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Journal of virology1992; 66(11); 6294-6303; doi: 10.1128/JVI.66.11.6294-6303.1992

Structural proteins of equine arteritis virus.

Abstract: We have recently shown that the genome of equine arteritis virus (EAV) contains seven open reading frames (ORFs). We now present data on the structural proteins of EAV and the assignment of their respective genes. Virions are composed of a 14-kDa nucleocapsid protein (N) and three membrane proteins designated M, GS, and GL. M is an unglycosylated protein of 16 kDa, and GS and GL are N-glycosylated proteins of 25 and 30 to 42 kDa, respectively. The broad size distribution of GL results from heterogeneous N-acetyllactosamine addition since it is susceptible to digestion by endo-beta-galactosidase. Using monospecific antisera as well as an antivirion serum, and by expression of individual ORFs, the genes for the structural proteins were identified: ORF 7 codes for N, ORF 6 for M, ORF 5 for GL, and ORF 2 for GS. With the exception of GS, the proteins are about equally abundant in EAV virions, being present at a molar ratio of 3 (N):2 (M):3 (GL). The GS protein, which is expressed at a level similar to that of M in infected cells, is strikingly underrepresented in virus particles (1 to 2%). Our data justify a distinct taxonomic position for EAV, together with lactate dehydrogenase-elevating virus and simian hemorrhagic fever virus; although coronavirus- and toroviruslike in features of transcription and translation, the virion architecture of EAV is fundamentally different.
Publication Date: 1992-11-01 PubMed ID: 1328669PubMed Central: PMC240121DOI: 10.1128/JVI.66.11.6294-6303.1992Google Scholar: Lookup
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Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

This research article delves into the structural proteins found in the equine arteritis virus (EAV), assigning each of them to their respective genes. The study establishes that EAV possesses a unique structural complexity among viruses, and thus merits its own distinct taxonomic categorisation.

Investigating the Structure of Equine Arteritis Virus

  • The research displays the composition of the equine arteritis virus, which is a virus affecting the horses’ respiratory system. The scientists used specific tools and processes to identify the structural proteins of the virus and link them to their respective genes.
  • The virus was found to be composed of one 14-kDa nucleocapsid protein (N) and three membrane proteins designated M, GS, and GL. These proteins were identified by using monospecific antisera, an antivirion serum, as well as by expressing individual open reading frames (ORFs), which are sections of the virus’s genome that can be decoded into proteins.

Discoveries about the Structural Proteins

  • The membrane protein M is unglycosylated and weighs 16 kDa. The other two proteins, GS and GL, are N-glycosylated and weigh 25 and between 30 to 42 kDa, respectively. The wide size range of GL is due to the heterogeneous addition of N-acetyllactosamine, which makes it susceptible to digestion by endo-beta-galactosidase.
  • The structural proteins were identified to specific genes, revealing that ORF 7 codes for N, ORF 6 for M, ORF 5 for GL, and ORF 2 for GS. These proteins have roughly the same presence in EAV virions, being present at a molar ratio of 3(N):2(M):3(GL), except for GS which is significantly less prevalent in virus particles.

Taxonomic Position of EAV

  • Although EAV shares characteristics with coronavirus and torovirus in terms of transcription and translation features, its virion structure is notably distinctive. This leads to the conclusion that EAV deserves its own unique classification, along with the lactate dehydrogenase-elevating virus and the simian hemorrhagic fever virus.
  • This research has therefore significantly contributed to understanding the structural characteristics of EAV, enriching the scientific community’s knowledge of this important equine virus and its place within virus taxonomy.

Cite This Article

APA
de Vries AA, Chirnside ED, Horzinek MC, Rottier PJ. (1992). Structural proteins of equine arteritis virus. J Virol, 66(11), 6294-6303. https://doi.org/10.1128/JVI.66.11.6294-6303.1992

Publication

ISSN: 0022-538X
NlmUniqueID: 0113724
Country: United States
Language: English
Volume: 66
Issue: 11
Pages: 6294-6303

Researcher Affiliations

de Vries, A A
  • Department of Infectious Diseases and Immunology, Veterinary Faculty, University of Utrecht, The Netherlands.
Chirnside, E D
    Horzinek, M C
      Rottier, P J

        MeSH Terms

        • Amino Acid Sequence
        • Antibodies, Viral
        • Capsid / analysis
        • Capsid / genetics
        • Chromosome Mapping
        • Cloning, Molecular
        • Equartevirus / chemistry
        • Equartevirus / genetics
        • Genes, Viral / genetics
        • Genome, Viral
        • Glycoproteins / analysis
        • Glycoproteins / genetics
        • Molecular Sequence Data
        • Open Reading Frames / genetics
        • Protein Conformation
        • Sequence Analysis, DNA
        • Viral Matrix Proteins / analysis
        • Viral Matrix Proteins / genetics
        • Viral Structural Proteins / chemistry
        • Viral Structural Proteins / genetics
        • Virion / genetics

