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Journal of virology2003; 77(11); 6216-6226; doi: 10.1128/jvi.77.11.6216-6226.2003

Formation of disulfide-linked complexes between the three minor envelope glycoproteins (GP2b, GP3, and GP4) of equine arteritis virus.

Abstract: Equine arteritis virus (EAV) is an enveloped, positive-stranded RNA virus belonging to the family Arteriviridae of the order NIDOVIRALES: Six transmembrane proteins have been identified in EAV particles: the nonglycosylated membrane protein M and the glycoprotein GP(5) (previously named G(L)), which occur as disulfide-bonded heterodimers and are the major viral envelope proteins; the unglycosylated small envelope protein E; and the minor glycoproteins GP(2b) (formerly designated G(S)), GP(3), and GP(4). Analysis of the appearance of the GP(2b), GP(3), and GP(4) proteins in viral particles by gel electrophoresis under reducing and nonreducing conditions revealed the occurrence of two different covalently linked oligomeric complexes between these proteins, i.e., heterodimers of GP(2b) and GP(4) and heterotrimers of GP(2b), GP(3), and GP(4). Shortly after their release from infected cells, virions contained mainly cystine-linked GP(2b)/GP(4) heterodimers, which were subsequently converted into disulfide-bonded GP(2b)/GP(3)/GP(4) trimers through the covalent recruitment of GP(3). This process occurred faster at a higher pH but was arrested at 4 degrees C. Furthermore, the conversion was almost instantaneous in the presence of the thiol oxidant diamide. In contrast, the sulfhydryl-modifying agent N-ethylmaleimide inhibited the formation of disulfide-bonded GP(2b)/GP(3)/GP(4) trimers. Using sucrose density gradients, we could not demonstrate a noncovalent association of GP(3) with the cystine-linked GP(2b)/GP(4) dimer in freshly released virions, nor did we observe higher-order structures of the GP(2b)/GP(4) or GP(2b)/GP(3)/GP(4) complexes. Nevertheless, the instantaneous diamide-induced formation of disulfide-bonded GP(2b)/GP(3)/GP(4) heterotrimers at 4 degrees C suggests that the three minor glycoproteins of EAV are assembled as trimeric complexes. The existence of a noncovalent interaction between the cystine-linked GP(2b)/GP(4) dimer and GP(3) was also inferred from coexpression experiments showing that the presence of GP(3) increased the electrophoretic mobility of the disulfide-bonded GP(2b)/GP(4) dimers. Our study reveals that the minor envelope proteins of arteriviruses enter into both covalent and noncovalent interactions, the function of which has yet to be established.
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Publication Date: 2003-05-14 PubMed ID: 12743278PubMed Central: PMC155002DOI: 10.1128/jvi.77.11.6216-6226.2003Google Scholar: Lookup
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  • Journal Article

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 examines how certain proteins in the equine arteritis virus (EAV) form bonds with each other. The study found two kinds of protein complexes, each with various combinations of proteins. The findings suggest that these proteins come together in specific ways, though the purpose of these interactions is not currently known.

Understanding the Equine Arteritis Virus (EAV)

  • Equine arteritis virus is an RNA virus with six identified proteins in its particles.
  • Five of these proteins are transmembrane proteins, including the major ones M and GP(5), and the minor ones GP(2b), GP(3), and GP(4).
  • The latter three are identified as glycoproteins, i.e., proteins that have sugar groups attached to them.

Protein Complexes in EAV

  • The research found that proteins GP(2b), GP(3), and GP(4) are able to bond together to form different types of protein complexes.
  • Two different types of this bonding were identified: heterodimers of GP(2b) and GP(4), and heterotrimers of GP(2b), GP(3), and GP(4).
  • These complexes were found to be “covalently linked”, meaning they shared electron pairs in a chemical bond.

Changes in Protein Structures

  • Post release from infected cells, the virus contained mostly GP(2b)/GP(4) heterodimers.
  • Interestingly, these heterodimers were found to convert into GP(2b)/GP(3)/GP(4) heterotrimers by incorporating GP(3) into the complex, through the recruitment of a covalent bond.
  • This process was found to occur faster at higher pH levels, but halt completely at temperatures of around 4 degrees celsius.

