Afr.
1 (Rabies) 1985 9142EST Estonia
1 (Rabies) 1986 8681IRA Iran
1 (Rabies) 1985 86118BRE Brazil
1 (Rabies) 1992 BBCAN Canada
1 (Rabies) 1992 MYCAN Canada
1 (Rabies) ? ? Chile
1 (Rabies) 1988 Insectivorous bat Chile
1 (Rabies) 1989 PA R89 USA
2 (Lagos bat) 1958 8619NGA Nigeria
3 (Mokola) ? Y09762 ?
3 (Mokola) 1981 MOK Zimbabwe
4 (Duvenhage) 1986 86132AS Rep. South Africa
5 (EBLV-1) 1985 8615POL Poland
5 (EBLV-1) 1989 8918FRA France
6 (EBLV-2) 1986 9007FIN Finland
6 (EBLV-2) 1986 9018HOL Holand
7 (ABLV) 1996 Ballina Australia
7 (ABLV) 1996 Insectivorous isolate Australia
? 1991 Aravan Kyrghystan
Yr
Genotype (a) isolated Host Accession no.
1 (Rabies) ? ? AF367863
1 (Rabies) 1983 Human U22653
1 (Rabies) ? ? AF374721
1 (Rabies) 1996 Jackal AB041964
1 (Rabies) 1992 Dog U22644
1 (Rabies) 1988 Arctic fox U22656
1 (Rabies) ? Vulpes vulpes L20676
1 (Rabies) 1987 Yellow mongoose U22628
1 (Rabies) 1985 Racoon dog U22476
1 (Rabies) 1986 Dog U22482
1 (Rabies) 1985 Vampire bat U22479
1 (Rabies) 1992 Eptesicus fuscus AF351833
1 (Rabies) 1992 Myotis lucifugus AF351839
1 (Rabies) ? Tadarida brasiliensis AF070450
1 (Rabies) 1988 Insectivorous bat AF351850
1 (Rabies) 1989 Raccoon U27221
2 (Lagos bat) 1958 Eidolon helvum U22842
3 (Mokola) ? ? Y09762
3 (Mokola) 1981 Cat U22843
4 (Duvenhage) 1986 Human U22848
5 (EBLV-1) 1985 Eptesicus serotinus U22844
5 (EBLV-1) 1989 E. serotinus U22845
6 (EBLV-2) 1986 Human U22846
6 (EBLV-2) 1986 M. dasycneme U22847
7 (ABLV) 1996 Pteropid alecto AF006497
7 (ABLV) 1996 Insectivorous bat AF081020
? 1991 M. blythi AB094438
(a) EBLV-1, European bat lyssavirus 1; EBLV-2, European bat lyssavirus
2; ABLV, Australian bat lyssavirus; MOK, strain name in Mokola virus
Table 2. Comparison of nucleotide and deduced amino acid
sequences of Aravan virus with other 13 lyssaviruses
Nucleotide sequence identity (%)
Amino acid
sequence Genotype 1 (rabies virus)
identity
(%) Aravan SRL1032 U22479BRE U22628AFS U22644CHAD
Aravan 100.0 75.6 76.0 76.2 74.8
SRL1032 90.9 100.0 84.7 85.9 86.9
U22479BRE 88.2 95.3 100.0 83.3 83.0 (a)
U22628AFS 89.3 96.7 93.3 (b) 100.0 85.6
U22644CHAD 88.7 96.9 93.1 (b) 94.4 100.0
U22653THA 89.8 97.1 94.2 95.1 95.3
AF351850 90.0 97.1 95.8 95.8 94.9
U27221 89.6 95.8 93.6 94.2 93.8
LBU22842 84.7 82.9 81.8 81.1 80.4
MKU22843 80.9 78.2 77.6 77.3 76.2
86132AS 91.8 88.9 86.9 87.6 87.1
8918FRA 92.0 88.9 86.7 87.8 87.8
9007FIN 88.9 88.0 86.2 87.1 86.9
AF006497 92.0 93.1 (d) 91.1 91.6 91.1
Nucleotide sequence identity (%)
Amino acid
sequence Genotype 1 (rabies virus)
identity
(%) Genotype 2 Genotype 3
U22653THA AF351850 U27221 LBU22842 MKU22843
Aravan 75.9 77.0 76.2 74.3 72.4
SRL1032 86.8 86.6 84.1 74.2 70.2
U22479BRE 83.3 89.9 83.7 73.8 69.6
U22628AFS 83.5 83.6 83.2 73.3 70.4
U22644CHAD 86.0 84.0 82.9 (a) 73.0 69.0
U22653THA 100.0 84.2 82.8 (a) 73.4 69.7
AF351850 95.6 100.0 86.1 73.5 70.1
U27221 94.9 95.1 100.0 73.1 70.2
LBU22842 81.8 83.1 82.2 100.0 74.8
MKU22843 77.6 78.4 77.8 84.4 100.0
86132AS 87.6 88.7 87.8 85.8 80.7
8918FRA 88.4 88.2 88.2 83.8 79.1
9007FIN 88.0 87.3 87.3 79.1 76.2
AF006497 91.8 91.8 91.3 82.0 79.8
Amino acid
sequence
identity Genotype 4 Genotype 5 Genotype 6 Genotype 7
(%)
86132AS 8918FRA 9007FIN AF006497
Aravan 78.2 77.9 77.2 76.9
SRL1032 73.9 75.6 74.7 78.0
U22479BRE 73.8 75.3 74.2 77.2
U22628AFS 73.9 75.3 74.6 78.0
U22644CHAD 73.1 75.0 74.8 77.2
U22653THA 74.4 75.9 74.7 77.3
AF351850 74.3 75.0 74.8 77.4
U27221 74.2 74.6 75.7 77.1
