{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T14:59:07Z","timestamp":1786719547624,"version":"build-2736575974"},"reference-count":48,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T00:00:00Z","timestamp":1786665600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T00:00:00Z","timestamp":1786665600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100002352","name":"Ain Shams University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100002352","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Virol J"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>Foot-and-mouth disease (FMD) considers as one of the extremely infectious diseases with major economic impact on cloven-hoofed animals globally. The causative agent of the disease, Foot and Mouth Disease Virus (FMDV), comprises seven serotypes designated as; O, A, SAT1, SAT2, SAT3, C, and Asia1. Over the past decades, studies have shown that glycyrrhizic acid, the primary bioactive compound of licorice root, and chitosan, as a biodegradable and biocompatible natural polymer derived from crustacean shells, are potent candidates for inhibition of FMDV activity. The current work evaluated the potential antiviral efficacy of glycyrrhizic acid nanoparticles (GA-NPs) and chitosan nanoparticles (Ch-NPs) against different FMDV serotypes through computational (molecular docking), in vitro and in vivo assays.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Materials and methods<\/jats:title>\n                    <jats:p>The study included the use of field isolates from local FMD viral outbreaks (O PanAsia-2, A Iran 05, A Africa 2020, A Africa G IV Egypt 2022, A Venezuela, and SAT2\/EGY\/2012); Glycyrrhizic acid and chitosan nanoparticles were prepared and characterized; antiviral activity was assessed by molecular docking analysis, BHK-21 cell assays, and experimental animal studies using representative FMDV serotypes.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Molecular docking showed that GA-NPs exhibited stronger binding affinity toward the FMDV RNA-dependent RNA polymerase (3D\n                      <jats:sup>pol<\/jats:sup>\n                      ) with binding energies of -8.974\u00a0kcal\/mol while Ch-NPs demonstrated superior binding affinity toward the FMDV VP3 protein with binding energies of -7.173\u00a0kcal\/mol, respectively. At a concentration of 2\u2009\u00d7\u200910\n                      <jats:sup>\u2212\u20091<\/jats:sup>\n                      mg\/ml, both nanoparticles significantly suppressed viral replication, yielding a reduction in viral titers 8, 7.5, 7, 7, 7.5, and 6.5 log\u2081\u2080 TCID\u2085\u2080\/ml for O PanAsia-2, A Iran 05, A Africa 2020, A Africa G IV Egypt 2022, A Venezuela, and SAT2\/EGY\/2012, respectively which can aid in the disease restriction during outbreaks. Otherwise, at 2\u2009\u00d7\u200910\n                      <jats:sup>\u2212\u20092<\/jats:sup>\n                      mg\/ml, treatment markedly reduced viral dissemination in treated guinea pigs, providing protection rates of 66.7%; 44.4%; 66.7%; 100%; and 77.8% and 100% against serotype O; A Iran 05; A Africa 2020; A Africa G IV Egypt 2022; A Venezuela; and SAT2, compared with untreated controls.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusion<\/jats:title>\n                    <jats:p>These findings highlighted that GA-NPs and Ch-NPs possess broad-spectrum antiviral activity against six FMDV serotypes, supporting their further development as nanoparticle-based antiviral agents for the control of FMD.