        References

        This article includes 45 references
        1. Intervirology. 1978;9(3):129-48
          pubmed: 618831
        2. Cornell Vet. 1957 Jan;47(1):3-41
          pubmed: 13397177
        3. Proc Soc Exp Biol Med. 1975 Dec;150(3):707-11
          pubmed: 1208594
        4. J Cell Biol. 1992 May;117(4):895-902
          pubmed: 1374413
        5. Virus Res. 1992 Apr;23(1-2):55-72
          pubmed: 1604932
        6. J Virol. 1991 Sep;65(9):5118-23
          pubmed: 1870216
        7. Aust Vet J. 1990 Dec;67(12):429-31
          pubmed: 1963771
        8. Virology. 1991 Jun;182(2):655-63
          pubmed: 2024492
        9. Nucleic Acids Res. 1990 Jun 11;18(11):3241-7
          pubmed: 2162519
        10. J Biol Chem. 1990 Nov 25;265(33):20476-87
          pubmed: 2243101
        11. EMBO J. 1985 Nov;4(11):2869-77
          pubmed: 2415353
        12. J Biol Chem. 1988 Apr 15;263(11):5314-8
          pubmed: 2451668
        13. J Virol. 1988 Sep;62(9):3210-6
          pubmed: 2457113
        14. Proc Natl Acad Sci U S A. 1987 Nov;84(21):7413-7
          pubmed: 2823261
        15. Equine Vet J. 1985 Jan;17(1):6-11
          pubmed: 2983980
        16. Biochemistry. 1986 Mar 25;25(6):1335-9
          pubmed: 3008826
        17. Virus Res. 1986 Feb;4(2):145-56
          pubmed: 3010596
        18. Mol Cell Biol. 1988 Mar;8(3):1186-96
          pubmed: 3367907
        19. Annu Rev Biochem. 1985;54:631-64
          pubmed: 3896128
        20. Arch Gesamte Virusforsch. 1971;33(3):306-18
          pubmed: 4107033
        21. Arch Gesamte Virusforsch. 1970;30(2):105-12
          pubmed: 4195609
        22. Virology. 1973 Sep;55(1):211-7
          pubmed: 4738049
        23. Nature. 1970 Aug 15;227(5259):680-5
          pubmed: 5432063
        24. Zentralbl Veterinarmed B. 1983 May;30(4):297-304
          pubmed: 6191473
        25. Proc Natl Acad Sci U S A. 1984 Mar;81(5):1421-5
          pubmed: 6324191
        26. J Mol Biol. 1982 May 5;157(1):105-32
          pubmed: 7108955
        27. Virology. 1976 Aug;73(1):200-5
          pubmed: 183352
        28. Adv Virus Res. 1992;41:99-192
          pubmed: 1315480
        29. J Biol Chem. 1992 Mar 5;267(7):4440-55
          pubmed: 1537830
        30. J Virol. 1991 Jun;65(6):2910-20
          pubmed: 1851863
        31. Aust Vet J. 1990 Dec;67(12):432-5
          pubmed: 1963772
        32. Nucleic Acids Res. 1991 Apr 25;19 Suppl:2241-5
          pubmed: 2041810
        33. Virology. 1990 Aug;177(2):768-71
          pubmed: 2371780
        34. J Gen Virol. 1986 Aug;67 ( Pt 8):1543-9
          pubmed: 2426393
        35. J Virol. 1988 Aug;62(8):2762-72
          pubmed: 2455818
        36. J Biol Chem. 1988 Sep 5;263(25):12461-71
          pubmed: 2970459
        37. Intervirology. 1985;24(3):125-39
          pubmed: 2999027
        38. Proc Natl Acad Sci U S A. 1986 Nov;83(21):8122-6
          pubmed: 3095828
        39. Nucleic Acids Res. 1986 Jun 11;14(11):4683-90
          pubmed: 3714490
        40. Mol Cell Biol. 1985 Dec;5(12):3403-9
          pubmed: 3939316
        41. Arch Gesamte Virusforsch. 1973;40(3):177-88
          pubmed: 4633581
        42. Eur J Biochem. 1974 Jul 1;46(1):83-8
          pubmed: 4850204
        43. Virology. 1982 Apr 30;118(2):345-52
          pubmed: 6283728
        44. Biochem J. 1983 Aug 1;213(2):485-94
          pubmed: 6412685
        45. J Virol. 1975 Aug;16(2):420-33
          pubmed: 1171266

