{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,20]],"date-time":"2026-09-20T05:52:02Z","timestamp":1789883522281,"version":"4.0.1"},"reference-count":60,"publisher":"MDPI AG","issue":"6","license":[{"start":{"date-parts":[[2019,6,14]],"date-time":"2019-06-14T00:00:00Z","timestamp":1560470400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"European Union\u2019s Horizon 2020","award":["n\u00b0633184"],"award-info":[{"award-number":["n\u00b0633184"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>The Porcine Reproductive and Respiratory Syndrome Virus (PRRSV) induces reproductive disorders in sows and respiratory illnesses in growing pigs and is considered as one of the main pathogenic agents responsible for economic losses in the porcine industry worldwide. Modified live PRRSV vaccines (MLVs) are very effective vaccine types against homologous strains but they present only partial protection against heterologous viral variants. With the goal to induce broad and cross-protective immunity, we generated DNA vaccines encoding B and T antigens derived from a European subtype 1 strain that include T-cell epitope sequences known to be conserved across strains. These antigens were expressed either in a native form or in the form of vaccibodies targeted to the endocytic receptor XCR1 and CD11c expressed by different types of antigen-presenting cells (APCs). When delivered in skin with cationic nanoparticles and surface electroporation, multiple DNA vaccinations as a stand-alone regimen induced substantial antibody and T-cell responses, which were not promoted by targeting antigens to APCs. Interestingly, a DNA-MLV prime\u2013boost strategy strongly enhanced the antibody response and broadened the T-cell responses over the one induced by MLV or DNA-only. The anti-nucleoprotein antibody response induced by the DNA-MLV prime\u2013boost was clearly promoted by targeting the antigen to CD11c and XCR1, indicating a benefit of APC-targeting on the B-cell response. In conclusion, a DNA-MLV prime\u2013boost strategy, by enhancing the potency and breadth of MLV vaccines, stands as a promising vaccine strategy to improve the control of PRRSV in infected herds.<\/jats:p>","DOI":"10.3390\/v11060551","type":"journal-article","created":{"date-parts":[[2019,6,14]],"date-time":"2019-06-14T11:19:58Z","timestamp":1560511198000},"page":"551","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":14,"title":["A DNA-Modified Live Vaccine Prime\u2013Boost Strategy Broadens the T-Cell Response and Enhances the Antibody Response against the Porcine Reproductive and Respiratory Syndrome Virus"],"prefix":"10.3390","volume":"11","author":[{"given":"Cindy","family":"Bernelin-Cottet","sequence":"first","affiliation":[{"name":"VIM, INRA, Universit\u00e9 Paris-Saclay, Domaine de Vilvert, 78350 Jouy-en-Josas, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"C\u00e9line","family":"Urien","sequence":"additional","affiliation":[{"name":"VIM, INRA, Universit\u00e9 Paris-Saclay, Domaine de Vilvert, 78350 Jouy-en-Josas, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elisabeth","family":"Stubsrud","sequence":"additional","affiliation":[{"name":"Vaccibody AS, Gaustadalleen 21, 0349 Oslo, Norway"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Virginie","family":"Jakob","sequence":"additional","affiliation":[{"name":"Vaccine Formulation Laboratory, University of Lausanne, Chemin des Boveresses 155, 1066 Epalinges, Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edwige","family":"Bouguyon","sequence":"additional","affiliation":[{"name":"VIM, INRA, Universit\u00e9 Paris-Saclay, Domaine de Vilvert, 78350 Jouy-en-Josas, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elise","family":"Bordet","sequence":"additional","affiliation":[{"name":"VIM, INRA, Universit\u00e9 Paris-Saclay, Domaine de Vilvert, 78350 Jouy-en-Josas, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"C\u00e9line","family":"Barc","sequence":"additional","affiliation":[{"name":"Plate-Forme d\u2019Infectiologie Exp\u00e9rimentale-PFIE-UE1277, INRA, 37380 Nouzilly, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Olivier","family":"Boulesteix","sequence":"additional","affiliation":[{"name":"Plate-Forme d\u2019Infectiologie Exp\u00e9rimentale-PFIE-UE1277, INRA, 37380 Nouzilly, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vanessa","family":"Contreras","sequence":"additional","affiliation":[{"name":"Immunology of viral infections and autoimmune diseases, IDMIT Department, IBFJ, INSERM U1184-CEA-Universit\u00e9 Paris Sud 11, 92260 Fontenay-Aux-Roses et 94270 Le Kremlin-Bic\u00eatre, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christophe","family":"Barnier-Quer","sequence":"additional","affiliation":[{"name":"Vaccine