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s12985-026-03271-6","type":"journal-article","created":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T13:45:02Z","timestamp":1786715102000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["In silico, in vitro, and in vivo evaluation of the antiviral potential of glycyrrhizic acid and chitosan nanoparticles against foot and mouth disease virus"],"prefix":"10.1186","volume":"23","author":[{"given":"Asmaa R.F.","family":"Eweis","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Khodeir M.","family":"Hassan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sahar A.H.","family":"Shoman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hoda A.","family":"Taha","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ehab E.","family":"Ibrahim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tahsin","family":"Shoala","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,8,14]]},"reference":[{"issue":"9\u201310","key":"3271_CR1","doi-asserted-by":"publisher","first-page":"1097","DOI":"10.1016\/S0264-410X(00)00337-6","volume":"19","author":"AV Iyer","year":"2001","unstructured":"Iyer AV, Ghosh S, Singh SN, Deshmukh RA. Evaluation of three \u2018ready to formulate\u2019 oil adjuvants for foot and mouth disease vaccine production. Vaccine. 2001;19(9\u201310):1097\u2013105. https:\/\/doi.org\/10.1016\/S0264-410X(00)00337-6.","journal-title":"Vaccine"},{"issue":"4","key":"3271_CR2","doi-asserted-by":"publisher","first-page":"1079","DOI":"10.1111\/tbed.12507","volume":"64","author":"TJD Knight-Jones","year":"2017","unstructured":"Knight-Jones TJD, McLaws M, Rushton J. Foot-and-Mouth Disease Impact on Smallholders - What Do We Know, What Don\u2019t We Know and How Can We Find Out More? Transbound Emerg Dis. 2017;64(4):1079\u201394. https:\/\/doi.org\/10.1111\/tbed.12507.","journal-title":"Transbound Emerg Dis"},{"issue":"3","key":"3271_CR3","doi-asserted-by":"publisher","first-page":"637","DOI":"10.20506\/rst.21.3.1356","volume":"21","author":"AD James","year":"2002","unstructured":"James AD, Rushton J. The economics of foot and mouth disease. Rev Sci Tech. 2002;21(3):637\u201344. https:\/\/doi.org\/10.20506\/rst.21.3.1356.","journal-title":"Rev Sci Tech"},{"key":"3271_CR4","doi-asserted-by":"publisher","unstructured":"Vosloo W, Thomson GR. Natural habitats in which foot-and-mouth disease viruses are maintained. Horizon Sci Press. 2004;383\u2013410. https:\/\/doi.org\/10.1201\/9781420037968.ch14.","DOI":"10.1201\/9781420037968.ch14"},{"issue":"1","key":"3271_CR5","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/s0168-1702(02)00259-9","volume":"91","author":"E Domingo","year":"2003","unstructured":"Domingo E, Escarmis C, Baronowski E, RuizJarabo CM, Carrillo E, Nunez J, Sorbino F. Evolution of foot and mouth disease virus. Virol Res. 2003;91(1):47\u201363. https:\/\/doi.org\/10.1016\/s0168-1702(02)00259-9.","journal-title":"Virol Res"},{"key":"3271_CR6","unstructured":"Ciara M, Zhidong Z, Melvyn Q, Jeannette K, Soren A. Int Symp. 2002;17-20 Sep., Turkey. App., 20."},{"key":"3271_CR7","unstructured":"King S, Jarvie H, Dobson P. nanoparticle. Encyclopedia Britannica. 2025. https:\/\/www.britannica.com\/science\/nanoparticle"},{"issue":"1","key":"3271_CR8","doi-asserted-by":"publisher","DOI":"10.1155\/2021\/6699560","volume":"2021","author":"SA Richard","year":"2021","unstructured":"Richard SA. Exploring the pivotal immunomodulatory and anti-inflammatory potentials of glycyrrhizic and glycyrrhetinic acids. Mediators Inflamm. 