Chemical Influences on Protein Complexes

  • This study also found that certain chemicals influenced how these protein complexes formed.
  • The presence of a thiol oxidant called diamide facilitated the almost instantaneous formation of the GP(2b)/GP(3)/GP(4) heterotrimers.
  • On the other hand, an agent called N-ethylmaleimide was found to inhibit trimer formation.

Keeping the Future in Mind

  • All these findings suggest that the minor glycoproteins of EAV are assembled in specific complex formations.
  • The exact functional purpose of these covalently bonded structures and their behaviour is not yet known, but this research provides a base for such understanding,
  • The research helps in piecing together the complexities of virus structure and function, serving as an important waypoint in confronting diseases like the EAV.

Cite This Article

APA
Wieringa R, de Vries AA, Rottier PJ. (2003). Formation of disulfide-linked complexes between the three minor envelope glycoproteins (GP2b, GP3, and GP4) of equine arteritis virus. J Virol, 77(11), 6216-6226. https://doi.org/10.1128/jvi.77.11.6216-6226.2003

Publication

Journal of virology
ISSN: 0022-538X
NlmUniqueID: 0113724
Country: United States
Language: English
Volume: 77
Issue: 11
Pages: 6216-6226

Researcher Affiliations

Wieringa, Roeland
  • Virology Division, Department of Infectious Diseases and Immunology, Faculty of Veterinary Medicine, Utrecht University, The Netherlands. r.wieringa@vet.uu.nl
de Vries, Antoine A F
    Rottier, Peter J M

      MeSH Terms

      • Animals
      • Cell Line
      • Centrifugation, Density Gradient
      • Cricetinae
      • Dimerization
      • Disulfides / chemistry
      • Electrophoresis, Agar Gel / methods
      • Equartevirus / metabolism
      • Horses
      • Precipitin Tests
      • Viral Envelope Proteins / chemistry
      • Viral Envelope Proteins / genetics
      • Viral Envelope Proteins / metabolism
      • Virion / metabolism