Formulation Laboratory, University of Lausanne, Chemin des Boveresses 155, 1066 Epalinges, Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicolas","family":"Collin","sequence":"additional","affiliation":[{"name":"Vaccine Formulation Laboratory, University of Lausanne, Chemin des Boveresses 155, 1066 Epalinges, Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2849-5140","authenticated-orcid":false,"given":"Ivan","family":"Trus","sequence":"additional","affiliation":[{"name":"Laboratory of Virology, Faculty of Veterinary Medicine, Ghent University, Salisburylaan 133, B-9820 Merelbeke, 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Rev. Anim. Biosci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1007\/s00705-015-2672-z","article-title":"Reorganization and expansion of the nidoviral family Arteriviridae","volume":"161","author":"Kuhn","year":"2016","journal-title":"Arch. Virol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.vetmic.2011.01.008","article-title":"Genetic and immunobiological diversities of porcine reproductive and respiratory syndrome genotype I strains","volume":"150","author":"Darwich","year":"2011","journal-title":"Vet. Microbiol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"3704","DOI":"10.1016\/j.vaccine.2009.04.022","article-title":"Challenges for porcine reproductive and respiratory syndrome virus (PRRSV) vaccinology","volume":"27","author":"Kimman","year":"2009","journal-title":"Vaccine"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"4069","DOI":"10.1016\/j.vaccine.2015.06.092","article-title":"Live porcine reproductive and respiratory syndrome virus vaccines: Current status and future direction","volume":"33","author":"Renukaradhya","year":"2015","journal-title":"Vaccine"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1016\/j.tvjl.2007.05.022","article-title":"The challenge of PRRS immunology","volume":"177","author":"Mateu","year":"2008","journal-title":"Vet. J."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Rahe, M.C., and Murtaugh, M.P. (2017). Mechanisms of Adaptive Immunity to Porcine Reproductive and Respiratory Syndrome Virus. Viruses, 9.","DOI":"10.3390\/v9060148"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1707","DOI":"10.2460\/javma.2005.226.1707","article-title":"Correlation of cell-mediated immunity against porcine reproductive and respiratory syndrome virus with protection against reproductive failure in sows during outbreaks of porcine reproductive and respiratory syndrome in commercial herds","volume":"226","author":"Lowe","year":"2005","journal-title":"J. Am. Vet. Med. Assoc."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.virusres.2014.12.026","article-title":"Host-pathogen interactions during porcine reproductive and respiratory syndrome virus 1 infection of piglets","volume":"202","author":"Salguero","year":"2015","journal-title":"Virus Res."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"432","DOI":"10.1016\/j.vetmic.2014.05.030","article-title":"Evaluation of the efficacy of a new modified live porcine reproductive and respiratory syndrome virus (PRRSV) vaccine (Fostera PRRS) against heterologous PRRSV challenge","volume":"172","author":"Park","year":"2014","journal-title":"Vet. Microbiol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/S0042-6822(03)00009-6","article-title":"Gradual development of the interferon-gamma response of swine to porcine reproductive and respiratory syndrome virus infection or vaccination","volume":"309","author":"Meier","year":"2003","journal-title":"Virology"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1191","DOI":"10.1007\/s007050050578","article-title":"IFN gamma inhibits porcine reproductive and respiratory syndrome virus replication in macrophages","volume":"144","author":"Bautista","year":"1999","journal-title":"Arch. Virol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1038\/nri820","article-title":"No one is naive: The significance of heterologous T-cell immunity","volume":"2","author":"Welsh","year":"2002","journal-title":"Nat. Rev. Immunol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"40","DOI":"10.3389\/fimmu.2016.00040","article-title":"The Non-structural Protein 5 and Matrix Protein Are Antigenic Targets of T Cell Immunity to Genotype 1 Porcine Reproductive and Respiratory Syndrome Viruses","volume":"7","author":"Mokhtar","year":"2016","journal-title":"Front. Immunol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"6828","DOI":"10.1016\/j.vaccine.2014.04.054","article-title":"Proteome-wide screening of the European porcine reproductive and respiratory syndrome virus