2021;6699560. https:\/\/doi.org\/10.1155\/2021\/6699560.","journal-title":"Mediators Inflamm"},{"issue":"2","key":"3271_CR9","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1016\/j.antiviral.2012.11.006","volume":"97","author":"A Milewska","year":"2013","unstructured":"Milewska A, Ciejka J, Kaminski K, Karewicz A, Bielska D, Zeglen S. Novel polymeric inhibitors of HCoV-NL63. Antivir Res. 2013;97(2):112\u201321. https:\/\/doi.org\/10.1016\/j.antiviral.2012.11.006.","journal-title":"Antivir Res"},{"issue":"8","key":"3271_CR10","doi-asserted-by":"publisher","first-page":"2999","DOI":"10.1016\/j.matpr.2020.10.055","volume":"49","author":"V Srivastava","year":"2022","unstructured":"Srivastava V, Yadav A, Sarkar P. Molecular docking and ADMET study of bioactive compounds of Glycyrrhiza glabra against main protease of SARS-CoV2. Mater Today Proc. 2022;49(8):2999\u20133007. https:\/\/doi.org\/10.1016\/j.matpr.2020.10.055.","journal-title":"Mater Today Proc"},{"issue":"6","key":"3271_CR11","doi-asserted-by":"publisher","first-page":"190","DOI":"10.7324\/JAPS.2025.197023","volume":"15","author":"AR Supekar","year":"2025","unstructured":"Supekar AR, Bhujbal S, Yadav R. Molecular docking and molecular dynamics simulation of glycyrrhizic acid in multitarget agents as potential inhibitors of respiratory influenza viruses. J App Pharm Sci. 2025;15(6):190\u2013204. https:\/\/doi.org\/10.7324\/JAPS.2025.197023.","journal-title":"J App Pharm Sci"},{"issue":"6","key":"3271_CR12","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1080\/08927022.2021.1880576","volume":"47","author":"DSNBK Prasanth","year":"2021","unstructured":"Prasanth DSNBK, Manikanta M, Chandramohan V, Bhavya G, Rao AL, Panda SP, Rao GSNK, Chakravarthi G, Teja N, Rani PS, Ashu G, Purnadurganjali C, Akhil P, Bhavani GV, Jaswitha T. In silico strategies of some selected phytoconstituents from Melissa officinalis as SARS CoV-2 main protease and spike protein (COVID-19) inhibitors. Mol Simul. 2021;47(6):457\u201370. . https:\/\/doi.org\/10.1080\/08927022.2021.1880576","journal-title":"Mol Simul"},{"issue":"5","key":"3271_CR13","doi-asserted-by":"publisher","first-page":"509","DOI":"10.3390\/jof8050509","volume":"8","author":"EO Hassan","year":"2022","unstructured":"Hassan EO, Shoala T, Attia AMF, Badr OAM, Mahmoud SYM, Farrag ESH, EL-Fiki IAI. Chitosan and Nano-Chitosan for Management of Harpophora maydis: Approaches for Investigating Antifungal Activity, Pathogenicity, Maize-Resistant Lines, and Molecular Diagnosis of Plant Infection. J Fungi. 2022;8(5):509. https:\/\/doi.org\/10.3390\/jof8050509.","journal-title":"J Fungi"},{"issue":"01","key":"3271_CR14","doi-asserted-by":"publisher","first-page":"47196","DOI":"10.1007\/s11356-021-14022-1","volume":"28","author":"A Amer","year":"2021","unstructured":"Amer A, Ghoneim M, Shoala T, Mohamed HI. Comparative studies of eco-friendly compounds like humic acid, salicylic, and glycyrrhizic acids and their nanocomposites on French basil (Ocimum basilicum L. cv. Grand verde). Environ Sci Pollut Res. 2021;28(01):47196\u2013212. https:\/\/doi.org\/10.1007\/s11356-021-14022-1.","journal-title":"Environ Sci Pollut Res"},{"issue":"25","key":"3271_CR15","doi-asserted-by":"publisher","first-page":"15775","DOI":"10.1039\/d2ra00905f","volume":"12","author":"SA Loutfy","year":"2022","unstructured":"Loutfy SA, Abdel-Salam AI, Moatasim Y, Gomaa MR, Abdel Fattah NF, Emam MH, et al. Antiviral activity of chitosan nanoparticles encapsulating silymarin (Sil\u2013CNPs) against SARS-CoV-2 (in silico and in vitro study). RSC Adv. 2022;12(25):15775\u201386. . https:\/\/doi.org\/10.1039\/d2ra00905f","journal-title":"RSC Adv"},{"key":"3271_CR16","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/0042-6822(62)90290-8","volume":"16","author":"JA Macpherson","year":"1962","unstructured":"Macpherson JA, Stocker NG. Polyoma transformation of hamster cell clones - an investigation of genetic factors affecting all competence. Virology. 1962;16:147\u201351. https:\/\/doi.org\/10.1016\/0042-6822(62)90290-8.","journal-title":"Virology"},{"key":"3271_CR17","unstructured":"Saad MAA, Fawzy HGIV. Antiviral action of ribavirin on foot and mouth disease virus replication in cell culture and laboratory animals. 