spring viraemia of carp virus
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- spring viraemia of carp virus
International Common Names
- English: rhabdovirus carpio; spring viremia of carp virus
Taxonomic TreeTop of page
- Domain: Virus
- Group: "Positive sense ssRNA viruses"
- Group: "RNA viruses"
- Order: Mononegavirales
- Family: Rhabdoviridae
- Genus: Vesiculovirus
- Species: spring viraemia of carp virus
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Botswana||Absent, No presence record(s)|
|Burundi||Absent, No presence record(s)|
|Cabo Verde||Absent, No presence record(s)|
|Cameroon||Absent, No presence record(s)|
|Central African Republic||Absent, No presence record(s)|
|Congo, Democratic Republic of the||Absent, No presence record(s)|
|Djibouti||Absent, No presence record(s)|
|Egypt||Absent, No presence record(s)|
|Eritrea||Absent, No presence record(s)|
|Eswatini||Absent, No presence record(s)|
|Ethiopia||Absent, No presence record(s)|
|Ghana||Absent, No presence record(s)|
|Kenya||Absent, No presence record(s)|
|Madagascar||Absent, No presence record(s)|
|Mauritius||Absent, No presence record(s)|
|South Africa||Absent, No presence record(s)|
|Sudan||Absent, No presence record(s)|
|Togo||Absent, No presence record(s)|
|Tunisia||Absent, No presence record(s)|
|Uganda||Absent, No presence record(s)|
|Zimbabwe||Absent, No presence record(s)|
|Bahrain||Absent, No presence record(s)|
|Brunei||Absent, No presence record(s)|
|Georgia||Absent, No presence record(s)|
|Hong Kong||Absent, No presence record(s)|
|Indonesia||Absent, No presence record(s)|
|Iran||Absent, No presence record(s)|
|Japan||Absent, No presence record(s)|
|Kazakhstan||Absent, No presence record(s)|
|Lebanon||Absent, No presence record(s)|
|-Peninsular Malaysia||Absent, No presence record(s)|
|Mongolia||Absent, No presence record(s)|
|North Korea||Absent, No presence record(s)|
|Singapore||Absent, No presence record(s)|
|South Korea||Absent, No presence record(s)|
|Sri Lanka||Absent, No presence record(s)|
|Syria||Absent, No presence record(s)|
|Taiwan||Absent, No presence record(s)|
|Thailand||Absent, No presence record(s)|
|Turkmenistan||Absent, No presence record(s)|
|Uzbekistan||Absent, No presence record(s)|
|Andorra||Absent, No presence record(s)|
|Bosnia and Herzegovina||Absent, No presence record(s)|
|Bulgaria||Absent, No presence record(s)|
|Cyprus||Absent, No presence record(s)|
|Estonia||Absent, No presence record(s)|
|Finland||Absent, No presence record(s)|
|Greece||Absent, No presence record(s)|
|Iceland||Absent, No presence record(s)|
|Ireland||Absent, No presence record(s)|
|Isle of Man||Absent, No presence record(s)|
|Jersey||Absent, No presence record(s)|
|Latvia||Absent, No presence record(s)|
|Liechtenstein||Absent, No presence record(s)|
|Luxembourg||Absent, No presence record(s)|
|Malta||Absent, No presence record(s)|
|Norway||Absent, No presence record(s)|
|Portugal||Absent, No presence record(s)|
|Romania||Absent, No presence record(s)|
|Sweden||Absent, No presence record(s)|
|Ukraine||Absent, No presence record(s)|
|-Northern Ireland||Absent, No presence record(s)|
|Barbados||Absent, No presence record(s)|
|Belize||Absent, No presence record(s)|
|Bermuda||Absent, No presence record(s)|
|British Virgin Islands||Absent, No presence record(s)|
|Canada||Absent, No presence record(s)|
|Cayman Islands||Absent, No presence record(s)|
|Costa Rica||Absent, No presence record(s)|
|Cuba||Absent, No presence record(s)|
|Dominica||Absent, No presence record(s)|
|Dominican Republic||Absent, No presence record(s)|
|Guatemala||Absent, No presence record(s)|
|Haiti||Absent, No presence record(s)|
|Honduras||Absent, No presence record(s)|
|Jamaica||Absent, No presence record(s)|
|Martinique||Absent, No presence record(s)|
|Mexico||Absent, No presence record(s)|
|Nicaragua||Absent, No presence record(s)|
|Panama||Absent, No presence record(s)|
|Saint Kitts and Nevis||Absent, No presence record(s)|
|Saint Vincent and the Grenadines||Absent, No presence record(s)|
|Trinidad and Tobago||Absent, No presence record(s)|
|Australia||Absent, No presence record(s)|
|French Polynesia||Absent, No presence record(s)|