reveals a broad range of T cell antigen reactivity","volume":"32","author":"Mokhtar","year":"2014","journal-title":"Vaccine"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"663","DOI":"10.3390\/v5020663","article-title":"Predicted peptides from non-structural proteins of porcine reproductive and respiratory syndrome virus are able to induce IFN-gamma and IL-10","volume":"5","author":"Diaz","year":"2013","journal-title":"Viruses"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Tregoning, J.S., and Kinnear, E. (2014). Using Plasmids as DNA Vaccines for Infectious Diseases. Microbiol. Spectr., 2.","DOI":"10.1128\/microbiolspec.PLAS-0028-2014"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"61","DOI":"10.3858\/emm.2009.41.2.008","article-title":"Improving vaccines by targeting antigens to dendritic cells","volume":"41","author":"Shortman","year":"2009","journal-title":"Exp. Mol. Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1111\/j.1365-2796.2011.02496.x","article-title":"Dendritic cell-targeted protein vaccines: A novel approach to induce T-cell immunity","volume":"271","author":"Trumpfheller","year":"2012","journal-title":"J. Intern. Med."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"259","DOI":"10.4161\/hv.22487","article-title":"New generation of dendritic cell vaccines","volume":"9","author":"Radford","year":"2013","journal-title":"Hum. Vaccin. Immunother."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1484","DOI":"10.3389\/fimmu.2018.01484","article-title":"Dendritic Cells in the Cross Hair for the Generation of Tailored Vaccines","volume":"9","author":"Gornati","year":"2018","journal-title":"Front. Immunol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1427","DOI":"10.1172\/JCI34224","article-title":"The efficacy of DNA vaccination is enhanced in mice by targeting the encoded protein to dendritic cells","volume":"118","author":"Nchinda","year":"2008","journal-title":"J. Clin. Invest."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"6988","DOI":"10.1016\/j.vaccine.2015.08.094","article-title":"Targeting of nucleoprotein to chemokine receptors by DNA vaccination results in increased CD8(+)-mediated cross protection against influenza","volume":"33","author":"Baranowska","year":"2015","journal-title":"Vaccine"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"624","DOI":"10.1002\/eji.201445080","article-title":"Vaccine molecules targeting Xcr1 on cross-presenting DCs induce protective CD8(+) T-cell responses against influenza virus","volume":"45","author":"Fossum","year":"2015","journal-title":"Eur. J. Immunol."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Grodeland, G., Mjaaland, S., Tunheim, G., Fredriksen, A.B., and Bogen, B. (2013). The Specificity of Targeted Vaccines for APC Surface Molecules Influences the Immune Response Phenotype. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0080008"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1283","DOI":"10.1084\/jem.20100223","article-title":"The XC chemokine receptor 1 is a conserved selective marker of mammalian cells homologous to mouse CD8alpha+ dendritic cells","volume":"207","author":"Crozat","year":"2010","journal-title":"J. Exp. Med."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"3313","DOI":"10.4049\/jimmunol.1000824","article-title":"Existence of CD8 \u03b1-Like Dendritic Cells with a Conserved Functional Specialization and a Common Molecular Signature in Distant Mammalian Species","volume":"185","author":"Contreras","year":"2010","journal-title":"J. Immunol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1038\/mi.2015.105","article-title":"The respiratory DC\/macrophage network at steady-state and upon influenza infection in the swine biomedical model","volume":"9","author":"Maisonnasse","year":"2016","journal-title":"Mucosal Immunol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.dci.2016.06.015","article-title":"Expanding the tools for identifying mononuclear phagocyte subsets in swine: Reagents to porcine CD11c and XCR1","volume":"65","author":"Deloizy","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2072","DOI":"10.1002\/eji.200838645","article-title":"CD11c: Not merely a murine DC marker, but also a useful vaccination target","volume":"38","author":"Kurts","year":"2008","journal-title":"Eur. J. Immunol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2263","DOI":"10.1002\/eji.200838302","article-title":"CD11c provides an effective immunotarget for the generation of both CD4 and CD8 T cell responses","volume":"38","author":"Castro","year":"2008","journal-title":"Eur. J. Immunol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1084\/jem.20121251","article-title":"Antigen delivery to early endosomes eliminates the superiority of human blood BDCA3+ dendritic cells at cross presentation","volume":"210","author":"Cohn","year":"2013","journal-title":"J. Exp. Med."