{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,26]],"date-time":"2026-04-26T08:57:05Z","timestamp":1777193825708,"version":"3.51.4"},"reference-count":58,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T00:00:00Z","timestamp":1773792000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000038","name":"U.S. Food and Drug Administration Medical Countermeasures Initiative","doi-asserted-by":"publisher","award":["75F40120C00085"],"award-info":[{"award-number":["75F40120C00085"]}],"id":[{"id":"10.13039\/100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000265","name":"Medical Research Council","doi-asserted-by":"publisher","award":["MR\/W005611\/1"],"award-info":[{"award-number":["MR\/W005611\/1"]}],"id":[{"id":"10.13039\/501100000265","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000265","name":"Medical Research Council","doi-asserted-by":"publisher","award":["MR\/Y004205\/1"],"award-info":[{"award-number":["MR\/Y004205\/1"]}],"id":[{"id":"10.13039\/501100000265","id-type":"DOI","asserted-by":"publisher"}]},{"name":"The Pandemic Institute"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>In this study, the phenotypic consequences of naturally occurring single nucleotide polymorphisms (SNPs) in the Middle East respiratory syndrome coronavirus (MERS-CoV) Spike protein were investigated. The impact of Spike mutations on the syncytia formation and neutralisation of contemporary MERS-CoV strains is not currently well understood. Mutations were identified by aligning 584 MERS-CoV Spike sequences from either human clinical isolates collected between 2012 and 2024 or from a clinical isolate that had been passaged in human cells. Fifteen SNPs of interest occurring in the N-terminal domain (NTD), receptor binding domain (RBD) and adjacent to the S1\/S2 cleavage site were selected for further characterisation based on their location in the Spike protein, frequency and identification in previous studies. A contemporary clade B, lineage 5 wildtype Spike sequence, obtained from a human MERS-CoV clinical isolate, was used as the backbone in this study. The mutations of interest were introduced to the wildtype backbone to generate Spike variants. Spike variants were characterised via cell\u2013cell fusion assays, and a lentiviral pseudotyping system was used to investigate the impact of these Spike mutations on neutralisation. The I529T, E536K and L745F mutations were shown to increase fusion and syncytia formation. The L411F, T424I, L506F, L745F and T746K mutations were found to increase resistance to neutralisation by pooled patient sera. This study has identified novel naturally occurring Spike mutations that resulted in phenotypic differences in the syncytia formation and neutralisation of contemporary MERS-CoV strains. Continued investigation of the phenotypic consequences of MERS-CoV Spike mutations is essential for assessing the risk to public health, especially given the pandemic potential of this virus.<\/jats:p>","DOI":"10.3390\/v18030377","type":"journal-article","created":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T09:10:04Z","timestamp":1773825004000},"page":"377","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Characterisation of Naturally Occurring MERS-CoV Spike Mutations and Their Impact on Fusion and Neutralisation"],"prefix":"10.3390","volume":"18","author":[{"given":"Rachael","family":"Dempsey","sequence":"first","affiliation":[{"name":"Institute of Infection, Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of Liverpool, Liverpool L69 7ZB, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4194-8714","authenticated-orcid":false,"given":"Hannah","family":"Goldswain","sequence":"additional","affiliation":[{"name":"Institute of Infection, Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of Liverpool, Liverpool L69 7ZB, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0717-8672","authenticated-orcid":false,"given":"Joseph","family":"Newman","sequence":"additional","affiliation":[{"name":"The Pirbright Institute, Pirbright, Woking, Surrey GU24 0NF, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4450-5911","authenticated-orcid":false,"given":"Nazia","family":"Thakur","sequence":"additional","affiliation":[{"name":"The Pirbright Institute, Pirbright, Woking, Surrey GU24 0NF, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tracy","family":"MacGill","sequence":"additional","affiliation":[{"name":"Office of Regulatory and Emerging Science, U.S. Food and Drug Administration, Silver Spring, MD 20993, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Todd","family":"Myers","sequence":"additional","affiliation":[{"name":"Office of Regulatory and Emerging Science, U.S. Food and Drug Administration, Silver Spring, MD 