      References

      This article includes 39 references
      1. Abell BA, Brown DT. Sindbis virus membrane fusion is mediated by reduction of glycoprotein disulfide bridges at the cell surface.. J. Virol. 67:5496-5501.
        pmc: PMC237952pubmed: 8350409
      2. Buchholz UJ, Finke S, Conzelmann KK. Generation of bovine respiratory syncytial virus (BRSV) from cDNA: BRSV NS2 is not essential for virus replication in tissue culture, and the human RSV leader region acts as a functional BRSV genome promoter.. J. Virol. 73:251-259.
        pmc: PMC103829pubmed: 9847328
      3. den Boon JA, Kleijnen MF, Spaan WJM, Snijder EJ. Equine arteritis virus subgenomic mRNA synthesis: analysis of leader-body junctions and replicative-form RNAs.. J. Virol. 70:4291-4298.
        pmc: PMC190361pubmed: 8676451
      4. den Boon JA, Snijder EJ, Chirnside ED, de Vries AAF, Horzinek MC, Spaan WJM. Equine arteritis virus is not a togavirus but belongs to the coronaviruslike superfamily.. J. Virol. 65:2910-2920.
        pmc: PMC240924pubmed: 1851863
      5. de Vries AAF, Chirnside ED, Bredenbeek PJ, Gravestein LA, Horzinek MC, Spaan WJM. All subgenomic mRNAs of equine arteritis virus contain a common leader sequence.. Nucleic Acids Res. 18:3241-3247.
        pmc: PMC330929pubmed: 2162519
      6. de Vries AAF, Chirnside ED, Horzinek MC, Rottier PJM. Structural proteins of equine arteritis virus.. J. Virol. 66:6294-6303.
        pmc: PMC240121pubmed: 1328669
      7. de Vries AAF, Post SM, Raamsman MJB, Horzinek MC, Rottier PJM. The two major envelope proteins of equine arteritis virus associate into disulfide-linked heterodimers.. J. Virol. 69:4668-4674.
        pmc: PMC189270pubmed: 7609031
      8. de Vries AAF, Raamsman MJB, van Dijk HA, Horzinek MC, Rottier PJM. The small envelope glycoprotein (GS) of equine arteritis virus folds into three distinct monomers and a disulfide-linked dimer.. J. Virol. 69:3441-3448.
        pmc: PMC189056pubmed: 7745690
      9. D'Halluin JC. Virus assembly.. Curr. Top. Microbiol. Immunol. 199:47-66.
        pubmed: 7555060
      10. Dobbe JC, van der Meer Y, Spaan WJM, Snijder EJ. Construction of chimeric arteriviruses reveals that the ectodomain of the major glycoprotein is not the main determinant of equine arteritis virus tropism in cell culture.. Virology 288:283-294.
        pubmed: 11601900
      11. 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. 69:613-617.
        pmc: PMC188620pubmed: 7983766
      12. Faaberg KS, Plagemann PG. ORF 3 of lactate dehydrogenase-elevating virus encodes a soluble, nonstructural, highly glycosylated, and antigenic protein.. Virology 227:245-251.
        pubmed: 9007081
      13. Fenouillet E, Barbouche R, Courageot J, Miquelis R. The catalytic activity of protein disulfide isomerase is involved in human immunodeficiency virus envelope-mediated membrane fusion after CD4 cell binding.. J. Infect. Dis. 183:744-752.
        pubmed: 11181151
      14. Fuerst TR, Niles EG, Studier FW, Moss B. Eukaryotic transient-expression system based on recombinant vaccinia virus that synthesizes bacteriophage T7 RNA polymerase.. Proc. Natl. Acad. Sci. USA 83:8122-8126.
        pmc: PMC386879pubmed: 3095828
      15. Gallagher TM. Murine coronavirus membrane fusion is blocked by modification of thiols buried within the spike protein.. J. Virol. 70:4683-4690.
        pmc: PMC190404pubmed: 8676494
      16. Gallagher TM, Rueckert RR. Assembly-dependent maturation cleavage in provirions of a small icosahedral insect ribovirus.. J. Virol. 62:3399-3406.
        pmc: PMC253463pubmed: 3404580
      17. Glomb-Reinmund S, Kielian M. The role of low pH and disulfide shuffling in the entry and fusion of Semliki Forest virus and Sindbis virus.. Virology 248:372-381.
        pubmed: 9721245
      18. Gonin P, Mardassi H, Gagnon CA, Massie B, Dea S. A nonstructural and antigenic glycoprotein is encoded by ORF3 of the IAF-Klop strain of porcine reproductive and respiratory syndrome virus.. Arch. Virol. 143:1927-1940.
        pmc: PMC7086821pubmed: 9856081