},{"key":"ref_33","first-page":"641","article-title":"A Universal Influenza Vaccine Can Lead to Disease Exacerbation or Viral Control Depending on Delivery Strategies","volume":"7","author":"Deloizy","year":"2016","journal-title":"Front. Immunol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"7852","DOI":"10.1016\/j.vaccine.2010.09.078","article-title":"In vivo electroporation enhances the potency of poly-lactide co-glycolide (PLG) plasmid DNA immunization","volume":"28","author":"Barbon","year":"2010","journal-title":"Vaccine"},{"key":"ref_35","unstructured":"Bernelin-Cottet, C., Urien, C., McCaffrey, J., Collins, D., Donadei, A., McDaid, D., Jakob, V., Barnier-Quer, C., Collin, N., and Bouguyon, E. (2019). Assessing the immunogenic properties of DNA vaccines delivered with nanoparticles, electroporation and dissolvable microneedle patches in pig skin, Submitted."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"4122","DOI":"10.1038\/s41598-017-04547-2","article-title":"Electroporation as a vaccine delivery system and a natural adjuvant to intradermal administration of plasmid DNA in macaques","volume":"7","author":"Todorova","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1089\/hgtb.2015.020","article-title":"Skin Transfection Patterns and Expression Kinetics of Electroporation-Enhanced Plasmid Delivery Using the CELLECTRA-3P, a Portable Next-Generation Dermal Electroporation Device","volume":"26","author":"Amante","year":"2015","journal-title":"Hum. Gene Ther. Methods"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1186\/s13567-015-0166-3","article-title":"Different clinical, virological, serological and tissue tropism outcomes of two new and one old Belgian type 1 subtype 1 porcine reproductive and respiratory virus (PRRSV) isolates","volume":"46","author":"Frydas","year":"2015","journal-title":"Vet. Res."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"6768","DOI":"10.1016\/j.vaccine.2014.10.005","article-title":"In vivo targeting of porcine reproductive and respiratory syndrome virus antigen through porcine DC-SIGN to dendritic cells elicits antigen-specific CD4T cell immunity in pigs","volume":"32","author":"Subramaniam","year":"2014","journal-title":"Vaccine"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1797","DOI":"10.1182\/blood-2006-06-032938","article-title":"Chemokine-idiotype fusion DNA vaccines are potentiated by bivalency and xenogeneic sequences","volume":"110","author":"Fredriksen","year":"2007","journal-title":"Blood"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"W435","DOI":"10.1093\/nar\/gkp254","article-title":"ANNIE: Integrated de novo protein sequence annotation","volume":"37","author":"Ooi","year":"2009","journal-title":"Nucleic Acids Res."},{"key":"ref_42","first-page":"299","article-title":"Defining Mononuclear Phagocyte Subset Homology Across Several Distant Warm-Blooded Vertebrates Through Comparative Transcriptomics","volume":"6","author":"Urien","year":"2015","journal-title":"Front. Immunol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1016\/j.rvsc.2018.10.005","article-title":"Comparison of six commercial ELISAs for the detection of antibodies against porcine reproductive and respiratory syndrome virus (PRRSV) in field serum samples","volume":"121","author":"Biernacka","year":"2018","journal-title":"Res. Vet. Sci."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Sattler, T., Wodak, E., Revilla-Fernandez, S., and Schmoll, F. (2014). Comparison of different commercial ELISAs for detection of antibodies against porcine respiratory and reproductive syndrome virus in serum. BMC Vet. Res., 10.","DOI":"10.1186\/s12917-014-0300-x"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/j.vetimm.2017.07.003","article-title":"Dendritic cell-targeted porcine reproductive and respiratory syndrome virus (PRRSV) antigens adjuvanted with polyinosinic-polycytidylic acid (poly (I:C)) induced non-protective immune responses against heterologous type 2 PRRSV challenge in pigs","volume":"190","author":"Subramaniam","year":"2017","journal-title":"Vet. Immunol. Immunopathol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/j.dci.2013.04.021","article-title":"Antigen targeting to APC: From mice to veterinary species","volume":"41","author":"Alvarez","year":"2013","journal-title":"Dev. Comp. Immunol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"2836","DOI":"10.1182\/blood-2012-09-452078","article-title":"Targeting dendritic cells--why