{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T00:05:43Z","timestamp":1788998743193,"version":"build-2803163510"},"reference-count":39,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,8,16]],"date-time":"2025-08-16T00:00:00Z","timestamp":1755302400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,8,16]],"date-time":"2025-08-16T00:00:00Z","timestamp":1755302400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100003509","name":"\u00d6rebro University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100003509","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Microbiol"],"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:sec>\n            <jats:title>Background<\/jats:title>\n            <jats:p>\n              <jats:italic>Neisseria meningitidis<\/jats:italic> is a human pathogen, carried asymptomatically in the nasopharynx, that can also cause invasive meningococcal disease. Understanding the carriage\/invasiveness balance is crucial, and bacterial genetic factors may impact this balance. A previous genome-wide association study reported that the gene <jats:italic>porB<\/jats:italic> class 3 was significantly associated with carriage isolates. This study aimed to examine the impact of <jats:italic>porB<\/jats:italic> variants on virulence in carriage and invasive meningococcal isolates.<\/jats:p>\n          <\/jats:sec>\n          <jats:sec>\n            <jats:title>Results<\/jats:title>\n            <jats:p>For this, 24 isolates were used (13 invasive and 11 carriage) belonging to different genogroups (B, C, W, Y, and cnl) and selected based on the presence of the genetic variant <jats:italic>porB<\/jats:italic> class 2 or class 3. Transgenic BALB\/c mice expressing human transferrin were infected intraperitoneally with these isolates. After 3 and 24\u00a0h of infection, clinical scores (fur quality, strength, and temperature) and bacterial load in blood were used to evaluate bacterial virulence. The concentrations of inflammatory cytokines were determined from blood. PorB<jats:sup>-<\/jats:sup> mutants were created from a carriage and an invasive isolate, and were tested in transgenic mice. The invasive isolates provoked significantly more severe infections compared to the carriage isolates, and the carriage isolates of <jats:italic>porB<\/jats:italic> class 3 were significantly less virulent than the invasive isolates of <jats:italic>porB<\/jats:italic> class 2 or 3. The invasive PorB<jats:sup>-<\/jats:sup> isolate caused milder infections than the parental isolate.<\/jats:p>\n          <\/jats:sec>\n          <jats:sec>\n            <jats:title>Conclusions<\/jats:title>\n            <jats:p>This study confirms the ability of invasive isolates of <jats:italic>N. meningitidis<\/jats:italic> to cause more severe infections than carriage isolates in transgenic mice. The <jats:italic>porB<\/jats:italic> expressed in invasive isolates seems to contribute to their higher virulence compared to carriage isolates, although this effect may depend on the genomic context. Notably, differences in virulence were mainly observed among serogroup C and W isolates.<\/jats:p>\n          <\/jats:sec>","DOI":"10.1186\/s12866-025-04246-3","type":"journal-article","created":{"date-parts":[[2025,8,16]],"date-time":"2025-08-16T09:34:16Z","timestamp":1755336856000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Inactivation of the porB gene reduces the virulence of Neisseria meningitidis in transgenic mice"],"prefix":"10.1186","volume":"25","author":[{"given":"Cecilia","family":"Klanger","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ala-Eddine","family":"Deghmane","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lorraine","family":"Eriksson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Olof","family":"S\u00e4ll","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sara