      19. Hedges JF, Balasuriya UBR, MacLachlan NJ. The open reading frame 3 of equine arteritis virus encodes an immunogenic glycosylated, integral membrane protein.. Virology 264:92-98.
        pubmed: 10544133
      20. Horzinek MC, Maess J, Laufs R. Studies on the substructure of togaviruses. II. Analysis of equine arteritis, rubella, bovine viral diarrhea, and hog cholera viruses.. Arch. Gesamte Virusforsch. 33:306-318.
        pubmed: 4107033
      21. Hyllseth B. Structural proteins of equine arteritis virus.. Arch. Gesamte Virusforsch. 40:177-188.
        pubmed: 4633581
      22. Kosower NS, Kosower EM. Formation of disulfides with diamide.. Methods Enzymol. 143:264-270.
        pubmed: 3657543
      23. Laemmli UK. Cleavage of structural proteins during the assembly of the head of bacteriophage T4.. Nature 227:680-685.
        pubmed: 5432063
      24. Magnusson P, Hyllseth B, Marusyk H. Morphological studies on equine arteritis virus.. Arch. Gesamte Virusforsch. 30:105-112.
        pubmed: 4195609
      25. 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. 72:6298-6306.
        pmc: PMC109768pubmed: 9658068
      26. Mirazimi A, Mousavi-Jazi M, Sundqvist VA, Svensson L. Free thiol groups are essential for infectivity of human cytomegalovirus.. J. Gen. Virol. 80:2861-2865.
        pubmed: 10580047
      27. Molenkamp R, van Tol H, Rozier BCD, van der Meer Y, Spaan WJM, Snijder EJ. The arterivirus replicase is the only viral protein required for genome replication and subgenomic mRNA transcription.. J. Gen. Virol. 81:2491-2496.
        pubmed: 10993938
      28. Nermut MV, Hockley DJ. Comparative morphology and structural classification of retroviruses.. Curr. Top. Microbiol. Immunol. 214:1-24.
        pubmed: 8791723
      29. Ryser HJ, Levy EM, Mandel R, DiSciullo GJ. Inhibition of human immunodeficiency virus infection by agents that interfere with thiol-disulfide interchange upon virus-receptor interaction.. Proc. Natl. Acad. Sci. USA 91:4559-4563.
        pmc: PMC43825pubmed: 8183947
      30. Schneemann A, Zhong W, Gallagher TM, Rueckert RR. Maturation cleavage required for infectivity of a nodavirus.. J. Virol. 66:6728-6734.
        pmc: PMC240169pubmed: 1404613
      31. Snijder EJ, van Tol H, Pedersen KW, Raamsman MJB, de Vries AAF. Identification of a novel structural protein of arteriviruses.. J. Virol. 73:6335-6345.
        pmc: PMC112712pubmed: 10400725
      32. Sturman LS, Ricard CS, Holmes KV. Conformational change of the coronavirus peplomer glycoprotein at pH 8.0 and 37°C correlates with virus aggregation and virus-induced cell fusion.. J. Virol. 64:3042-3050.
        pmc: PMC249489pubmed: 2159562
      33. van Berlo MF, Rottier PJM, Spaan WJM, Horzinek MC. Equine arteritis virus-induced polypeptide synthesis.. J. Gen. Virol. 67:1543-1549.
        pubmed: 2426393
      34. van Nieuwstadt AP, Meulenberg JJM, van Essen-Zanbergen A, Petersen-den Besten A, Bende RJ, Moormann RJM, 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. 70:4767-4772.
        pmc: PMC190414pubmed: 8676504
      35. Verheije MH, Welting TJM, Jansen HT, Rottier PJM, Meulenberg JJM. Chimeric arteriviruses generated by swapping of the M protein ectodomain rule out a role of this domain in viral targeting.. Virology 302:364-373.
        pubmed: 12490397
      36. Vogt VM. Proteolytic processing and particle maturation.. Curr. Top. Microbiol. Immunol. 214:95-131.
        pubmed: 8791726
      37. Weiland E, Bolz S, Weiland F, Herbst W, Raamsman MJB, Rottier PJM, de Vries AAF. Monoclonal antibodies directed against conserved epitopes on the nucleocapsid protein and the major envelope glycoprotein of equine arteritis virus.. J. Clin. Microbiol. 38:2065-2075.
        pmc: PMC86730pubmed: 10834955
      38. Wieringa R, de Vries AAF, Raamsman MJB, Rottier PJM. Characterization of two new structural glycoproteins, GP3 and GP4, of equine arteritis virus.. J. Virol. 76:10829-10840.
        pmc: PMC136612pubmed: 12368326
      39. Zeegers JJW, van der Zeijst BAM, Horzinek MC. The structural proteins of equine arteritis virus.. Virology 73:200-205.
        pubmed: 183352