bother?","volume":"121","author":"Kreutz","year":"2013","journal-title":"Blood"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1038\/nri3727","article-title":"Dendritic cell-targeted vaccines\u2014hope or hype?","volume":"14","author":"Kastenmuller","year":"2014","journal-title":"Nat. Rev. Immunol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"39250","DOI":"10.1038\/srep39250","article-title":"The presence of T cell epitopes is important for induction of antibody responses against antigens directed to DEC205+ dendritic cells","volume":"6","author":"Amorim","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.jhep.2014.08.026","article-title":"HCV epitope, homologous to multiple human protein sequences, induces a regulatory T cell response in infected patients","volume":"62","author":"Losikoff","year":"2015","journal-title":"J. Hepatol."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Tenbusch, M., Ignatius, R., Nchinda, G., Trumpfheller, C., Salazar, A.M., Topfer, K., Sauermann, U., Wagner, R., Hannaman, D., and Tenner-Racz, K. (2012). Immunogenicity of DNA vaccines encoding simian immunodeficiency virus antigen targeted to dendritic cells in rhesus macaques. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0039038"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1038\/s41541-017-0033-5","article-title":"Enhancing vaccine antibody responses by targeting Clec9A on dendritic cells","volume":"2","author":"Park","year":"2017","journal-title":"NPJ Vaccines"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2785","DOI":"10.4049\/jimmunol.1601881","article-title":"Targeting Influenza Virus Hemagglutinin to Xcr1+ Dendritic Cells in the Absence of Receptor-Mediated Endocytosis Enhances Protective Antibody Responses","volume":"198","author":"Gudjonsson","year":"2017","journal-title":"J. Immunol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"7639","DOI":"10.1038\/s41598-017-07372-9","article-title":"The anti-influenza M2e antibody response is promoted by XCR1 targeting in pig skin","volume":"7","author":"Deloizy","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.virusres.2018.01.015","article-title":"Porcine reproductive and respiratory syndrome virus neutralizing antibodies provide in vivo cross-protection to PRRSV1 and PRRSV2 viral challenge","volume":"248","author":"Robinson","year":"2018","journal-title":"Virus Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.virusres.2012.06.024","article-title":"Location of T-cell epitopes in nonstructural proteins 9 and 10 of type-II porcine reproductive and respiratory syndrome virus","volume":"169","author":"Parida","year":"2012","journal-title":"Virus Res."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1016\/j.vaccine.2015.12.043","article-title":"Transdermal delivery of plasmid encoding truncated nucleocapsid protein enhanced PRRSV-specific immune responses","volume":"34","author":"Suradhat","year":"2015","journal-title":"Vaccine"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Chung, C.J., Cha, S.H., Grimm, A.L., Ajithdoss, D., Rzepka, J., Chung, G., Yu, J., Davis, W.C., and Ho, C.S. (2018). Pigs that recover from porcine reproduction and respiratory syndrome virus infection develop cytotoxic CD4+CD8+ and CD4+CD8- T-cells that kill virus infected cells. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0203482"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1264","DOI":"10.1016\/j.vaccine.2005.09.041","article-title":"A DNA vaccine coding for gB and gD of pseudorabies virus (suid herpes type 1) primes the immune system in the presence of maternal immunity more efficiently than conventional vaccines","volume":"24","author":"Middel","year":"2006","journal-title":"Vaccine"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/j.vetmic.2016.07.014","article-title":"Maternally-derived antibodies (MDAs) impair piglets\u2019 humoral and cellular immune responses to vaccination against porcine reproductive and respiratory syndrome (PRRS)","volume":"192","author":"Fablet","year":"2016","journal-title":"Vet. Microbiol."}],"container-title":["Viruses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1999-4915\/11\/6\/551\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T12:58:31Z","timestamp":1760187511000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1999-4915\/11\/6\/551"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,6,14]]},"references-count":60,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2019,6]]}},"alternative-id":["v11060551"],"URL":"https:\/\/doi.org\/10.3390\/v11060551","relation":{},"ISSN":["1999-4915"],"issn-type":[{"value":"1999-4915","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,6,14]]}}}