20993, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Robert","family":"Orr","sequence":"additional","affiliation":[{"name":"Office of Regulatory and Emerging Science, U.S. Food and Drug Administration, Silver Spring, MD 20993, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Dalan","family":"Bailey","sequence":"additional","affiliation":[{"name":"The Pirbright Institute, Pirbright, Woking, Surrey GU24 0NF, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8928-2037","authenticated-orcid":false,"given":"James P.","family":"Stewart","sequence":"additional","affiliation":[{"name":"Institute of Infection, Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of Liverpool, Liverpool L69 7ZB, UK"},{"name":"The Pandemic Institute, Liverpool L7 3FA, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Waleed","family":"Aljabr","sequence":"additional","affiliation":[{"name":"Institute of Infection, Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of Liverpool, Liverpool L69 7ZB, UK"},{"name":"Public Health Authority, Riyadh 13351, Saudi Arabia"},{"name":"King Fahad Medical City, Riyadh 12231, Saudi Arabia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Julian A.","family":"Hiscox","sequence":"additional","affiliation":[{"name":"Institute of Infection, Veterinary and Ecological Sciences, Faculty of Health and Life Sciences, University of Liverpool, Liverpool L69 7ZB, UK"},{"name":"The Pandemic Institute, Liverpool L7 3FA, UK"},{"name":"A*STAR Infectious Diseases (ID) Labs, Singapore 138648, Singapore"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2026,3,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1016\/j.tim.2016.03.003","article-title":"Epidemiology, Genetic Recombination, and Pathogenesis of Coronaviruses","volume":"24","author":"Su","year":"2016","journal-title":"Trends Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Mostafa, A., Kandeil, A., Shehata, M., El Shesheny, R., Samy, A.M., Kayali, G., and Ali, M.A. (2020). Middle East Respiratory Syndrome Coronavirus (MERS-CoV): State of the Science. Microorganisms, 8.","DOI":"10.3390\/microorganisms8070991"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"3144","DOI":"10.1073\/pnas.1718769115","article-title":"MERS coronaviruses from camels in Africa exhibit region-dependent genetic diversity","volume":"115","author":"Chu","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"57","DOI":"10.3201\/eid3101.241030","article-title":"Ongoing Evolution of Middle East Respiratory Syndrome Coronavirus, Saudi Arabia, 2023\u20132024","volume":"31","author":"Hassan","year":"2025","journal-title":"Emerg. Infect. Dis."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1080\/22221751.2021.2019559","article-title":"Enhanced replication fitness of MERS-CoV clade B over clade A strains in camelids explains the dominance of clade B strains in the Arabian Peninsula","volume":"11","author":"Te","year":"2022","journal-title":"Emerg. Microbes Infect."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1007\/s44197-025-00426-6","article-title":"Recurrent MERS-CoV Transmission in Saudi Arabia- Renewed Lessons in Healthcare Preparedness and Surveillance","volume":"15","author":"Memish","year":"2025","journal-title":"J. Epidemiol. Glob. Health"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"5324","DOI":"10.1038\/s41467-021-25519-1","article-title":"Functional comparison of MERS-coronavirus lineages reveals increased replicative fitness of the recombinant lineage 5","volume":"12","author":"Schroeder","year":"2021","journal-title":"Nat. Commun."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1093\/cid\/ciu812","article-title":"An observational, laboratory-based study of outbreaks of middle East respiratory syndrome coronavirus in Jeddah and Riyadh, kingdom of Saudi Arabia, 2014","volume":"60","author":"Drosten","year":"2015","journal-title":"Clin. Infect. Dis."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/j.jhin.2016.10.008","article-title":"Middle East respiratory syndrome coronavirus (MERS-CoV) outbreak in South Korea, 2015: Epidemiology, characteristics and public health implications","volume":"95","author":"Kim","year":"2017","journal-title":"J. Hosp. Infect."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"e2015049","DOI":"10.4178\/epih\/e2015049","article-title":"The first case of the 2015 Korean Middle East Respiratory Syndrome outbreak","volume":"37","author":"Park","year":"2015","journal-title":"Epidemiol. Health"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Lee, J.Y., Kim, Y.J., Chung, E.H., Kim, D.W., Jeong, I., Kim, Y., Yun, M.R., Kim, S.S., Kim, G., and Joh, J.S. (2017). The clinical and virological features of the first imported case causing MERS-CoV outbreak in South Korea, 2015. BMC Infect. Dis., 17.","DOI":"10.1186\/s12879-017-2576-5"},{"key":"ref_12","first-page":"846","article-title":"MERS-CoV Outbreak in Jeddah\u2014A Link to Health Care Facilities","volume":"372","year":"2014","journal-title":"N. Engl. J. Med."