        Citations

        This article has been cited 97 times.
        1. Zhang M, Qian B, Kunec D, Veit M. Development of GFP-expressing infectious clones for PRRSV using TAR cloning for antiviral drug screening. Npj Viruses 2025 Sep 5;3(1):66.
          doi: 10.1038/s44298-025-00148-3pubmed: 40913022google scholar: lookup
        2. Treffers EE, Tas A, Scholte FEM, de Ru AH, Snijder EJ, van Veelen PA, van Hemert MJ. The alphavirus nonstructural protein 2 NTPase induces a host translational shut-off through phosphorylation of eEF2 via cAMP-PKA-eEF2K signaling. PLoS Pathog 2023 Feb;19(2):e1011179.
          doi: 10.1371/journal.ppat.1011179pubmed: 36848386google scholar: lookup
        3. Veit M, Gadalla MR, Zhang M. Using Alphafold2 to Predict the Structure of the Gp5/M Dimer of Porcine Respiratory and Reproductive Syndrome Virus. Int J Mol Sci 2022 Oct 30;23(21).
          doi: 10.3390/ijms232113209pubmed: 36361998google scholar: lookup
        4. Cai Y, Yu S, Fang Y, Bollinger L, Li Y, Lauck M, Postnikova EN, Mazur S, Johnson RF, Finch CL, Radoshitzky SR, Palacios G, Friedrich TC, Goldberg TL, O'Connor DH, Jahrling PB, Kuhn JH. Development and Characterization of a cDNA-Launch Recombinant Simian Hemorrhagic Fever Virus Expressing Enhanced Green Fluorescent Protein: ORF 2b' Is Not Required for In Vitro Virus Replication. Viruses 2021 Apr 7;13(4).
          doi: 10.3390/v13040632pubmed: 33917085google scholar: lookup
        5. Zhang M, Han X, Osterrieder K, Veit M. Palmitoylation of the envelope membrane proteins GP5 and M of porcine reproductive and respiratory syndrome virus is essential for virus growth. PLoS Pathog 2021 Apr;17(4):e1009554.
          doi: 10.1371/journal.ppat.1009554pubmed: 33891658google scholar: lookup
        6. Guo R, Yan X, Li Y, Cui J, Misra S, Firth AE, Snijder EJ, Fang Y. A swine arterivirus deubiquitinase stabilizes two major envelope proteins and promotes production of viral progeny. PLoS Pathog 2021 Mar;17(3):e1009403.
          doi: 10.1371/journal.ppat.1009403pubmed: 33735221google scholar: lookup
        7. Ding G, Liu J, Shao Q, Wang B, Feng J, Li Y, Li L, Cao S, Cong F, Zhao Y, Liu S, Xiao Y. Porcine Reproductive and Respiratory Syndrome Virus Structural Protein GP3 Regulates Claudin 4 To Facilitate the Early Stages of Infection. J Virol 2020 Sep 29;94(20).
          doi: 10.1128/JVI.00124-20pubmed: 32759320google scholar: lookup
        8. de Vries AAF, Horzinek MC, Rottier PJM, de Groot RJ. The Genome Organization of the Nidovirales: Similarities and Differences between Arteri-, Toro-, and Coronaviruses. Semin Virol 1997 Feb;8(1):33-47.
          doi: 10.1006/smvy.1997.0104pubmed: 32288441google scholar: lookup
        9. de Wilde AH, Boomaars-van der Zanden AL, de Jong AWM, Bárcena M, Snijder EJ, Posthuma CC. Adaptive Mutations in Replicase Transmembrane Subunits Can Counteract Inhibition of Equine Arteritis Virus RNA Synthesis by Cyclophilin Inhibitors. J Virol 2019 Sep 15;93(18).
          doi: 10.1128/JVI.00490-19pubmed: 31243130google scholar: lookup
        10. Nam B, Mekuria Z, Carossino M, Li G, Zheng Y, Zhang J, Cook RF, Shuck KM, Campos JR, Squires EL, Troedsson MHT, Timoney PJ, Balasuriya UBR. Intrahost Selection Pressure Drives Equine Arteritis Virus Evolution during Persistent Infection in the Stallion Reproductive Tract. J Virol 2019 Jun 15;93(12).
          doi: 10.1128/JVI.00045-19pubmed: 30918077google scholar: lookup
        11. Veit M, Matczuk AK, Sinhadri BC, Krause E, Thaa B. Membrane proteins of arterivirus particles: structure, topology, processing and function. Virus Res 2014 Dec 19;194:16-36.
        12. Butler JE, Lager KM, Golde W, Faaberg KS, Sinkora M, Loving C, Zhang YI. Porcine reproductive and respiratory syndrome (PRRS): an immune dysregulatory pandemic. Immunol Res 2014 Aug;59(1-3):81-108.
          doi: 10.1007/s12026-014-8549-5pubmed: 24981123google scholar: lookup
        13. Go YY, Li Y, Chen Z, Han M, Yoo D, Fang Y, Balasuriya UB. Equine arteritis virus does not induce interferon production in equine endothelial cells: identification of nonstructural protein 1 as a main interferon antagonist. Biomed Res Int 2014;2014:420658.
          doi: 10.1155/2014/420658pubmed: 24967365google scholar: lookup
        14. Balasuriya UB, Go YY, MacLachlan NJ. Equine arteritis virus. Vet Microbiol 2013 Nov 29;167(1-2):93-122.
          doi: 10.1016/j.vetmic.2013.06.015pubmed: 23891306google scholar: lookup