|New Zealand||Absent, No presence record(s)|
|Vanuatu||Absent, No presence record(s)|
|Argentina||Absent, No presence record(s)|
|Bolivia||Absent, No presence record(s)|
|Brazil||Absent, No presence record(s)|
|Chile||Absent, No presence record(s)|
|Colombia||Absent, No presence record(s)|
|Falkland Islands||Absent, No presence record(s)|
|Guyana||Absent, No presence record(s)|
|Paraguay||Absent, No presence record(s)|
|Peru||Absent, No presence record(s)|
|Uruguay||Absent, No presence record(s)|
|Venezuela||Absent, No presence record(s)|
Pathogen CharacteristicsTop of page
The aetiological role of a virus in SVC was confirmed by Baudouy (1975), Rudikov et al. (1975), Ahne (1977), Hill (1977), Tesarcík et al. (1977), Bèkèsi and Szabó (1979), Bucke and Finlay (1979), and Osadcaja and Rudenko (1981). Ahne (1973) and Bachmann and Ahne (1973, 1974) reported a rhabdovirus from carp with clinical signs of swim bladder inflammation (SBI), established its pathogenicity and named it SBI virus. They noted its serological similarity with Rhabdovirus carpio (Bachmann and Ahne, 1974). The virus induced haemorrhage and inflammation of the swim-bladder, a typical clinical sign in the initial stages of SBI. However, serological comparison by de Kinkelin and Le Berre (1974) and Hill (1975) showed the SBI virus to be indistinguishable from Rhabdovirus carpio. Furthermore, Krizanac et al. (1981) could not isolate virus from typical (Arshaniza and Bauer, 1973) cases of SBI.
Spring viraemia of carp virus is 60-90 nm wide and 90-180 nm long. It has the typical bullet form of the family Rhabdoviridae. Virions bear a regular array of spicules on the surface. They pass through filters of 450 and 200 nm porosity, but are retained by 100 nm membranes. The inner nucleocapsid has helical symmetry, consists of a ribonucleic acid (RNA)-protein (polymerase [L], nucleocapsid [N] and non-structural [NS]) complex and measures about 50 nm in diameter. The nucleocapsid is surrounded by a lipid-containing envelope (matrix [M] protein) with spikes (glycoprotein [G protein]).
Defective interfering particles of SVCV (de Kinkelin and Le Berre, 1974) usually appear in cell cultures infected with a high virus input. Production of normal virions in cell culture can be secured by dilution of inocula to ensure a low multiplicity of infection. These particles measure about two-thirds of the length of infective virions (Bishop and Smith, 1977).
The purified SVCV has a buoyant density of 1.195-1.200 g ml-1 in caesium chloride (Ahne, 1973; Bachmann and Ahne, 1973) and 1.16 g ml-1 in a linear 15-60% sucrose gradient (Lenoir, 1973). The single-stranded (ss) linear RNA molecule sediments at 38-40 S in a 5-25% sucrose gradient (Hill, 1975). Structural proteins of SVCV are similar to those of vesicular stomatitis virus (VSV) (Lenoir, 1973; Sokol et al., 1974; Roy and Clewley, 1978a; Deuter et al., 1982). In these studies the molecular mass for L protein varied from 90 to 190 kDa, for the G protein from 70 to 88 kDa, for the N protein from 40 to 52 kDa, for the NS protein from 43 to 53 kDa and for the M protein from 19 to 27 kDa. The RNA polymerase of the virus is RNA-dependent, with optimum in vitro enzymatic activity at 22°C; its 5' nucleotide is pppAP (Roy and Clewley, 1978b). The glycoprotein on the virion's surface is responsible for its infectivity and immunogenicity (Bishop and Smith, 1977). The NS protein contains two related phosphoproteins, NS1 and NS2 (Roy, 1981). Studies on the 5'-terminal structure of RNA (Gupta et al., 1979), the mechanisms of messenger RNA (mRNA) capping (Gupta and Roy, 1980, 1981), the base sequence at the 3¢ end of the RNA (Roy et al., 1984) and the sequence analysis of mRNA coding for the M protein (Kiuchi and Roy, 1984) have elucidated properties and mechanism of synthesis of SVCV RNA and its similarities with VSV. They indicate a common ancestor of SVCV and VSV (Kiuchi and Roy, 1984). A close relationship between SVCV and VSV was confirmed by comparison of the nucleotide sequences of their L- and G-genes, their gene junctions and their transcription initiation sequences and transcription/polyadenylation sequences (Björklund et al., 1995, 1996). The M protein of SVCV, like that of VSV, completely blocks the transport of snRNAs, spliced mRNAs and snRNPs, as well as slowing the transport of other molecules through nuclear pore complexes; the effect of this is uncertain, but may reduce the host cell's ability to produce interferons following virus infection (Petersen et al., 2002). The virus lacks the non-virion (NV) gene which exists in three lyssa-type fish rhabdoviruses (Kurath et al., 1994, 1997) assigned to the recently created genus Novirhabdovirus (van Regenmortel et al., 2000). The complete genomic sequence of SVCV has been determined by Hoffmann et al. (2002). The genome comprises 111,019 nucleotides, which is consistent with the predicted size of rhabdovirus genomes, and contains 5 open reading frames coding for the N, P, M, G, and L genes. The deduced aminio acid sequences of SVCV were compared with deduced sequences of other rhabdoviruses, and the highest homologies were with VSV, confirming the placement of SVCV in the Vesiculovirus genus.
Spring viraemia of carp virus is inactivated by lipid solvents, heating (60°C for 15 min), glycerol, ozone and diethylpyrocarbonate, as well as by pH below 4 and above 10. Formalin (3%), sodium hydroxide (NaOH) (2%), chlorine (500 mg L-1), actomar (0.01% I2), gamma irradiation (103 krads) and ultraviolet (UV) irradiation (254 nm) inactivate the virus within 10 min (Ahne, 1982a,b). Normal handling in the laboratory does not cause undue loss of infectivity. Serum concentrations of 2 or 5% have a pronounced protective effect on the infectivity during storage at 4°C and at room and freezing temperatures, freeze-thaw cycles and lyophilization (Ahne, 1973; de Kinkelin and Le Berre, 1974). The infectivity is retained in tap water at 10°C, in mud (pH 7.4) at 4°C for 42 days, in stream water at 10°C for 14 days and after drying at 4-21°C for 21 days (Ahne, 1982a,b). The SVCV adsorbs to the plasma membrane and enters the host cell by receptor-mediated endocytosis. The first sign of viral replication is the formation of inclusion bodies in the cytoplasm. A budding process at the plasma membrane and, later in infection, at membranes of dilated Golgi vesicles secure maturation and release of virus (Granzow et al., 1997).
The virus replicates in a variety of fish and other vertebrate primary cell cultures and cell lines, causing a clear cytopathic effect. The best systems for viral replication are cell lines from cyprinid fishes, such as Epithelioma papulosum cyprini (EPC) (Fijan et al., 1983), fathead minnow (FHM) (Gravell and Malsberger, 1965) and carp leucocyte culture (CLC) (Faisal and Ahne, 1990) cells. Each produces high virus yields (108-109 tissue culture infective dose at 50% end-point (TCID50 ml-1). Channel catfish ovary (CCO) cells (Bowser and Plumb, 1980a,b) are also quite susceptible to SVCV (N. Fijan, unpublished data). A number of other widely used fish cell lines, such as bluegill fry (BF-2), brown bullhead (BB) and rainbow trout gonad (RTG-2), also support replication of SVCV, but the yields are somewhat or much lower. Chicken embryo cells and the mammalian cell lines fetal calf kidney, pig kidney, baby hamster kidney (BHK/21), Vero, MDCK, SK (Ahne, 1973; Bachmann and Ahne, 1974), human diploid lung (WI-38) and several reptilian cell lines (Clark and Soriano, 1974), are also susceptible if incubated at 20-22°C. SVCV caused apoptosis in EPC cells, as demonstrated by changes in cell morphology and DNA fragmentation (Björklund et al., 1997). Apoptosis was inhibited by human endogenous acid cysteine proteinase inhibitor. Zebrafish liver (ZFL) cells were used in molecular and functional studies on a zebrafish interferon gene (Altmann et al., 2003), using SVCV as challenge virus. ZFL cells transfected with zebrafish interferon were partially protected against infection with SVCV. The effect of pesticides (atrazine and lindane) on the growth of SVCV in EPC cell cultures was investigated, as they could possibly contaminate fish by run-off from land into water courses. However, they were found not to influence virus titres (Cossarini-Dunier and Hattenberger, 1988).