LBU22842 73.4 74.4 72.5 72.6
MKU22843 71.6 69.9 69.2 71.0
86132AS 100.0 79.8 (c) 75.9 77.0
8918FRA 93.3 (d) 100.0 78.0 76.9
9007FIN 86.2 88.0 100.0 77.2
AF006497 90.0 89.8 87.8 100.0
(a) The values were shown as maximum variation of
nucleotide sequence identities (%) within genotype 1.
(b) The values were shown as maximum variation of
amino acid sequence identities (%) within genotype 1.
(c) Thresholds of nucleotide sequence
identities % as different genotypes.
(d) Thresholds of amino acid sequence
identities % as different genotypes.
Acknowledgments We thank C. E. Rupprecht for providing the fluorescein/ isothiocyanate-labeled W502 and I. Kurane for helpful advice in preparing the manuscript. We also thank an anonymous reviewer, whose constructive criticism and many useful suggestions improved the manuscript. The work was partly supported by the Russian Foundation for Basic Research, grant (00-04-48004). References (1.) Gould AR, Hyatt AD, Lunt R, Kattenbelt JA, Hengstberger S, Blacksell SD. Characterization of a novel lyssavirus isolated from Pteropid bats in Australia. Virus Res 1998;54:165-87. (2.) Badrane H, Bahloul C, Perrin P, Tordo N. Evidence of two lyssavirus phylogroups with distinct pathogenicity and immunogenicity immunogenicity /im·mu·no·ge·nic·i·ty/ (-je-nis´it-e) the property enabling a substance to provoke an immune response, or the degree to which a substance possesses this property. . J Virol 2001;75:3268-76. (3.) Boulger LR, Porterfield JS. Isolation of a virus from Nigerian fruit bats. 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(22.) Mccoll KA, Gould AR, Selleck PW, Hooper PT, Westbury HA, Smith JS. Polymerase chain reaction polymerase chain reaction (pŏl`ĭmərās') (PCR), laboratory process in which a particular DNA segment from a mixture of DNA chains is rapidly replicated, producing a large, readily analyzed sample of a piece of DNA; the process is and other laboratory techniques in the diagnosis of long incubation rabies in Australia. Aust Vet J 1993;70:84-9. (23.) Kissi B, Tordo N, Bourhy H. Genetic polymorphism polymorphism, of minerals, property of crystallizing in two or more distinct forms. Calcium carbonate is dimorphous (two forms), crystallizing as calcite or aragonite. Titanium dioxide is trimorphous; its three forms are brookite, anatase (or octahedrite), and rutile. in the rabies virus nucleoprotein gene. Virology virology, study of viruses and their role in disease. 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Molecular epidemiology molecular epidemiology Molecular medicine An evolving field that combines the tools of standard epidemiology–case studies, questionnaires and monitoring of exposure to external factors with the tools of molecular biology–eg, restriction endonucleases, of rabies in Thailand. Microbiol Immunol 1999;43:551-9. (27.) Pal SR, Arora B, Chuttani PN, Broor S, Choudhury S, Joshi RM, et al. Rabies virus infection of a flying fox bat, Pterous policephalus in Chandigarh, North India. Trop Geogr Med 1980;32:265-7. (28.) Smith PC, Lawhaswasdi K, Vick WE, Stanton JS. Isolation of rabies virus from fruit bats in Thailand. Nature 1967;216:384. (29.) Arguin PM, Murray-Lillibridge K, Miranda MEG, Smith JS, Calaor AB, Rupprecht CE. Serologic se·rol·o·gy n. pl. se·rol·o·gies 1. The science that deals with the properties and reactions of serums, especially blood serum. 