7th Veterinary Medicine-Zagazig Conference (21-23 July, 2004), Sharm El-Sheikh."},{"issue":"1","key":"3271_CR18","doi-asserted-by":"publisher","first-page":"24","DOI":"10.1128\/aac.14.1.24","volume":"14","author":"JT Richards","year":"1978","unstructured":"Richards JT, Kern ER, Glasgow LA, Overall JJ, Deign EF, Hatch MT. Antiviral activity of extracts from marine algae. Antimicrob Agents Chemother. 1978;14(1):24\u201330. https:\/\/doi.org\/10.1128\/aac.14.1.24.","journal-title":"Antimicrob Agents Chemother"},{"issue":"2","key":"3271_CR19","first-page":"221","volume":"15","author":"EM Abeer","year":"2005","unstructured":"Abeer EM, Mansour. Trials for production of FMD cell culture vaccine grown in medium containing serum replacement. Beni-Suef Vet Med J Vol. 2005;15(2):221\u20136.","journal-title":"Beni-Suef Vet Med J Vol"},{"issue":"3","key":"3271_CR20","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1093\/oxfordjournals.aje.a118408","volume":"27","author":"LJ Reed","year":"1938","unstructured":"Reed LJ, Muench H. A simple method for estimating 50% endpoints. Am J Epidemiol. 1938;27(3):493\u20137. https:\/\/doi.org\/10.1093\/oxfordjournals.aje.a118408.","journal-title":"Am J Epidemiol"},{"key":"3271_CR21","unstructured":"Boseila AAH, Hatab EA. Evaluation of the antiviral activity of sage leaves (Salvia officinalis) extract against foot-and-mouth disease virus. Agric Relat. (FAR). Egypt. 2011;3-5 October. pp:301-308. https:\/\/api.semanticscholar.org\/CorpusID:83097672"},{"issue":"12","key":"3271_CR22","doi-asserted-by":"publisher","first-page":"052","DOI":"10.7324\/JAPS.2013.31209","volume":"3","author":"FK El-Baz","year":"2013","unstructured":"El-Baz FK, El-Senousy WM, El-Sayed AB, Kamel MM. In vitro antiviral and antimicrobial activities of Spirulina platensis. J Appl Pharm Sci. 2013;3(12):052\u20136. https:\/\/doi.org\/10.7324\/JAPS.2013.31209.","journal-title":"J Appl Pharm Sci"},{"issue":"6","key":"3271_CR23","first-page":"2250","volume":"6","author":"AM Mansour","year":"2016","unstructured":"Mansour AM, Elfiky AA, Fahmy A, Diab A. Therapeutic effect of HBV formulation in the treatment of FMD viral infection: preclinical and clinical evaluation. Int J Sci Res Publ. 2016;6(6):2250\u20133153.","journal-title":"Int J Sci Res Publ"},{"key":"3271_CR24","doi-asserted-by":"crossref","unstructured":"Zhao Z, Xiao Y, Xu L, Liu Y, Jiang G, Wang W, Li B, Zhu T, Tan Q, Tang L, Zhou H, Huang X, Shan H. Glycyrrhizic Acid Nanoparticles as Antiviral and Anti-inflammatory Agents for COVID-19 Treatment. ACS Appl Mater Interfaces., Dilnawaz F, Acharya S, Kanungo A. A clinical perspective of chitosan nanoparticles for infectious disease management. Polym Bull. 2023;81(2):1071-1095. DOI: 10.1007\/s00289-023-04755-z.","DOI":"10.1007\/s00289-023-04755-z"},{"key":"3271_CR25","doi-asserted-by":"crossref","unstructured":"Dilnawaz F, Acharya S, Kanungo A. A clinical perspective of chitosan nanoparticles for infectious disease management. Polym Bull. 2023;81(2):1071-1095. DOI: 10.1007\/s00289-023-04755-z","DOI":"10.1007\/s00289-023-04755-z"},{"issue":"12","key":"3271_CR26","doi-asserted-by":"publisher","first-page":"11520","DOI":"10.1074\/jbc.M413254200","volume":"280","author":"JR Birtley","year":"2005","unstructured":"Birtley JR, Knox SR, Jaulent AM, Brick P, Leatherbarrow RJ, Curry S. Crystal structure of foot-and-mouth disease virus 3C protease new insights into catalytic mechanism and cleavage specificity. J Biol Chem. 2005;280(12):11520\u20137. . https:\/\/doi.org\/10.1074\/jbc.M413254200","journal-title":"J Biol Chem"},{"issue":"1","key":"3271_CR27","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1146\/annurev.mi.44.100190.003131","volume":"44","author":"AC Palmenberg","year":"1990","unstructured":"Palmenberg AC. Proteolytic processing of picornaviral polyprotein. Annu Rev Microbiol. 