Replication in cell cultures takes place between 4 and 32°C, with the optimum at 20-22°C. The cytopathic effect is characterized by rounding, detachment and lysis of cells. Margination of nuclear chromatin and cytoplasmic vacuolation precede rounding of cells and can be seen in fixed and stained cell sheets. Fathead minnow cells incubated at 20°C synthesize first progeny virus 4-6 h after infection and peak titres of both cell-associated and cell-free virus are reached between 10 and 22 h. One growth cycle lasts 8-10 h (Bachmann and Ahne, 1974; de Kinkelin and Le Berre, 1974). Well-defined plaques are formed after 3 days at 20°C.
Antigenic studies with rabbit polyclonal neutralizing antibodies indicate that all isolates examined belong to a single serotype, although Bachmannn and Ahne (1974) suggested that one isolate from carp may represent a serological sub-type of SVCV. Dikkeboom et al. (2004) reported that an SVCV isolate from the USA did not react in an immunocytochemical test with an antiserum prepared against a European isolate, and was not neutralized to the same degree as a European isolate, but it did cross-react strongly in an enzyme-linked immunosorbent assay (ELISA). Spring viraemia of carp virus is serologically unrelated to salmonid rhabdoviruses which cause viral haemorrhagic septicaemia (VHS virus) and infectious haematopoietic necrosis (IHN virus) and to other fish rhabdoviruses, except pike fry rhabdovirus (PFR). The possibility of a serological relationship between these two rhabdoviruses of the Vesiculovirus genus was confirmed by Jørgensen et al. (1989). Immunochemical and biological examinations of 22 rhabdovirus isolates from Cyprinidae, Esocidae and Siluridae show that SVCV and PFR share common antigenic determinants on the G, N and M proteins which prevent the two viruses being distinguished by an indirect fluorescent antibody test (IFAT). Spring viraemia of carp virus and PFR can be differentiated by an ELISA and by a virus neutralization test, if the rabbit antiserum is heat-inactivated and no complement is added. Jørgensen et al. (1989) suggested that SVCV and PFR are two serotypes of a single virus species. Heat-inactivated hyperimmune carp anti-SVCV serum can also distinguish SVCV from PFR (Petrinec, 1984).
The virulence of individual isolates may vary with the species or age of fish from which the virus was isolated (Ahne, 1986; Shchelkunov and Shchelkunova, 1989). It can be reduced by passage in mammalian and fish cell lines (Fijan et al., 1977b; Kölbl, 1980).
Host AnimalsTop of page
Vectors and Intermediate HostsTop of page
ReferencesTop of page
Ahne W, 1973. Zellkulturen aus verschiedenen Süsswasserteleosteergeweben und Untersuchung über die Ätiologie der Schwimmblasenentzündung der Karpfen. PhD dissertation. Munich: Ludwig-Maximilians Universität.
Ahne W, 1977. Evidence for the systemic character of Rhabdovirus carpio infection. (Summary). Bulletin de l'Office International des Epizooties, 87(5/6):435-436.
Ahne W, 1982. Survival of fish viruses. Fortschritte der Veterinarmedizin, No.35:305-309.
Ahne W, 1986. Different biological properties of four rhabdoviruses isolated from cyprinid fish. Journal of Veterinary Medicine, B (Infectious Diseases, Immunology, Food Hygiene, Veterinary Public Health), 33(4):253-259.
Arshaniza NM; Bauer ON, 1973. The epizootiology, diagnosis and prophylaxis of swim-bladder inflammation of cyprinids (SBI). In: Dill WA, ed. Symposium on the Major Communicable Fish Diseases in Europe and Their Control. European Inland Fisheries Advisory Commission. Technical Paper 17, Supplement 2, Rome, 140-144.
Bachmann PA; Ahne W, 1973. Isolation and characterization of agent causing swim bladder inflammation in carp. Nature, 244(No.5413):235-237.
Baudouy AM, 1975. Virémie printanière de la carpe: premiers isolements du virus en France. Bulletin de l’Office International des Epizooties, 83:717-722.
Békési L; Szabó E, 1979. A rhabdovirus isolated from carps in Hungary. Experimental infection of carps and resistance to the virus. Acta Microbiol Acad Sci Hung, 26:193-197.