2. evidence of lyssavirus infections among bats, the Philippines. Emerg Infect Dis 2002;8:258-62. (30.) Kuzmin IV, Botvinkin AD, Rybin SN, Bayaliev AB. Lyssavirus with usual antigenic structure isolated from bat in Southern Kyrghyzstan. Vopr Virusol 1992;37:256-9 (in Russian.). (31.) Kuzmin IV, Botvinkin AD. The behaviour of bats Pipistrellus pipistrellus after experimental inoculation inoculation, in medicine, introduction of a preparation into the tissues or fluids of the body for the purpose of preventing or curing certain diseases. The preparation is usually a weakened culture of the agent causing the disease, as in vaccination against with rabies and rabies-like viruses rabies-like viruses Mokola, Lagos bat, Duvenhage, European and Australian bat lyssaviruse. So called because they cause rabies-like disease in humans and animals. and some aspects of the pathogenesis. Myotis 1996;34:939. (32.) Botvinkin AD, Kuzmin IV, Rybin SN. The unusual bat lyssavirus Aravan from Central Asia. Myotis 1996;34:101-4. (33.) Arai YT, Kuzmin IV, Kurane I. Phylogenetic analysis of Aravan virus isolated from the bat in Central Asia: natural foci infections of human. Proceedings of the Conference on the 80th Anniversary of the Institute for Natural Foci Infections, Omsk, Russia, 2001. p. 65-9. (34.) Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG. The Clustal X windows interface: flexible strategies for multiple sequence alignment A multiple sequence alignment (MSA) is a sequence alignment of three or more biological sequences, generally protein, DNA, or RNA. In general, the input set of query sequences are assumed to have an evolutionary relationship by which they share a lineage and are descended from a aided by quality analysis tools. Nucleic Acids Nucleic acids The cellular molecules DNA and RNA that act as coded instructions for the production of proteins and are copied for transmission of inherited traits. Res 1997;25:4876-82. (35.) Kumar S, Tamura K, Jakobsen IB, Nei M. MEGA2: Molecular evolutionary genetics analysis software. Tempe (AZ): Arizona State University Arizona State University, at Tempe; coeducational; opened 1886 as a normal school, became 1925 Tempe State Teachers College, renamed 1945 Arizona State College at Tempe. Its present name was adopted in 1958. ; 2001. (36.) Saitou N, Nei M. The neighbor-joining method: a new method for reconstructing phylogenetic trees. Mol Biol Evol 1987;4:406-25. (37.) Badrane H, Tordo N. Host switching in Lyssavirus history from the Chroptera to the Carnivora orders. J Virol 2001;75:8096-104. (38.) Bourhy H, Kissi B, Tordo N. Molecular diversity of the Lyssavirus genus. Virology 1993;194: 70-81. Address for correspondence: Yohko T. Arai, Department of Virology 1, National Institute of Infectious Diseases, Toyama 1-23-1, Shinjuku-ku, Tokyo 162, Japan; fax: +813-52851188; e-mail: yarai@nih.go.jp Yohko T. Arai, * Ivan V. Kuzmin, ([dagger]) Yosuke Kameoka, * and Alexandr D. Botvinkin ([double dagger]) * National Institute of Infectious Diseases, Tokyo, Japan; ([dagger]) Institute for Natural Foci Infections, Omsk, Russia; and ([double dagger]) Antiplague Research Institute of Siberia and the Far East, Irkutsk, Russia Dr. Arai is a research scientist at Department of Virology 1, National Institute of Infectious Diseases, Tokyo, Japan. Her research interests focus on the molecular epidemiology of lyssaviruses and the quality control of rabies vaccines. |
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