1990;44(1):603\u201323. https:\/\/doi.org\/10.1146\/annurev.mi.44.100190.003131.","journal-title":"Annu Rev Microbiol"},{"issue":"2","key":"3271_CR28","doi-asserted-by":"publisher","first-page":"364","DOI":"10.1016\/0042-6822(90)90090-E","volume":"174","author":"M Tesar","year":"1990","unstructured":"Tesar M, Marquardt O. Foot-and-mouth disease virus protease 3C inhibits cellular transcription and mediates cleavage of histone H3. Virology. 1990;174(2):364\u201374. https:\/\/doi.org\/10.1016\/0042-6822(90)90090-E.","journal-title":"Virology"},{"issue":"1","key":"3271_CR29","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1023\/A:1008174100886","volume":"22","author":"G Manju","year":"2001","unstructured":"Manju G, Ramamurthy V, Bramhadev P, Aniket S, Gurumurthy CB, Divakar H, et al. Sequence analysis of the RNA polymerase gene of foot-and-mouth disease virus serotype Asia1. Virus Genes. 2001;22(1):21\u20136. https:\/\/doi.org\/10.1023\/A:1008174100886.","journal-title":"Virus Genes"},{"issue":"1","key":"3271_CR30","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1128\/jvi.46.1.311-316.1983","volume":"46","author":"BH Robertson","year":"1983","unstructured":"Robertson BH, Moore DM, Grubman MJ, Kleid DG. Identification of an exposed region of the immunogenic capsid polypeptide VP1 on foot-and-mouth disease virus. J Virol. 1983;46(1):311\u20136. https:\/\/doi.org\/10.1128\/jvi.46.1.311-316.1983.","journal-title":"J Virol"},{"issue":"1","key":"3271_CR31","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1186\/s12985-016-0561-z","volume":"13","author":"Y Gao","year":"2016","unstructured":"Gao Y, Sun SQ, Guo HC. Biological function of Foot-and-mouth disease virus non-structural proteins and non-coding elements. Virol J. 2016;13(1):107. https:\/\/doi.org\/10.1186\/s12985-016-0561-z.","journal-title":"Virol J"},{"issue":"4","key":"3271_CR32","doi-asserted-by":"publisher","first-page":"880","DOI":"10.1038\/sj.emboj.7600971","volume":"25","author":"C Ferrer-Orta","year":"2006","unstructured":"Ferrer-Orta C, Arias A, Agudo R, P\u00e9rez\u2010Luque R, Escarm\u00eds C, Domingo E, et al. The structure of a protein primer\u2013polymerase complex in the initiation of genome replication. EMBO J. 2006;25(4):880\u20138. https:\/\/doi.org\/10.1038\/sj.emboj.7600971.","journal-title":"EMBO J"},{"issue":"8","key":"3271_CR33","doi-asserted-by":"publisher","first-page":"1109","DOI":"10.1016\/S0969-2126(97)00261-X","volume":"5","author":"JL Hansen","year":"1997","unstructured":"Hansen JL, Long AM, Schultz SC. Structure of the RNA-dependent RNA polymerase of poliovirus. Structure. 1997;5(8):1109\u201322. https:\/\/doi.org\/10.1016\/S0969-2126(97)00261-X.","journal-title":"Structure"},{"issue":"7","key":"3271_CR34","doi-asserted-by":"publisher","first-page":"1909","DOI":"10.1099\/vir.0.80730-0","volume":"86","author":"EE Fry","year":"2005","unstructured":"Fry EE, Newman JW, Curry S, Najjam S, Jackson T, Blakemore W, Lea SM, Miller L, Burman A, King AM, Stuart DI. Structure of Foot-and-mouth disease virus serotype A1061 alone and complexed with oligosaccharide receptor: receptor conservation in the face of antigenic variation. J Gen Virol. 2005;86(7):1909\u201320. https:\/\/doi.org\/10.1099\/vir.0.80730-0.","journal-title":"J Gen Virol"},{"issue":"3","key":"3271_CR35","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1099\/0022-1317-4-3-313","volume":"4","author":"TF Wild","year":"1969","unstructured":"Wild TF, Burroughs JN, Brown F. Surface structure of foot-and-mouth disease virus. J Gen Virol. 