        15. Zhao S, Qi T, Guo W, Lu G, Xiang W. Identification of a conserved B-cell epitope in the equine arteritis virus (EAV) N protein using the pepscan technique. Virus Genes 2013 Oct;47(2):292-7.
          doi: 10.1007/s11262-013-0943-xpubmed: 23813249google scholar: lookup
        16. de Wilde AH, Li Y, van der Meer Y, Vuagniaux G, Lysek R, Fang Y, Snijder EJ, van Hemert MJ. Cyclophilin inhibitors block arterivirus replication by interfering with viral RNA synthesis. J Virol 2013 Feb;87(3):1454-64.
          doi: 10.1128/JVI.02078-12pubmed: 23152531google scholar: lookup
        17. Knoops K, Bárcena M, Limpens RW, Koster AJ, Mommaas AM, Snijder EJ. Ultrastructural characterization of arterivirus replication structures: reshaping the endoplasmic reticulum to accommodate viral RNA synthesis. J Virol 2012 Mar;86(5):2474-87.
          doi: 10.1128/JVI.06677-11pubmed: 22190716google scholar: lookup
        18. van Kasteren PB, Beugeling C, Ninaber DK, Frias-Staheli N, van Boheemen S, García-Sastre A, Snijder EJ, Kikkert M. Arterivirus and nairovirus ovarian tumor domain-containing Deubiquitinases target activated RIG-I to control innate immune signaling. J Virol 2012 Jan;86(2):773-85.
          doi: 10.1128/JVI.06277-11pubmed: 22072774google scholar: lookup
        19. Firth AE, Zevenhoven-Dobbe JC, Wills NM, Go YY, Balasuriya UBR, Atkins JF, Snijder EJ, Posthuma CC. Discovery of a small arterivirus gene that overlaps the GP5 coding sequence and is important for virus production. J Gen Virol 2011 May;92(Pt 5):1097-1106.
          doi: 10.1099/vir.0.029264-0pubmed: 21307223google scholar: lookup
        20. Silva PA, Pereira CF, Dalebout TJ, Spaan WJ, Bredenbeek PJ. An RNA pseudoknot is required for production of yellow fever virus subgenomic RNA by the host nuclease XRN1. J Virol 2010 Nov;84(21):11395-406.
          doi: 10.1128/JVI.01047-10pubmed: 20739539google scholar: lookup
        21. Nedialkova DD, Gorbalenya AE, Snijder EJ. Arterivirus Nsp1 modulates the accumulation of minus-strand templates to control the relative abundance of viral mRNAs. PLoS Pathog 2010 Feb 19;6(2):e1000772.
          doi: 10.1371/journal.ppat.1000772pubmed: 20174607google scholar: lookup
        22. Zevenhoven-Dobbe JC, Wassenaar AL, van der Meer Y, Snijder EJ. Production of monospecific rabbit antisera recognizing nidovirus proteins. Methods Mol Biol 2008;454:205-26.
          doi: 10.1007/978-1-59745-181-9_16pubmed: 19057875google scholar: lookup
        23. Nitschke M, Korte T, Tielesch C, Ter-Avetisyan G, Tünnemann G, Cardoso MC, Veit M, Herrmann A. Equine arteritis virus is delivered to an acidic compartment of host cells via clathrin-dependent endocytosis. Virology 2008 Aug 1;377(2):248-54.
          doi: 10.1016/j.virol.2008.04.041pubmed: 18570963google scholar: lookup
        24. Veit M, Kabatek A, Tielesch C, Hermann A. Characterization of equine arteritis virus particles and demonstration of their hemolytic activity. Arch Virol 2008;153(2):351-6.
          doi: 10.1007/s00705-007-1094-ypubmed: 18219439google scholar: lookup
        25. Go YY, Wong SJ, Branscum AJ, Demarest VL, Shuck KM, Vickers ML, Zhang J, McCollum WH, Timoney PJ, Balasuriya UB. Development of a fluorescent-microsphere immunoassay for detection of antibodies specific to equine arteritis virus and comparison with the virus neutralization test. Clin Vaccine Immunol 2008 Jan;15(1):76-87.
          doi: 10.1128/CVI.00388-07pubmed: 18032597google scholar: lookup
        26. Li G, Huang J, Jiang P, Li Y, Jiang W, Wang X. Suppression of porcine reproductive and respiratory syndrome virus replication in MARC-145 cells by shRNA targeting ORF1 region. Virus Genes 2007 Dec;35(3):673-9.
          doi: 10.1007/s11262-007-0134-8pubmed: 17671836google scholar: lookup
        27. Echeverría MG, Díaz S, Metz GE, Serena MS, Panei CJ, Nosetto E. Genetic typing of equine arteritis virus isolates from Argentina. Virus Genes 2007 Oct;35(2):313-20.
          doi: 10.1007/s11262-007-0081-4pubmed: 17294142google scholar: lookup
        28. Schommer SK, Kleiboeker SB. Use of a PRRSV infectious clone to evaluate in vitro quasispecies evolution. Adv Exp Med Biol 2006;581:435-8.
          doi: 10.1007/978-0-387-33012-9_78pubmed: 17037574google scholar: lookup
        29. Glaser AL, Chirnside ED, Horzinek MC, de Vries AA. Equine arteritis virus. Theriogenology 1997 Apr 15;47(6):1275-95.