Thulin","family":"Hedberg","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paula","family":"M\u00f6lling","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Muhamed-Kheir","family":"Taha","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,8,16]]},"reference":[{"key":"4246_CR1","doi-asserted-by":"publisher","first-page":"1378","DOI":"10.1056\/NEJM200105033441807","volume":"344","author":"NE Rosenstein","year":"2001","unstructured":"Rosenstein NE, Perkins BA, Stephens DS, Popovic T, Hughes JM. Meningococcal disease. N Engl J Med. 2001;344:1378\u201388.","journal-title":"N Engl J Med"},{"key":"4246_CR2","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/978-1-61779-346-2_1","volume":"799","author":"NG Rouphael","year":"2012","unstructured":"Rouphael NG, Stephens DS. Neisseria meningitidis: biology, microbiology, and epidemiology. Methods Mol Biol. 2012;799:1\u201320.","journal-title":"Methods Mol Biol"},{"key":"4246_CR3","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1016\/j.jinf.2020.05.079","volume":"81","author":"SR Parikh","year":"2020","unstructured":"Parikh SR, Campbell H, Bettinger JA, Harrison LH, Marshall HS, Martinon-Torres F, et al. The everchanging epidemiology of meningococcal disease worldwide and the potential for prevention through vaccination. J Infect. 2020;81:483\u201398.","journal-title":"J Infect"},{"key":"4246_CR4","doi-asserted-by":"publisher","first-page":"148","DOI":"10.1186\/s12879-022-07129-4","volume":"22","author":"SS Voss","year":"2022","unstructured":"Voss SS, Nielsen J, Valentiner-Branth P. Risk of sequelae after invasive meningococcal disease. BMC Infect Dis. 2022;22:148.","journal-title":"BMC Infect Dis"},{"key":"4246_CR5","doi-asserted-by":"publisher","first-page":"B64","DOI":"10.1016\/j.vaccine.2009.04.061","volume":"27","author":"DA Caugant","year":"2009","unstructured":"Caugant DA, Maiden MCJ. Meningococcal carriage and disease\u2014population biology and evolution. Vaccine. 2009;27:B64\u201370.","journal-title":"Vaccine"},{"key":"4246_CR6","doi-asserted-by":"publisher","first-page":"e80","DOI":"10.1017\/S0950268820000734","volume":"148","author":"SV Watle","year":"2020","unstructured":"Watle SV, Caugant DA, Tunheim G, Bekkevold T, Laake I, Brynildsrud OB, et al. Meningococcal carriage in Norwegian teenagers: strain characterisation and assessment of risk factors. Epidemiol Infect. 2020;148:e80.","journal-title":"Epidemiol Infect"},{"key":"4246_CR7","doi-asserted-by":"publisher","first-page":"3140","DOI":"10.1073\/pnas.95.6.3140","volume":"95","author":"MC Maiden","year":"1998","unstructured":"Maiden MC, Bygraves JA, Feil E, Morelli G, Russell JE, Urwin R, et al. Multilocus sequence typing: a portable approach to the identification of clones within populations of pathogenic microorganisms. Proc Natl Acad Sci U S A. 1998;95:3140\u20135.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"4246_CR8","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1111\/j.1469-0691.2008.01955.x","volume":"14","author":"ML Zarantonelli","year":"2008","unstructured":"Zarantonelli ML, Lancellotti M, Deghmane AE, Giorgini D, Hong E, Ruckly C, et al. Hyperinvasive genotypes of Neisseria meningitidis in France. Clin Microbiol Infect. 2008;14:467\u201372.","journal-title":"Clin Microbiol Infect"},{"key":"4246_CR9","doi-asserted-by":"publisher","first-page":"558","DOI":"10.1016\/j.meegid.2008.04.002","volume":"8","author":"DA Caugant","year":"2008","unstructured":"Caugant DA. Genetics and evolution of Neisseria meningitidis: importance for the epidemiology of meningococcal disease. Infect Genet Evol. 2008;8:558\u201365.","journal-title":"Infect Genet Evol"},{"key":"4246_CR10","doi-asserted-by":"publisher","first-page":"977","DOI":"10.1016\/j.jaci.2016.04.062","volume":"139","author":"J-F Jusot","year":"2017","unstructured":"Jusot J-F, Neill DR, Waters EM, Bangert M, Collins M, Moreno LB, et al. Airborne dust and high temperatures are risk factors for invasive bacterial disease. J Allergy Clin Immunol. 