1969;4(3):313\u201320. https:\/\/doi.org\/10.1099\/0022-1317-4-3-313.","journal-title":"J Gen Virol"},{"issue":"11","key":"3271_CR36","doi-asserted-by":"publisher","first-page":"4949","DOI":"10.1128\/jvi.74.11.4949-4956.2000","volume":"74","author":"T Jackson","year":"2000","unstructured":"Jackson T, Sheppard D, Denyer M, Blakemore W, King AM. The epithelial integrin \u03b1v\u03b26 is a receptor for foot-and-mouth disease virus. J Virol. 2000;74(11):4949\u201356. https:\/\/doi.org\/10.1128\/jvi.74.11.4949-4956.2000.","journal-title":"J Virol"},{"issue":"10","key":"3271_CR37","doi-asserted-by":"publisher","first-page":"2769","DOI":"10.1099\/vir.0.81172-0","volume":"86","author":"P Monaghan","year":"2005","unstructured":"Monaghan P, Gold S, Simpson J, Zhang Z, Weinreb PH, Violette SM, et al. The \u03b1v\u03b26 integrin receptor for foot-and-mouth disease virus is expressed constitutively on the epithelial cells targeted in cattle. J Gen Virol. 2005;86(10):2769\u201380. https:\/\/doi.org\/10.1099\/vir.0.81172-0.","journal-title":"J Gen Virol"},{"key":"3271_CR38","doi-asserted-by":"publisher","unstructured":"Curry S, Fry E, Blakemore W, Ghazaleh RA, Jackson T, King A, Lea S, Newman J, Stuart D, Logan D, Abu-Ghazaleh R, Blakemore W, Curry S, Jackson T, King A, Lea S, Lewis R, Newman J, Parry N, Rowlands D. Dissecting the roles of VP0 cleavage and RNA packaging in picornavirus capsid stabilization: the structure of empty capsids of foot-and-mouth disease virus. J VirolNature. 19971993;71(12):9743\u20139752. https:\/\/doi.org\/10.1128\/jvi.71.12.9743-9752.1997","DOI":"10.1128\/jvi.71.12.9743-9752.1997"},{"key":"3271_CR39","doi-asserted-by":"publisher","unstructured":"Logan D, Abu-Ghazaleh R, Blakemore W, Curry S, Jackson T, King A, Lea S, Lewis R, Newman J, Parry N, Rowlands D. Structure of a major immunogenic site on foot-and mouth disease virus. Nature. 1993;362(6420): 566-568.https:\/\/doi.org\/10.1038\/362566a0","DOI":"10.1038\/362566a0"},{"issue":"3","key":"3271_CR40","doi-asserted-by":"publisher","first-page":"369","DOI":"10.1099\/0022-1317-19-3-369","volume":"19","author":"P Talbot","year":"1973","unstructured":"Talbot P, Rowlands DJ, Burroughs JN, Sangar DV, Brown F. Evidence for a group protein in foot-and-mouth disease virus particles. J Gen Virol. 1973;19(3):369\u201380. https:\/\/doi.org\/10.1099\/0022-1317-19-3-369.","journal-title":"J Gen Virol"},{"issue":"45","key":"3271_CR41","doi-asserted-by":"publisher","first-page":"47212","DOI":"10.1074\/jbc.M405465200","volume":"279","author":"C Ferrer-Orta","year":"2004","unstructured":"Ferrer-Orta C, Arias A, Perez-Luque R, Escarm\u0131\u00b4s C, Domingo E, Verdaguer N. Structure of Foot-and-Mouth Disease Virus RNA-dependent RNA Polymerase and Its Complex with a Template-Primer RNA. JBC. 2004;279(45):47212\u201321. https:\/\/doi.org\/10.1074\/jbc.M405465200.","journal-title":"JBC"},{"issue":"22","key":"3271_CR42","doi-asserted-by":"publisher","first-page":"9463","DOI":"10.1073\/pnas.0700518104","volume":"104","author":"C Ferrer-Orta","year":"2007","unstructured":"Ferrer-Orta C, Arias A, Perez-Luque R, Escarm\u0131\u00b4s C, Domingo E, Verdaguer N. Sequential structures provide insights into the fidelity of RNA replication. PNAS. 2007;104(22):9463\u20138. https:\/\/doi.org\/10.1073\/pnas.0700518104.","journal-title":"PNAS"},{"issue":"6","key":"3271_CR43","doi-asserted-by":"publisher","first-page":"446","DOI":"10.1007\/s13238-021-00828-9","volume":"13","author":"H Dong","year":"2022","unstructured":"Dong H, Liu P, Bai M, Wang K, Feng R, Zhu D, Sun Y, Mu S, Li H, Harmsen M, Sun S, Wang X, Guo H. Structural and molecular basis for foot-and-mouth disease virus neutralization by two potent protective antibodies. Protein Cell. 2022;13(6):446\u201353. https:\/\/doi.org\/10.1007\/s13238-021-00828-9.","journal-title":"Protein