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1126\/science.aac8608","article-title":"Co-circulation of three camel coronavirus species and recombination of MERS-CoVs in Saudi Arabia","volume":"351","author":"Sabir","year":"2016","journal-title":"Science"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Karani, A., Ombok, C., Situma, S., Breiman, R., Mureithi, M., Jaoko, W., Njenga, M.K., and Ngere, I. (2025). Low-Level Zoonotic Transmission of Clade C MERS-CoV in Africa: Insights from Scoping Review and Cohort Studies in Hospital and Community Settings. Viruses, 17.","DOI":"10.3390\/v17010125"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"114530","DOI":"10.1016\/j.celrep.2024.114530","article-title":"Mapping immunodominant sites on the MERS-CoV spike glycoprotein targeted by infection-elicited antibodies in humans","volume":"43","author":"Addetia","year":"2024","journal-title":"Cell Rep."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"16292","DOI":"10.1038\/s41598-019-52730-4","article-title":"Humoral Immunogenicity and Efficacy of a Single Dose of ChAdOx1 MERS Vaccine Candidate in Dromedary Camels","volume":"9","author":"Alharbi","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1126\/science.aad1283","article-title":"An orthopoxvirus-based vaccine reduces virus excretion after MERS-CoV infection in dromedary camels","volume":"351","author":"Haagmans","year":"2016","journal-title":"Science"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"e11","DOI":"10.1016\/S2666-5247(21)00193-2","article-title":"Safety and immunogenicity of ChAdOx1 MERS vaccine candidate in healthy Middle Eastern adults (MERS002): An open-label, non-randomised, dose-escalation, phase 1b trial","volume":"3","author":"Bosaeed","year":"2022","journal-title":"Lancet Microbe"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"827","DOI":"10.1016\/S1473-3099(20)30248-6","article-title":"Safety and immunogenicity of a modified vaccinia virus Ankara vector vaccine candidate for Middle East respiratory syndrome: An open-label, phase 1 trial","volume":"20","author":"Koch","year":"2020","journal-title":"Lancet Infect. Dis."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1013","DOI":"10.1016\/S1473-3099(19)30266-X","article-title":"Safety and immunogenicity of an anti-Middle East respiratory syndrome coronavirus DNA vaccine: A phase 1, open-label, single-arm, dose-escalation trial","volume":"19","author":"Modjarrad","year":"2019","journal-title":"Lancet Infect. Dis."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1016\/S1473-3099(24)00423-7","article-title":"Safety, immunogenicity, and optimal dosing of a modified vaccinia Ankara-based vaccine against MERS-CoV in healthy adults: A phase 1b, double-blind, randomised placebo-controlled clinical trial","volume":"25","author":"Raadsen","year":"2025","journal-title":"Lancet Infect. Dis."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"So, R.T.Y., Hui, K.P.Y., Ho, J.C.W., Lau, K.K.M., Zhou, Z., Chan, M.C.W., Poon, L.L.M., and Peiris, M. (2026). The impact of clade B lineage 5 MERS coronaviruses spike mutations from 2015 to 2023 on virus entry and replication competence. PLoS Pathog., 22.","DOI":"10.1371\/journal.ppat.1013336"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"e01381-18","DOI":"10.1128\/JVI.01381-18","article-title":"Mutations in the Spike Protein of Middle East Respiratory Syndrome Coronavirus Transmitted in Korea Increase Resistance to Antibody-Mediated Neutralization","volume":"93","author":"Wang","year":"2019","journal-title":"J. Virol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"15092","DOI":"10.1038\/ncomms15092","article-title":"Cryo-EM structures of MERS-CoV and SARS-CoV spike glycoproteins reveal the dynamic receptor binding domains","volume":"8","author":"Yuan","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"15214","DOI":"10.1073\/pnas.1407087111","article-title":"Host cell entry of Middle East respiratory syndrome coronavirus after two-step, furin-mediated activation of the spike protein","volume":"111","author":"Millet","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"12262","DOI":"10.1073\/pnas.1608147113","article-title":"Proteolytic processing of Middle East respiratory syndrome coronavirus spikes expands virus tropism","volume":"113","author":"Park","year":"2016","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"9114","DOI":"10.1128\/JVI.01133-16","article-title":"Carcinoembryonic Antigen-Related Cell Adhesion Molecule 5 Is an Important Surface Attachment Factor That Facilitates Entry of Middle East Respiratory Syndrome Coronavirus","volume":"90","author":"Chan","year":"2016","journal-title":"J. Virol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"11709","DOI":"10.1074\/jbc.RA118.001897","article-title":"Middle East respiratory syndrome coronavirus and bat coronavirus HKU9 both can utilize GRP78 for attachment onto host cells","volume":"293","author":"Chu","year":"2018","journal-title":"J. Biol. Chem."