      Citations

      This article has been cited 31 times.
      1. Maloney SM, Shaw TM, Nennig KM, Larsen MS, Shah A, Kumar A, Marcotrigiano J, Grove J, Snijder EJ, Kirchdoerfer RN, Bailey AL. CD81 is a receptor for equine arteritis virus (family: Arteriviridae). mBio 2025 Jul 9;16(7):e0062325.
        doi: 10.1128/mbio.00623-25pubmed: 40422661google scholar: lookup
      2. Amona FM, Pang Y, Gong X, Wang Y, Fang X, Zhang C, Chen X. Mechanism of PRRSV infection and antiviral role of polyphenols. Virulence 2024 Dec;15(1):2417707.
        doi: 10.1080/21505594.2024.2417707pubmed: 39432383google scholar: lookup
      3. Matczuk AK, Zhang M, Veit M, Ugorski M. Expression of the Heterotrimeric GP2/GP3/GP4 Spike of an Arterivirus in Mammalian Cells. Viruses 2022 Apr 1;14(4).
        doi: 10.3390/v14040749pubmed: 35458479google scholar: lookup
      4. Pavesi A. Origin, Evolution and Stability of Overlapping Genes in Viruses: A Systematic Review. Genes (Basel) 2021 May 26;12(6).
        doi: 10.3390/genes12060809pubmed: 34073395google scholar: lookup
      5. Wang H, Shen L, Chen J, Liu X, Tan T, Hu Y, Bai X, Li Y, Tian K, Li N, Hu X. Deletion of CD163 Exon 7 Confers Resistance to Highly Pathogenic Porcine Reproductive and Respiratory Viruses on Pigs. Int J Biol Sci 2019;15(9):1993-2005.
        doi: 10.7150/ijbs.34269pubmed: 31523199google scholar: lookup
      6. Matczuk AK, Chodaczek G, Ugorski M. Production of Recombinant EAV with Tagged Structural Protein Gp3 to Study Artervirus Minor Protein Localization in Infected Cells. Viruses 2019 Aug 9;11(8).
        doi: 10.3390/v11080735pubmed: 31404947google scholar: lookup
      7. Xie J, Trus I, Oh D, Kvisgaard LK, Rappe JCF, Ruggli N, Vanderheijden N, Larsen LE, Lefèvre F, Nauwynck HJ. A Triple Amino Acid Substitution at Position 88/94/95 in Glycoprotein GP2a of Type 1 Porcine Reproductive and Respiratory Syndrome Virus (PRRSV1) Is Responsible for Adaptation to MARC-145 Cells. Viruses 2019 Jan 8;11(1).
        doi: 10.3390/v11010036pubmed: 30626009google scholar: lookup
      8. Pavesi A, Vianelli A, Chirico N, Bao Y, Blinkova O, Belshaw R, Firth A, Karlin D. Overlapping genes and the proteins they encode differ significantly in their sequence composition from non-overlapping genes. PLoS One 2018;13(10):e0202513.
        doi: 10.1371/journal.pone.0202513pubmed: 30339683google scholar: lookup
      9. Zhang M, Krabben L, Wang F, Veit M. Glycoprotein 3 of Porcine Reproductive and Respiratory Syndrome Virus Exhibits an Unusual Hairpin-Like Membrane Topology. J Virol 2018 Aug 1;92(15).
        doi: 10.1128/JVI.00660-18pubmed: 29769343google scholar: lookup
      10. Burkard C, Lillico SG, Reid E, Jackson B, Mileham AJ, Ait-Ali T, Whitelaw CB, Archibald AL. Precision engineering for PRRSV resistance in pigs: Macrophages from genome edited pigs lacking CD163 SRCR5 domain are fully resistant to both PRRSV genotypes while maintaining biological function. PLoS Pathog 2017 Feb;13(2):e1006206.
        doi: 10.1371/journal.ppat.1006206pubmed: 28231264google scholar: lookup
      11. Caì Y, Postnikova EN, Bernbaum JG, Yú SQ, Mazur S, Deiuliis NM, Radoshitzky SR, Lackemeyer MG, McCluskey A, Robinson PJ, Haucke V, Wahl-Jensen V, Bailey AL, Lauck M, Friedrich TC, O'Connor DH, Goldberg TL, Jahrling PB, Kuhn JH. Simian hemorrhagic fever virus cell entry is dependent on CD163 and uses a clathrin-mediated endocytosis-like pathway. J Virol 2015 Jan;89(1):844-56.
        doi: 10.1128/JVI.02697-14pubmed: 25355889google scholar: lookup
      12. 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.
        doi: 10.1016/j.virusres.2014.09.010pubmed: 25278143google scholar: lookup
      13. Vatter HA, Di H, Donaldson EF, Baric RS, Brinton MA. Each of the eight simian hemorrhagic fever virus minor structural proteins is functionally important. Virology 2014 Aug;462-463:351-62.
        doi: 10.1016/j.virol.2014.06.001pubmed: 25036340google scholar: lookup