Bishop DHL; Smith MS, 1977. Rhabdoviruses. In: Nayak DP, ed. The Molecular Biology of Animal Viruses. New York and Basle: Marcel Dekker, 167-280.
Björklund HV; Emmenegger EJ; Kurath G, 1995. Comparison of the polymerases (L genes) of spring viremia of carp virus and infectious hematopoietic necrosis virus. Veterinary Research, 26(5/6):394-398; 12 ref.
Björklund HV; Higman KH; Kurath G, 1996. The glycoprotein genes and gene junctions of the fish rhabdoviruses spring viremia of carp virus and hirame rhabdovirus: analysis of relationships with other rhabdoviruses. Virus Research, 42(1/2):65-80; 55 ref.
Björklund HV; Johansson TR; Rinne A, 1997. Rhabdovirus-induced apoptosis in a fish cell line is inhibited by a human endogenous acid cysteine proteinase inhibitor. Journal of Virology, 71:5658-5662.
Bowser P; Plumb JA, 1980. Fish cell lines: establishment of a line from ovaries of channel catfish. In Vitro, 16:365-368.
Bowser PR; Plumb JA, 1980. Growth rates of a new cell line from channel catfish ovary and channel catfish virus replication at different temperatures. Canadian Journal of Fisheries and Aquatic Sciences, 37(5):871-873.
Clark HF; Soriano EZ, 1974. Fish rhabdovirus replication in non-piscine cell culture: new system for the study of rhabdovirus-cell interaction in which the virus and cell have different temperature optima. Infection and Immunology, 10:180-188.
Deuter A; Enzmann P-J; Bigott K, 1982. Comparison of two fish pathogenic rhabdoviruses: L-(75Se) selenomethionine-labelled proteins. Zentralblatt fur Bakteriologie Mikrobiologie und Hygiene, I. Abt. Originale, A, 253(1):10-11; [abstract].
Dikkeboom AL; Radi C; Toohey-Kurth K; Marcquenski SV; Engel M; Goodwin AE; Way K; Stone DM; Longshaw CB, 2004. First report of spring viremia of carp virus (SVCV) in wild common carp in North America. Journal of Aquatic Animal Health, 16:169-178.
Faisal M; Ahne W, 1990. A cell line (CLC) of adherent peripheral blood mononuclear leucocytes of normal common carp Cyprinus carpio.. Developmental and Comparative Immunology, 14(2):255-260.
Fijan N; Petrinec Z; Stancl Z; Dorson M; Berre Mle, 1977. Hyperimmunization of carp with Rhabdovirus carpio. Bulletin de l'Office International des Epizooties, 87(5/6):439-440.
Fijan N; Petrinec Z; Stancl Z; Kezic N; Teskeredzic E, 1977. Vaccination of carp against spring viraemia: comparison of intraperitoneal and peroral application of live virus to fish kept in ponds. Bulletin de l'Office International des Epizooties, 87(5/6):441-442.
Fijan N; Sulimanovic D; Bearzotti M; Muzinic D; Zwillenberg LO; Chilmonczyk S; Vautherot JF; DeKinkelin P, 1983. Some properties of the epithelioma papulosum Cyprini (EPC) cell line. Annales de Virologie, Institut Pasteur, 134E:207-220.
Gravell M; Malsberger R, 1965. A permanent cell line from the fathead minnow (Pimephales promelas). Annals of the New York Academy of Sciences, 126:555-565.
Gupta KC; Bishop DHL; Roy P, 1979. 5-Terminal sequences of spring viremia of carp virus RNA synthesized in vitro. Journal of Virology, 30:735-745.
Gupta KC; Roy P, 1980. Alternate capping mechanisms for transcription of spring viremia of carp virus: evidence for independent mRNA initiation. Journal of Virology, 33(1):292-303.
Gupta KC; Roy P, 1981. Synthesis of capped and uncapped methylated oligonucleotides by the virion transcriptase of spring viremia of carp virus, a rhabdovirus. Proceedings of the National Academy of Sciences, 78(8):4758-4762.
Hill BJ, 1977. Studies on SVC [spring viraemia of carp] virulence and immunization. Bulletin de l'Office International des Epizooties, 87(5/6):455-456.
Hoffmann B; Schütze H; Mettenleiter TC, 2002. Determination of the complete genomic sequence and analysis of the gene products of the virus of Spring Viremia of Carp, a fish rhabdovirus. Virus Research, 84:89-100.