          doi: 10.1016/s0093-691x(97)00107-6pubmed: 16728076google scholar: lookup
        30. van Aken D, Snijder EJ, Gorbalenya AE. Mutagenesis analysis of the nsp4 main proteinase reveals determinants of arterivirus replicase polyprotein autoprocessing. J Virol 2006 Apr;80(7):3428-37.
        31. Posthuma CC, Nedialkova DD, Zevenhoven-Dobbe JC, Blokhuis JH, Gorbalenya AE, Snijder EJ. Site-directed mutagenesis of the Nidovirus replicative endoribonuclease NendoU exerts pleiotropic effects on the arterivirus life cycle. J Virol 2006 Feb;80(4):1653-61.
        32. van den Born E, Posthuma CC, Gultyaev AP, Snijder EJ. Discontinuous subgenomic RNA synthesis in arteriviruses is guided by an RNA hairpin structure located in the genomic leader region. J Virol 2005 May;79(10):6312-24.
        33. Wieringa R, de Vries AA, van der Meulen J, Godeke GJ, Onderwater JJ, van Tol H, Koerten HK, Mommaas AM, Snijder EJ, Rottier PJ. Structural protein requirements in equine arteritis virus assembly. J Virol 2004 Dec;78(23):13019-27.
        34. Takahashi-Omoe H, Omoe K, Sakaguchi M, Kameoka Y, Matsushita S, Inada T. Analysis of protein expression by mammalian cell lines stably expressing lactate dehydrogenase-elevating virus ORF 5 and ORF 6 proteins. Comp Immunol Microbiol Infect Dis 2004 Mar;27(2):81-92.
          doi: 10.1016/S0147-9571(03)00053-5pubmed: 14690718google scholar: lookup
        35. Takahashi-Omoe H, Omoe K, Sakaguchi M, Kameoka Y, Matsushita S, Inada T. Production of virus-specific antiserum corresponding to sequences in the lactate dehydrogenase-elevating virus (LDV) ORF6 protein. Comp Immunol Microbiol Infect Dis 2004 Jan;27(1):47-55.
          doi: 10.1016/S0147-9571(03)00035-3pubmed: 14656541google scholar: lookup
        36. Wieringa R, De Vries AA, Post SM, Rottier PJ. Intra- and intermolecular disulfide bonds of the GP2b glycoprotein of equine arteritis virus: relevance for virus assembly and infectivity. J Virol 2003 Dec;77(24):12996-3004.
        37. Castillo-Olivares J, Wieringa R, Bakonyi T, de Vries AA, Davis-Poynter NJ, Rottier PJ. Generation of a candidate live marker vaccine for equine arteritis virus by deletion of the major virus neutralization domain. J Virol 2003 Aug;77(15):8470-80.
        38. Wieringa R, de Vries AA, Rottier PJ. Formation of disulfide-linked complexes between the three minor envelope glycoproteins (GP2b, GP3, and GP4) of equine arteritis virus. J Virol 2003 Jun;77(11):6216-26.
        39. Snijder EJ, Dobbe JC, Spaan WJ. Heterodimerization of the two major envelope proteins is essential for arterivirus infectivity. J Virol 2003 Jan;77(1):97-104.
          doi: 10.1128/jvi.77.1.97-104.2003pubmed: 12477814google scholar: lookup
        40. Wieringa R, de Vries AA, Raamsman MJ, Rottier PJ. Characterization of two new structural glycoproteins, GP(3) and GP(4), of equine arteritis virus. J Virol 2002 Nov;76(21):10829-40.
        41. Jeronimo C, Archambault D. Importance of M-protein C terminus as substrate antigen for serodetection of equine arteritis virus infection. Clin Diagn Lab Immunol 2002 May;9(3):698-703.
          doi: 10.1128/cdli.9.3.698-703.2002pubmed: 11986280google scholar: lookup
        42. Plagemann PG. An ORF-2a protein is not present at a significant level in virions of the arterivirus lactate dehydrogenase-elevating virus. Virus Res 2001 Apr;74(1-2):47-52.
          doi: 10.1016/s0168-1702(00)00247-1pubmed: 11226573google scholar: lookup
        43. Tijms MA, van Dinten LC, Gorbalenya AE, Snijder EJ. A zinc finger-containing papain-like protease couples subgenomic mRNA synthesis to genome translation in a positive-stranded RNA virus. Proc Natl Acad Sci U S A 2001 Feb 13;98(4):1889-94.
          doi: 10.1073/pnas.98.4.1889pubmed: 11172046google scholar: lookup
        44. Pasternak AO, Gultyaev AP, Spaan WJ, Snijder EJ. Genetic manipulation of arterivirus alternative mRNA leader-body junction sites reveals tight regulation of structural protein expression. J Virol 2000 Dec;74(24):11642-53.
        45. Balasuriya UB, Heidner HW, Hedges JF, Williams JC, Davis NL, Johnston RE, MacLachlan NJ. Expression of the two major envelope proteins of equine arteritis virus as a heterodimer is necessary for induction of neutralizing antibodies in mice immunized with recombinant Venezuelan equine encephalitis virus replicon particles. J Virol 2000 Nov;74(22):10623-30.