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"E8508","DOI":"10.1073\/pnas.1712592114","article-title":"Identification of sialic acid-binding function for the Middle East respiratory syndrome coronavirus spike glycoprotein","volume":"114","author":"Li","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1038\/nature12005","article-title":"Dipeptidyl peptidase 4 is a functional receptor for the emerging human coronavirus-EMC","volume":"495","author":"Raj","year":"2013","journal-title":"Nature"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1038\/nature12328","article-title":"Molecular basis of binding between novel human coronavirus MERS-CoV and its receptor CD26","volume":"500","author":"Lu","year":"2013","journal-title":"Nature"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"516","DOI":"10.1080\/22221751.2019.1597644","article-title":"Towards a solution to MERS: Protective human monoclonal antibodies targeting different domains and functions of the MERS-coronavirus spike glycoprotein","volume":"8","author":"Widjaja","year":"2019","journal-title":"Emerg. Microbes Infect."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"7712","DOI":"10.1038\/ncomms8712","article-title":"Evaluation of candidate vaccine approaches for MERS-CoV","volume":"6","author":"Wang","year":"2015","journal-title":"Nat. Commun."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"e4249","DOI":"10.21769\/BioProtoc.4249","article-title":"Production of Recombinant Replication-defective Lentiviruses Bearing the SARS-CoV or SARS-CoV-2 Attachment Spike Glycoprotein and Their Application in Receptor Tropism and Neutralisation Assays","volume":"11","author":"Thakur","year":"2021","journal-title":"Bio. Protoc."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1016\/j.mex.2015.09.003","article-title":"An optimised method for the production of MERS-CoV spike expressing viral pseudotypes","volume":"2","author":"Grehan","year":"2015","journal-title":"MethodsX"},{"key":"ref_36","unstructured":"WHO (2025). Middle East Respiratory Syndrome Coronavirus (MERS-CoV) Dashboard, World Health Organisation. Available online: https:\/\/data.who.int\/dashboards\/mers\/cases."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"101282","DOI":"10.1016\/j.onehlt.2025.101282","article-title":"MERS-COV in the Middle East, a one health concept approach","volume":"21","author":"Dandachi","year":"2025","journal-title":"One Health"},{"key":"ref_38","unstructured":"WHO (2025). Disease Outbreak News: Middle East Respiratory Syndrome Coronavirus\u2014Kingdom of Saudi Arabia, World Health Organisation. Available online: https:\/\/www.who.int\/emergencies\/disease-outbreak-news\/item\/2025-DON569."},{"key":"ref_39","unstructured":"Rist, S. (2025, December 10). Two Cases of MERS-CoV, Returning from Abroad, Identified on National Territory, Available online: https:\/\/sante.gouv.fr\/actualites-presse\/presse\/communiques-de-presse\/article\/deux-cas-de-mers-cov-de-retour-de-l-etranger-identifies-sur-le-territoire."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"e00219","DOI":"10.1128\/mSphere.00219-21","article-title":"Amplicon and Metagenomic Analysis of Middle East Respiratory Syndrome (MERS) Coronavirus and the Microbiome in Patients with Severe MERS","volume":"6","author":"Aljabr","year":"2021","journal-title":"mSphere"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Qian, Z., Dominguez, S.R., and Holmes, K.V. (2013). Role of the Spike Glycoprotein of Human Middle East Respiratory Syndrome Coronavirus (MERS-CoV) in Virus Entry and Syncytia Formation. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0076469"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"871","DOI":"10.1038\/nbt0997-871","article-title":"Multiply attenuated lentiviral vector achieves efficient gene delivery in vivo","volume":"15","author":"Zufferey","year":"1997","journal-title":"Nat. Biotechnol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1089\/10430340252898984","article-title":"High-Level Transduction and Gene Expression in Hematopoietic Repopulating Cells Using a Human Imunodeficiency Virus Type 1-Based Lentiviral Vector Containing an Internal Spleen Focus Forming Virus Promoter","volume":"13","author":"Demaison","year":"2002","journal-title":"Hum. Gene Ther."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1180","DOI":"10.1038\/s41564-022-01163-3","article-title":"Neutralizing antibody activity against 21 SARS-CoV-2 variants in older adults vaccinated with BNT162b2","volume":"7","author":"Newman","year":"2022","journal-title":"Nat. Microbiol."