Jorgensen PEV; Olesen NJ; Ahne W; Lorenzen N, 1989. SVCV and PFR viruses: serological examination of 22 isolates indicates close relationship between the two fish rhabdoviruses. Viruses of lower vertebrates., 349-366; [1st International Symposium on Viruses of Lower Vertebrates, Munich, August 1988].
Kinkelin Pde; Le Berre M, 1974. Rhabdovirus des poissons. II. Proprietes in vitro du virus de la viremie printaniere de la carpe. Annales de Microbiologie (Paris), 125A:113-124.
Kiuchi A; Roy P, 1984. Comparison of the primary sequence of spring viremia of carp virus M protein with that of vesicular stomatitis virus. Virology, 134(1):238-243; 16 ref.
Kolbl O, 1980. Diagnosis of spring viraemia of carp and an immunization experiment against this disease. Bulletin de l'Office International des Epizooties, 92(9/10):1055-1068.
Kurath G; Highman KH; Bjorklund H, 1994. Comparative analysis of nonvirion (NV) genes and glycoprotein-polymerase gene junctions in the fish rhabdoviruses IHNV, HRV and SVCV. In: International Symposium on Aquatic Animal Health. Program and Abstracts. Davis, California, USA, W-4.4.
Lenoir G, 1973. Structural proteins of spring viremia virus of carp. Biochemical and Biophysical Research Communications, 51:895-898.
OIE Handistatus, 2005. World Animal Health Publication and Handistatus II (data set for 2004). Paris, France: Office International des Epizooties.
Osadcaja EF; Rudenko AP, 1981. Patogennost virusov, vydelennyh pri krasnuhe (vesennej viremii) karpov i kliniko-morfologicheskaja harakteristika estestvennogo techenija bolezni i v eksperimente. Rybnoe hozjajstvo, Kiev, 32:66-71.
Petersen JM; Her L-S; Dahlberg JE, 2002. Multiple vesiculoviral matrix proteins inhibit both nuclear export and import. Proceedings of the National Academy of Science, USA, 98:8590-8595.
Petrinec Z, 1984. Istrazivanje svojstava Rhabdovirus carpio. DSci dissertation. Zagreb: University of Zagreb.
Regenmortel MHVvan; Fauquet CM; Bishop DHL; Carstens EB; Estes MK; Lemon SM; Maniloff J; Mayo MA; McGeoch DJ; Pringle CR; Wickner RB, 2000. Virus taxonomy: classification and nomenclature of viruses. Seventh report of the International Committee on Taxonomy of Viruses. xii + 1162 pp.; many ref.
Roy P, 1981. Phosphoproteins of spring viremia of carp virus. Virology, 112(1):274-281.
Roy P; Clewley JP, 1978. Phosphoproteins of spring viremia carp virus and other rhabdoviruses. In: Mahy BWJ, Barry RD, eds. Negative Strand Viruses and Host Cells. New York: Academic Press, 116-125.
Roy P; Clewley JP, 1978. Spring viremia of carp virus RNA and virion-associated transcriptase activity. Journal of Virology, 25(3):912-916.
Roy P; Gupta KC; Kiuchi A, 1984. Characterization of spring viremia of carp virus mRNA species and the 3
Rudikov NI; Griscenko LI; Lobuncov KA, 1975. Vesennaja virusnaja boleznj ryb. Bjulletin Vsesojuznogo Ordeni Lenina Institut Eksperimentaljnoj Veterinarii, Moskva, 20:16-19.
Shchelkunov IS; Shchelkunova TI, 1989. Rhabdovirus carpio in herbivorous fishes: isolation, pathology and comparative susceptibility of fishes. Viruses of lower vertebrates., 333-348; [1st International Symposium on Viruses of Lower Vertebrates, Munich, August 1988].
Sokol F; Clark HF; Wiktor TJ; McFalls ML; Bishop DHL; Obijeski JF, 1974. Structural phosphoproteins associated with ten rhabdoviruses. Journal of General Virology, 24:433-445.
Tesarcík J; Macura B; Dedek L, 1977. Izolace virového agens z prípadu jarní virémie kapru v CSR. Buletin VURH Vodnany, 13:10-14.
OIE Handistatus, 2005. World Animal Health Publication and Handistatus II (dataset for 2004)., Paris, France: Office International des Epizooties.
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