        46. Molenkamp R, Greve S, Spaan WJ, Snijder EJ. Efficient homologous RNA recombination and requirement for an open reading frame during replication of equine arteritis virus defective interfering RNAs. J Virol 2000 Oct;74(19):9062-70.
        47. Weiland E, Bolz S, Weiland F, Herbst W, Raamsman MJ, Rottier PJ, De Vries AA. Monoclonal antibodies directed against conserved epitopes on the nucleocapsid protein and the major envelope glycoprotein of equine arteritis virus. J Clin Microbiol 2000 Jun;38(6):2065-75.
        48. Meng XJ. Heterogeneity of porcine reproductive and respiratory syndrome virus: implications for current vaccine efficacy and future vaccine development. Vet Microbiol 2000 Jun 12;74(4):309-29.
          doi: 10.1016/s0378-1135(00)00196-6pubmed: 10831854google scholar: lookup
        49. van Dinten LC, van Tol H, Gorbalenya AE, Snijder EJ. The predicted metal-binding region of the arterivirus helicase protein is involved in subgenomic mRNA synthesis, genome replication, and virion biogenesis. J Virol 2000 Jun;74(11):5213-23.
        50. de Haan CA, Vennema H, Rottier PJ. Assembly of the coronavirus envelope: homotypic interactions between the M proteins. J Virol 2000 Jun;74(11):4967-78.
        51. Molenkamp R, Rozier BC, Greve S, Spaan WJ, Snijder EJ. Isolation and characterization of an arterivirus defective interfering RNA genome. J Virol 2000 Apr;74(7):3156-65.
        52. Cho HJ, Entz SC, Deregt D, Jordan LT, Timoney PJ, McCollum WH. Detection of antibodies to equine arteritis virus by a monoclonal antibody-based blocking ELISA. Can J Vet Res 2000 Jan;64(1):38-43.
          pubmed: 10680655
        53. Raamsman MJ, Locker JK, de Hooge A, de Vries AA, Griffiths G, Vennema H, Rottier PJ. Characterization of the coronavirus mouse hepatitis virus strain A59 small membrane protein E. J Virol 2000 Mar;74(5):2333-42.
        54. de Haan CA, Smeets M, Vernooij F, Vennema H, Rottier PJ. Mapping of the coronavirus membrane protein domains involved in interaction with the spike protein. J Virol 1999 Sep;73(9):7441-52.
        55. Snijder EJ, van Tol H, Pedersen KW, Raamsman MJ, de Vries AA. Identification of a novel structural protein of arteriviruses. J Virol 1999 Aug;73(8):6335-45.
        56. van Marle G, van Dinten LC, Spaan WJ, Luytjes W, Snijder EJ. Characterization of an equine arteritis virus replicase mutant defective in subgenomic mRNA synthesis. J Virol 1999 Jul;73(7):5274-81.
        57. Hedges JF, Balasuriya UB, Timoney PJ, McCollum WH, MacLachlan NJ. Genetic divergence with emergence of novel phenotypic variants of equine arteritis virus during persistent infection of stallions. J Virol 1999 May;73(5):3672-81.
        58. van Dinten LC, Rensen S, Gorbalenya AE, Snijder EJ. Proteolytic processing of the open reading frame 1b-encoded part of arterivirus replicase is mediated by nsp4 serine protease and Is essential for virus replication. J Virol 1999 Mar;73(3):2027-37.
        59. Pedersen KW, van der Meer Y, Roos N, Snijder EJ. Open reading frame 1a-encoded subunits of the arterivirus replicase induce endoplasmic reticulum-derived double-membrane vesicles which carry the viral replication complex. J Virol 1999 Mar;73(3):2016-26.
        60. Kheyar A, St-Laurent G, Diouri M, Archambault D. Nucleotide sequence and genetic analysis of the leader region of Canadian, American and European equine arteritis virus isolates. Can J Vet Res 1998 Jul;62(3):224-30.
          pubmed: 9684053
        61. van der Meer Y, van Tol H, Locker JK, Snijder EJ. ORF1a-encoded replicase subunits are involved in the membrane association of the arterivirus replication complex. J Virol 1998 Aug;72(8):6689-98.
        62. Mardassi H, Gonin P, Gagnon CA, Massie B, Dea S. A subset of porcine reproductive and respiratory syndrome virus GP3 glycoprotein is released into the culture medium of cells as a non-virion-associated and membrane-free (soluble) form. J Virol 1998 Aug;72(8):6298-306.
        63. Kheyar A, Martin S, St-Laurent G, Timoney PJ, McCollum WH, Archambault D. Expression cloning and humoral immune response to the nucleocapsid and membrane proteins of equine arteritis virus. Clin Diagn Lab Immunol 1997 Nov;4(6):648-52.
          doi: 10.1128/cdli.4.6.648-652.1997pubmed: 9384283google scholar: lookup
        64. Wassenaar AL, Spaan WJ, Gorbalenya AE, Snijder EJ. Alternative proteolytic processing of the arterivirus replicase ORF1a polyprotein: evidence that NSP2 acts as a cofactor for the NSP4 serine protease. J Virol 1997 Dec;71(12):9313-22.