2017;139:977.","journal-title":"J Allergy Clin Immunol"},{"key":"4246_CR11","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1007\/s11910-015-0524-6","volume":"15","author":"R Dwilow","year":"2015","unstructured":"Dwilow R, Fanella S. Invasive meningococcal disease in the 21st century\u2014an update for the clinician. Curr Neurol Neurosci Rep. 2015;15:2.","journal-title":"Curr Neurol Neurosci Rep"},{"key":"4246_CR12","doi-asserted-by":"publisher","first-page":"1858","DOI":"10.1080\/21645515.2020.1849518","volume":"17","author":"M-K Taha","year":"2021","unstructured":"Taha M-K, Weil-Olivier C, Bou\u00e9e S, Emery C, Nachbaur G, Pribil C, et al. Risk factors for invasive meningococcal disease: a retrospective analysis of the French National public health insurance database. Hum Vaccines Immunother. 2021;17:1858\u201366.","journal-title":"Hum Vaccines Immunother"},{"key":"4246_CR13","doi-asserted-by":"publisher","first-page":"B71","DOI":"10.1016\/j.vaccine.2009.04.070","volume":"27","author":"DS Stephens","year":"2009","unstructured":"Stephens DS. Biology and pathogenesis of the evolutionarily successful, obligate human bacterium Neisseria meningitidis. Vaccine. 2009;27:B71-7.","journal-title":"Vaccine"},{"key":"4246_CR14","first-page":"759","volume":"42","author":"Y-L Tzeng","year":"2016","unstructured":"Tzeng Y-L, Thomas J, Stephens DS. Regulation of capsule in neisseria meningitidis. Crit Rev Microbiol. 2016;42:759\u201372.","journal-title":"Crit Rev Microbiol"},{"key":"4246_CR15","doi-asserted-by":"publisher","first-page":"001124","DOI":"10.1099\/mgen.0.001124","volume":"9","author":"L Eriksson","year":"2023","unstructured":"Eriksson L, Johannesen TB, Stenmark B, Jacobsson S, S\u00e4ll O, Hedberg ST, et al. Genetic variants linked to the phenotypic outcome of invasive disease and carriage of neisseria meningitidis. Microb Genomics. 2023;9:001124.","journal-title":"Microb Genomics"},{"key":"4246_CR16","doi-asserted-by":"publisher","first-page":"6811","DOI":"10.1073\/pnas.0912115107","volume":"107","author":"M Tanabe","year":"2010","unstructured":"Tanabe M, Nimigean CM, Iverson TM. Structural basis for solute transport, nucleotide regulation, and immunological recognition of Neisseria meningitidis PorB. Proc Natl Acad Sci U S A. 2010;107:6811\u20136.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"4246_CR17","first-page":"363","volume":"12","author":"NH Smith","year":"1995","unstructured":"Smith NH, Maynard Smith J, Spratt BG. Sequence evolution of the PorB gene of Neisseria gonorrhoeae and Neisseria meningitidis: evidence of positive darwinian selection. Mol Biol Evol. 1995;12:363\u201370.","journal-title":"Mol Biol Evol"},{"key":"4246_CR18","doi-asserted-by":"publisher","first-page":"5609","DOI":"10.1128\/IAI.00781-07","volume":"75","author":"M-L Zarantonelli","year":"2007","unstructured":"Zarantonelli M-L, Szatanik M, Giorgini D, Hong E, Huerre M, Guillou F, et al. Transgenic mice expressing human transferrin as a model for meningococcal infection. Infect Immun. 2007;75:5609\u201314.","journal-title":"Infect Immun"},{"key":"4246_CR19","doi-asserted-by":"publisher","first-page":"e22210","DOI":"10.1371\/journal.pone.0022210","volume":"6","author":"M Szatanik","year":"2011","unstructured":"Szatanik M, Hong E, Ruckly C, Ledroit M, Giorgini D, Jopek K, et al. Experimental meningococcal sepsis in congenic transgenic mice expressing human transferrin. PLoS One. 2011;6:e22210.","journal-title":"PLoS One"},{"key":"4246_CR20","doi-asserted-by":"publisher","first-page":"e25","DOI":"10.1017\/S0950268823000018","volume":"151","author":"S Olof","year":"2023","unstructured":"Olof S, Lorraine E, Asfaw Idosa B, Alexander P, Anders M, Sara TH, et al. Prevalence and persistence of neisseria meningitidis carriage in Swedish university students. Epidemiol Infect. 