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Conceicao, C., Thakur, N., Human, S., Kelly, J.T., Logan, L., Bialy, D., Bhat, S., Stevenson-Leggett, P., Zagrajek, A.K., and Hollinghurst, P. (2020). The SARS-CoV-2 Spike protein has a broad tropism for mammalian ACE2 proteins. PLoS Biol., 18.","DOI":"10.1101\/2020.06.17.156471"},{"key":"ref_46","unstructured":"Mattiuzzo, G., Bentley, E., Seo, S.H., Cho, N.H., Kim, J.O., Richardson, S., Hassall, M., Atkinson, E., Hockley, J., and Kim, Y.S. (2020). Establishment of 1st WHO International Standard for Anti-MERS-CoV Antibody, World Health Organisation. WHO\/BS\/2020.2398."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1249","DOI":"10.1093\/infdis\/jiy311","article-title":"Ultrapotent Human Neutralizing Antibody Repertoires Against Middle East Respiratory Syndrome Coronavirus From a Recovered Patient","volume":"218","author":"Niu","year":"2018","journal-title":"J. Infect. Dis."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"3395","DOI":"10.1016\/j.celrep.2019.08.052","article-title":"Structural Definition of a Neutralization-Sensitive Epitope on the MERS-CoV S1-NTD","volume":"28","author":"Wang","year":"2019","journal-title":"Cell Rep."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"7045","DOI":"10.1128\/JVI.00433-14","article-title":"A conformation-dependent neutralizing monoclonal antibody specifically targeting receptor-binding domain in Middle East respiratory syndrome coronavirus spike protein","volume":"88","author":"Du","year":"2014","journal-title":"J. Virol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Ferrara, F., and Temperton, N. (2018). Pseudotype Neutralization Assays: From Laboratory Bench to Data Analysis. Methods Protoc., 1.","DOI":"10.3390\/mps1010008"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1038\/cr.2013.92","article-title":"Structure of MERS-CoV spike receptor-binding domain complexed with human receptor DPP4","volume":"23","author":"Wang","year":"2013","journal-title":"Cell Res."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"E2018","DOI":"10.1073\/pnas.1402074111","article-title":"Identification of human neutralizing antibodies against MERS-CoV and their role in virus adaptive evolution","volume":"111","author":"Tang","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"e00019","DOI":"10.1128\/mBio.00019-16","article-title":"Spread of Mutant Middle East Respiratory Syndrome Coronavirus with Reduced Affinity to Human CD26 during the South Korean Outbreak","volume":"7","author":"Kim","year":"2016","journal-title":"mBio"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1161","DOI":"10.3201\/eid2506.181722","article-title":"Sequential Emergence and Wide Spread of Neutralization Escape Middle East Respiratory Syndrome Coronavirus Mutants, South Korea, 2015","volume":"25","author":"Kim","year":"2019","journal-title":"Emerg. Infect. Dis."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"e2102983118","DOI":"10.1073\/pnas.2102983118","article-title":"Middle East respiratory syndrome coronavirus Spike protein variants exhibit geographic differences in virulence","volume":"118","author":"Wong","year":"2021","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"e0130524","DOI":"10.1128\/jvi.01305-24","article-title":"Acquisition of a multibasic cleavage site does not increase MERS-CoV entry into Calu-3 human lung cells","volume":"98","author":"Hoffmann","year":"2024","journal-title":"J. Virol."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Peacock, T.P., Brown, J.C., Zhou, J., Thakur, N., Sukhova, K., Newman, J., Kugathasan, R., Yan, A.W.C., Furnon, W., and De Lorenzo, G. (2022). The altered entry pathway and antigenic distance of the SARS-CoV-2 Omicron variant map to separate domains of spike protein. bioRxiv.","DOI":"10.1101\/2021.12.31.474653"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"480","DOI":"10.1038\/s41467-025-68248-5","article-title":"Two-year persistence of MERS-CoV-specific antibody and T cell responses after MVA-MERS-S vaccination in healthy adults","volume":"17","author":"Mayer","year":"2026","journal-title":"Nat. Commun."}],"container-title":["Viruses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1999-4915\/18\/3\/377\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T09:13:18Z","timestamp":1773825198000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1999-4915\/18\/3\/377"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,18]]},"references-count":58,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2026,3]]}},"alternative-id":["v18030377"],"URL":"https:\/\/doi.org\/10.3390\/v18030377","relation":{},"ISSN":["1999-4915"],"issn-type":[{"value":"1999-4915","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,18]]}}}