        65. Gilbert SA, Timoney PJ, McCollum WH, Deregt D. Detection of equine arteritis virus in the semen of carrier stallions by using a sensitive nested PCR assay. J Clin Microbiol 1997 Aug;35(8):2181-3.
        66. Meulenberg JJ, van Nieuwstadt AP, van Essen-Zandbergen A, Langeveld JP. Posttranslational processing and identification of a neutralization domain of the GP4 protein encoded by ORF4 of Lelystad virus. J Virol 1997 Aug;71(8):6061-7.
        67. Meulenberg JJ, Petersen den Besten A, de Kluyver E, van Nieuwstadt A, Wensvoort G, Moormann RJ. Molecular characterization of Lelystad virus. Vet Microbiol 1997 Apr;55(1-4):197-202.
          doi: 10.1016/s0378-1135(96)01335-1pubmed: 9220614google scholar: lookup
        68. Smith SL, Wang X, Godeny EK. Sequence of the 3' end of the simian hemorrhagic fever virus genome. Gene 1997 Jun 3;191(2):205-10.
          doi: 10.1016/s0378-1119(97)00061-9pubmed: 9218721google scholar: lookup
        69. Plana Duran J, Climent I, Sarraseca J, Urniza A, Cortés E, Vela C, Casal JI. Baculovirus expression of proteins of porcine reproductive and respiratory syndrome virus strain Olot/91. Involvement of ORF3 and ORF5 proteins in protection. Virus Genes 1997;14(1):19-29.
          doi: 10.1023/a:1007931322271pubmed: 9208452google scholar: lookup
        70. van Dinten LC, den Boon JA, Wassenaar AL, Spaan WJ, Snijder EJ. An infectious arterivirus cDNA clone: identification of a replicase point mutation that abolishes discontinuous mRNA transcription. Proc Natl Acad Sci U S A 1997 Feb 4;94(3):991-6.
          doi: 10.1073/pnas.94.3.991pubmed: 9023370google scholar: lookup
        71. St-Laurent G, Lepage N, Carman S, Archambault D. Genetic and amino acid analysis of the GL protein of Canadian, American and European equine arteritis virus isolates. Can J Vet Res 1997 Jan;61(1):72-6.
          pubmed: 9008807
        72. Lepage N, St-Laurent G, Carman S, Archambault D. Comparison of nucleic and amino acid sequences and phylogenetic analysis of the Gs protein of various equine arteritis virus isolates. Virus Genes 1996;13(1):87-91.
          doi: 10.1007/BF00576983pubmed: 8938984google scholar: lookup
        73. Chirnside ED, Francis PM, Mumford JA. Expression cloning and antigenic analysis of the nucleocapsid protein of equine arteritis virus. Virus Res 1995 Dec;39(2-3):277-88.
          doi: 10.1016/0168-1702(95)00098-4pubmed: 8837890google scholar: lookup
        74. Hedges JF, Balasuriya UB, Timoney PJ, McCollum WH, MacLachlan NJ. Genetic variation in open reading frame 2 of field isolates and laboratory strains of equine arteritis virus. Virus Res 1996 Jun;42(1-2):41-52.
          doi: 10.1016/0168-1702(96)01294-4pubmed: 8806173google scholar: lookup
        75. van Dinten LC, Wassenaar AL, Gorbalenya AE, Spaan WJ, Snijder EJ. Processing of the equine arteritis virus replicase ORF1b protein: identification of cleavage products containing the putative viral polymerase and helicase domains. J Virol 1996 Oct;70(10):6625-33.
        76. Bautista EM, Meulenberg JJ, Choi CS, Molitor TW. Structural polypeptides of the American (VR-2332) strain of porcine reproductive and respiratory syndrome virus. Arch Virol 1996;141(7):1357-65.
          doi: 10.1007/BF01718837pubmed: 8774694google scholar: lookup
        77. van Nieuwstadt AP, Meulenberg JJ, van Essen-Zanbergen A, Petersen-den Besten A, Bende RJ, Moormann RJ, Wensvoort G. Proteins encoded by open reading frames 3 and 4 of the genome of Lelystad virus (Arteriviridae) are structural proteins of the virion. J Virol 1996 Jul;70(7):4767-72.
        78. den Boon JA, Kleijnen MF, Spaan WJ, Snijder EJ. Equine arteritis virus subgenomic mRNA synthesis: analysis of leader-body junctions and replicative-form RNAs. J Virol 1996 Jul;70(7):4291-8.
        79. Loemba HD, Mounir S, Mardassi H, Archambault D, Dea S. Kinetics of humoral immune response to the major structural proteins of the porcine reproductive and respiratory syndrome virus. Arch Virol 1996;141(3-4):751-61.
          doi: 10.1007/BF01718333pubmed: 8645111google scholar: lookup
        80. Plagemann PG, Rowland RR, Even C, Faaberg KS. Lactate dehydrogenase-elevating virus: an ideal persistent virus?. Springer Semin Immunopathol 1995;17(2-3):167-86.
          doi: 10.1007/BF00196164pubmed: 8571167google scholar: lookup