2023;151:e25.","journal-title":"Epidemiol Infect"},{"key":"4246_CR21","doi-asserted-by":"publisher","first-page":"1274","DOI":"10.1128\/jb.85.6.1274-1279.1963","volume":"85","author":"DS Kellogg","year":"1963","unstructured":"Kellogg DS, Peacock WL, Deacon WE, Brown L, Pirkle DI. Neisseria gonorrhoeae. i. Virulence genetically linked to clonal variation. J Bacteriol. 1963;85:1274\u20139.","journal-title":"J Bacteriol"},{"key":"4246_CR22","doi-asserted-by":"publisher","first-page":"e1000405","DOI":"10.1371\/journal.ppat.1000405","volume":"5","author":"A-E Deghmane","year":"2009","unstructured":"Deghmane A-E, Veckerl\u00e9 C, Giorgini D, Hong E, Ruckly C, Taha M-K. Differential modulation of TNF-\u03b1\u2013induced apoptosis by Neisseria meningitidis. PLoS Pathog. 2009;5:e1000405.","journal-title":"PLoS Pathog"},{"key":"4246_CR23","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1016\/j.steroids.2015.05.013","volume":"101","author":"M Levy","year":"2015","unstructured":"Levy M, Antunes A, Fiette L, Deghmane A-E, Taha M-K. Impact of corticosteroids on experimental meningococcal sepsis in mice. Steroids. 2015;101:96\u2013102.","journal-title":"Steroids"},{"key":"4246_CR24","volume-title":"AVMA guidelines for the euthanasia of animals: 2020 edition","author":"American Veterinary Medical Association","year":"2020","unstructured":"American Veterinary Medical Association, AVMA. AVMA guidelines for the euthanasia of animals: 2020 edition. Schaumburg, IL: AVMA; 2020."},{"key":"4246_CR25","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a016295","volume":"6","author":"T Tanaka","year":"2014","unstructured":"Tanaka T, Narazaki M, Kishimoto T. IL-6 in inflammation, immunity, and disease. Cold Spring Harb Perspect Biol. 2014;6: a016295.","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"4246_CR26","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1615\/CritRevEukarGeneExpr.v20.i2.10","volume":"20","author":"N Parameswaran","year":"2010","unstructured":"Parameswaran N, Patial S. Tumor necrosis factor-\u03b1 signaling in macrophages. Crit Rev Eukaryot Gene Expr. 2010;20:87\u2013103.","journal-title":"Crit Rev Eukaryot Gene Expr"},{"key":"4246_CR27","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1136\/adc.88.7.601","volume":"88","author":"N Pathan","year":"2003","unstructured":"Pathan N. Pathophysiology of meningococcal meningitis and septicaemia. Arch Dis Child. 2003;88:601\u20137.","journal-title":"Arch Dis Child"},{"key":"4246_CR28","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1084\/jem.169.1.333","volume":"169","author":"A Waage","year":"1989","unstructured":"Waage A, Brandtzaeg P, Halstensen A, Kierulf P, Espevik T. The complex pattern of cytokines in serum from patients with meningococcal septic shock. Association between interleukin 6, interleukin 1, and fatal outcome. J Exp Med. 1989;169:333\u20138.","journal-title":"J Exp Med"},{"key":"4246_CR29","doi-asserted-by":"publisher","first-page":"1797","DOI":"10.1084\/jem.171.5.1797","volume":"171","author":"B Moser","year":"1990","unstructured":"Moser B, Clark-Lewis I, Zwahlen R, Baggiolini M. Neutrophil-activating properties of the melanoma growth-stimulatory activity. J Exp Med. 1990;171:1797\u2013802.","journal-title":"J Exp Med"},{"key":"4246_CR30","doi-asserted-by":"publisher","first-page":"33123","DOI":"10.1038\/srep33123","volume":"6","author":"KV Sawant","year":"2016","unstructured":"Sawant KV, Poluri KM, Dutta AK, Sepuru KM, Troshkina A, Garofalo RP, et al. Chemokine CXCL1 mediated neutrophil recruitment: role of glycosaminoglycan interactions. Sci Rep. 2016;6:33123.","journal-title":"Sci Rep"},{"key":"4246_CR31","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1186\/s12866-020-01760-4","volume":"20","author":"L Eriksson","year":"2020","unstructured":"Eriksson L, Stenmark B, Deghmane A-E, Thulin Hedberg S, S\u00e4ll O, Fredlund H, et al. Difference in virulence between Neisseria meningitidis serogroups W and Y in transgenic mice. BMC Microbiol. 