Cell"},{"issue":"2","key":"3271_CR44","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1016\/j.jmb.2009.10.048","volume":"395","author":"PA Zunszain","year":"2010","unstructured":"Zunszain PA, Knox SR, Sweeney TR, Yang J, Roqu\u00e9-Rosell N, Belsham GJ, Leatherbarrow RJ, Curry S. Insights into Cleavage Specificity from the Crystal Structure of Foot-and-Mouth Disease Virus 3C Protease Complexed with a Peptide Substrate. J Mol Biol. 2010;395(2):375\u201389. . https:\/\/doi.org\/10.1016\/j.jmb.2009.10.048","journal-title":"J Mol Biol"},{"issue":"5","key":"3271_CR45","doi-asserted-by":"publisher","first-page":"1111","DOI":"10.1007\/s13204-018-0758-0","volume":"8","author":"S Jardon","year":"2018","unstructured":"Jardon S, Garc\u00eda CG, Quintanar D, Nieto JL, Ju\u00e1rez ML, Mendoza SE. Effect of two glycyrrhizinic acid nanoparticle carriers on MARC-145 cells actin filaments. Appl Nanosci. 2018;8(5):1111\u201321. https:\/\/doi.org\/10.1007\/s13204-018-0758-0.","journal-title":"Appl Nanosci"},{"key":"3271_CR46","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijpx.2025.100340","volume":"9","author":"H Li","year":"2025","unstructured":"Li H, Lin B, Wang S, Zhang Y, Zhou Z, Wen D, et al. Glycyrrhizic acid and its carrier-free micellar formulation: unraveling the potential for enhanced oral prevention of hearing loss. Int J Pharm X. 2025;9:100340. https:\/\/doi.org\/10.1016\/j.ijpx.2025.100340.","journal-title":"Int J Pharm X"},{"issue":"Pt 1","key":"3271_CR47","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2025.145623","volume":"320","author":"ASA Mohamed","year":"2025","unstructured":"Mohamed ASA, Abdelaziz AA, Abo-Kamar AM, Al-Madboly LA, El-Maradny YA, El-Fakharany EM. Multifunctional efficacy of the fabricated chitosan-coated silver nanoparticles as an antiviral agent against SARS-CoV-2: potent and safe mechanistic insights. Int J Biol Macromol. 2025;320(Pt 1):145623. https:\/\/doi.org\/10.1016\/j.ijbiomac.2025.145623.","journal-title":"Int J Biol Macromol"},{"issue":"107870","key":"3271_CR48","doi-asserted-by":"publisher","first-page":"0882","DOI":"10.1016\/j.micpath.2025.107870","volume":"207","author":"G Gopalakrishnan","year":"2025","unstructured":"Gopalakrishnan G, Deepa PM, Rajasekhar R, Vergis J, Bipin KC, Rathish RL, Janus A. In vitro antiviral activity of chitosan and curcumin-loaded chitosan nanoparticles against virulent Newcastle disease virus isolates. Microb Pathog. 2025;207(107870):0882\u20134010. . https:\/\/doi.org\/10.1016\/j.micpath.2025.107870","journal-title":"Microb Pathog"}],"container-title":["Virology Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12985-026-03271-6","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12985-026-03271-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12985-026-03271-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T14:07:05Z","timestamp":1786716425000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12985-026-03271-6"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,14]]},"references-count":48,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["3271"],"URL":"https:\/\/doi.org\/10.1186\/s12985-026-03271-6","relation":{},"ISSN":["1743-422X"],"issn-type":[{"value":"1743-422X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,14]]},"assertion":[{"value":"25 May 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 August 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 August 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"This work was confirmed by the Animal Ethics Committee of the Veterinary Serum and Vaccine Research Institute (VSVRI), and Research Ethics Committee of Ain Shams University, Microbiology Department with number: ASU-SCI\/MICR\/2024\/5\/4.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"197"}}