      14. 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
      15. Matczuk AK, Kunec D, Veit M. Co-translational processing of glycoprotein 3 from equine arteritis virus: N-glycosylation adjacent to the signal peptide prevents cleavage. J Biol Chem 2013 Dec 6;288(49):35396-405.
        doi: 10.1074/jbc.M113.505420pubmed: 24142700google scholar: lookup
      16. 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
      17. Stantchev TS, Paciga M, Lankford CR, Schwartzkopff F, Broder CC, Clouse KA. Cell-type specific requirements for thiol/disulfide exchange during HIV-1 entry and infection. Retrovirology 2012 Dec 3;9:97.
        doi: 10.1186/1742-4690-9-97pubmed: 23206338google scholar: lookup
      18. Kabatek A, Veit M. Folding and oligomerization of the gp2b/gp3/gp4 spike proteins of equine arteritis virus in vitro. Viruses 2012 Mar;4(3):414-23.
        doi: 10.3390/v4030414pubmed: 22590679google scholar: lookup
      19. Tian D, Wei Z, Zevenhoven-Dobbe JC, Liu R, Tong G, Snijder EJ, Yuan S. Arterivirus minor envelope proteins are a major determinant of viral tropism in cell culture. J Virol 2012 Apr;86(7):3701-12.
        doi: 10.1128/JVI.06836-11pubmed: 22258262google scholar: lookup
      20. Du Y, Pattnaik AK, Song C, Yoo D, Li G. Glycosyl-phosphatidylinositol (GPI)-anchored membrane association of the porcine reproductive and respiratory syndrome virus GP4 glycoprotein and its co-localization with CD163 in lipid rafts. Virology 2012 Mar 1;424(1):18-32.
        doi: 10.1016/j.virol.2011.12.009pubmed: 22222209google scholar: lookup
      21. Dokland T. The structural biology of PRRSV. Virus Res 2010 Dec;154(1-2):86-97.
        doi: 10.1016/j.virusres.2010.07.029pubmed: 20692304google scholar: lookup
      22. Yoo D, Song C, Sun Y, Du Y, Kim O, Liu HC. Modulation of host cell responses and evasion strategies for porcine reproductive and respiratory syndrome virus. Virus Res 2010 Dec;154(1-2):48-60.
        doi: 10.1016/j.virusres.2010.07.019pubmed: 20655963google scholar: lookup
      23. Go YY, Zhang J, Timoney PJ, Cook RF, Horohov DW, Balasuriya UB. Complex interactions between the major and minor envelope proteins of equine arteritis virus determine its tropism for equine CD3+ T lymphocytes and CD14+ monocytes. J Virol 2010 May;84(10):4898-911.
        doi: 10.1128/JVI.02743-09pubmed: 20219931google scholar: lookup
      24. 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
      25. Du Y, Zuckermann FA, Yoo D. Myristoylation of the small envelope protein of porcine reproductive and respiratory syndrome virus is non-essential for virus infectivity but promotes its growth. Virus Res 2010 Feb;147(2):294-9.
        doi: 10.1016/j.virusres.2009.11.016pubmed: 19951726google scholar: lookup
      26. 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
      27. Wissink EH, Kroese MV, van Wijk HA, Rijsewijk FA, Meulenberg JJ, Rottier PJ. Envelope protein requirements for the assembly of infectious virions of porcine reproductive and respiratory syndrome virus. J Virol 2005 Oct;79(19):12495-506.
        doi: 10.1128/JVI.79.19.12495-12506.2005pubmed: 16160177google scholar: lookup
      28. Wu WH, Fang Y, Rowland RR, Lawson SR, Christopher-Hennings J, Yoon KJ, Nelson EA. The 2b protein as a minor structural component of PRRSV. Virus Res 2005 Dec;114(1-2):177-81.
        doi: 10.1016/j.virusres.2005.06.014pubmed: 16095746google scholar: lookup
      29. 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.
        doi: 10.1128/JVI.78.23.13019-13027.2004pubmed: 15542653google scholar: lookup
      30. 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.
        doi: 10.1128/jvi.77.24.12996-13004.2003pubmed: 14645556google scholar: lookup
      31. 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.
        doi: 10.1128/jvi.77.15.8470-8480.2003pubmed: 12857916google scholar: lookup
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