        81. Cavanagh D, Brien DA, Brinton M, Enjuanes L, Holmes KV, Horzinek MC, Lai MMC, Laude H, Plagemann PGW, Siddell S, Spaan WJM, Taguchi F, Talbot PJ. Revision of the taxonomy of the Coronavirus, Torovirus and Arterivirus genera. Arch Virol 1994;135(1-2):227-37.
          doi: 10.1007/BF01309782pubmed: 8198447google scholar: lookup
        82. St-Laurent G, Morin G, Archambault D. Detection of equine arteritis virus following amplification of structural and nonstructural viral genes by reverse transcription-PCR. J Clin Microbiol 1994 Mar;32(3):658-65.
          doi: 10.1128/jcm.32.3.658-665.1994pubmed: 8195375google scholar: lookup
        83. Mardassi H, Athanassious R, Mounir S, Dea S. Porcine reproductive and respiratory syndrome virus: morphological, biochemical and serological characteristics of Quebec isolates associated with acute and chronic outbreaks of porcine reproductive and respiratory syndrome. Can J Vet Res 1994 Jan;58(1):55-64.
          pubmed: 8143254
        84. Horzinek MC. Snowdon Lecture 1993. New virus diseases: visible evolution. Aust Vet J 1993 Dec;70(12):433-6.
        85. Snijder EJ, Wassenaar AL, Spaan WJ. Proteolytic processing of the replicase ORF1a protein of equine arteritis virus. J Virol 1994 Sep;68(9):5755-64.
        86. Faaberg KS, Even C, Palmer GA, Plagemann PG. Disulfide bonds between two envelope proteins of lactate dehydrogenase-elevating virus are essential for viral infectivity. J Virol 1995 Jan;69(1):613-7.
          doi: 10.1128/JVI.69.1.613-617.1995pubmed: 7983766google scholar: lookup
        87. Godeny EK, Zeng L, Smith SL, Brinton MA. Molecular characterization of the 3' terminus of the simian hemorrhagic fever virus genome. J Virol 1995 Apr;69(4):2679-83.
        88. Chen Z, Faaberg KS, Plagemann PG. Detection of negative-stranded subgenomic RNAs but not of free leader in LDV-infected macrophages. Virus Res 1994 Nov;34(2):167-77.
          doi: 10.1016/0168-1702(94)90098-1pubmed: 7856308google scholar: lookup
        89. Meulenberg JJ, Petersen-den Besten A, De Kluyver EP, Moormann RJ, Schaaper WM, Wensvoort G. Characterization of proteins encoded by ORFs 2 to 7 of Lelystad virus. Virology 1995 Jan 10;206(1):155-63.
          doi: 10.1016/s0042-6822(95)80030-1pubmed: 7831770google scholar: lookup
        90. de Vries AA, Raamsman MJ, van Dijk HA, Horzinek MC, Rottier PJ. The small envelope glycoprotein (GS) of equine arteritis virus folds into three distinct monomers and a disulfide-linked dimer. J Virol 1995 Jun;69(6):3441-8.
        91. Murtaugh MP, Elam MR, Kakach LT. Comparison of the structural protein coding sequences of the VR-2332 and Lelystad virus strains of the PRRS virus. Arch Virol 1995;140(8):1451-60.
          doi: 10.1007/BF01322671pubmed: 7661696google scholar: lookup
        92. Mardassi H, Mounir S, Dea S. Molecular analysis of the ORFs 3 to 7 of porcine reproductive and respiratory syndrome virus, Québec reference strain. Arch Virol 1995;140(8):1405-18.
          doi: 10.1007/BF01322667pubmed: 7661693google scholar: lookup
        93. Morozov I, Meng XJ, Paul PS. Sequence analysis of open reading frames (ORFs) 2 to 4 of a U.S. isolate of porcine reproductive and respiratory syndrome virus. Arch Virol 1995;140(7):1313-9.
          doi: 10.1007/BF01322758pubmed: 7646363google scholar: lookup
        94. Anderson GW, Rowland RR, Palmer GA, Even C, Plagemann PG. Lactate dehydrogenase-elevating virus replication persists in liver, spleen, lymph node, and testis tissues and results in accumulation of viral RNA in germinal centers, concomitant with polyclonal activation of B cells. J Virol 1995 Aug;69(8):5177-85.
        95. de Vries AA, Post SM, Raamsman MJ, Horzinek MC, Rottier PJ. The two major envelope proteins of equine arteritis virus associate into disulfide-linked heterodimers. J Virol 1995 Aug;69(8):4668-74.
        96. Chirnside ED, Francis PM, de Vries AA, Sinclair R, Mumford JA. Development and evaluation of an ELISA using recombinant fusion protein to detect the presence of host antibody to equine arteritis virus. J Virol Methods 1995 Jul;54(1):1-13.
          doi: 10.1016/0166-0934(95)00020-upubmed: 7559853google scholar: lookup
        97. Nelson EA, Christopher-Hennings J, Drew T, Wensvoort G, Collins JE, Benfield DA. Differentiation of U.S. and European isolates of porcine reproductive and respiratory syndrome virus by monoclonal antibodies. J Clin Microbiol 1993 Dec;31(12):3184-9.