2020;20:92.","journal-title":"BMC Microbiol"},{"key":"4246_CR32","doi-asserted-by":"publisher","first-page":"e00339","DOI":"10.1128\/mBio.00339-13","volume":"4","author":"LA Lewis","year":"2013","unstructured":"Lewis LA, Vu DM, Vasudhev S, Shaughnessy J, Granoff DM, Ram S. Factor H-dependent alternative pathway inhibition mediated by porin B contributes to virulence of Neisseria meningitidis. mBio. 2013;4:e00339-13.","journal-title":"mBio"},{"key":"4246_CR33","doi-asserted-by":"publisher","first-page":"2574","DOI":"10.1128\/IAI.01517-14","volume":"82","author":"LA Lewis","year":"2014","unstructured":"Lewis LA, Vu DM, Granoff DM, Ram S. Inhibition of the alternative pathway of nonhuman infant complement by Porin B2 contributes to virulence of Neisseria meningitidis in the infant rat model. Infect Immun. 2014;82:2574\u201384.","journal-title":"Infect Immun"},{"key":"4246_CR34","doi-asserted-by":"publisher","first-page":"e1009992","DOI":"10.1371\/journal.ppat.1009992","volume":"17","author":"SG Earle","year":"2021","unstructured":"Earle SG, Lobanovska M, Lavender H, Tang C, Exley RM, Ramos-Sevillano E, et al. Genome-wide association studies reveal the role of polymorphisms affecting factor H binding protein expression in host invasion by Neisseria meningitidis. PLoS Pathog. 2021;17:e1009992.","journal-title":"PLoS Pathog"},{"key":"4246_CR35","doi-asserted-by":"publisher","unstructured":"Ren X, Eccles DA, Greig GA, Clapham J, Wheeler NE, Lindgreen S, et al. Genomic, transcriptomic, and phenotypic analyses of neisseria meningitidis isolates from disease patients and their household contacts. mSystems. 2017;2. https:\/\/doi.org\/10.1128\/msystems.00127.17.","DOI":"10.1128\/msystems.00127.17"},{"key":"4246_CR36","doi-asserted-by":"publisher","first-page":"5510","DOI":"10.1073\/pnas.1802298115","volume":"115","author":"OB Brynildsrud","year":"2018","unstructured":"Brynildsrud OB, Eldholm V, Bohlin J, Uadiale K, Obaro S, Caugant DA. Acquisition of virulence genes by a carrier strain gave rise to the ongoing epidemics of meningococcal disease in West Africa. Proc Natl Acad Sci U S A. 2018;115:5510\u20135.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"4246_CR37","first-page":"3287","volume":"191","author":"MG M\u00fcller","year":"2013","unstructured":"M\u00fcller MG, Ing JY, Cheng MK-W, Flitter BA, Moe GR. Identification of a phage-encoded Ig-binding protein from invasive Neisseria meningitidis. J Immunol Baltim Md 1950. 2013;191:3287\u201396.","journal-title":"J Immunol Baltim Md 1950"},{"key":"4246_CR38","doi-asserted-by":"publisher","first-page":"e3000410","DOI":"10.1371\/journal.pbio.3000410","volume":"18","author":"N Percie du Sert","year":"2020","unstructured":"Percie du Sert N, Hurst V, Ahluwalia A, Alam S, Avey MT, Baker M, et al. The ARRIVE guidelines 2.0: updated guidelines for reporting animal research. PLoS Biol. 2020;18:e3000410.","journal-title":"PLoS Biol"},{"key":"4246_CR39","unstructured":"PubMLST. Neisseria spp. https:\/\/pubmlst.org\/organisms\/neisseria-spp (accessed 5 Nov 2024)."}],"container-title":["BMC Microbiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-025-04246-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12866-025-04246-3\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-025-04246-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,9]],"date-time":"2025-09-09T14:59:19Z","timestamp":1757429959000},"score":1,"resource":{"primary":{"URL":"https:\/\/bmcmicrobiol.biomedcentral.com\/articles\/10.1186\/s12866-025-04246-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,16]]},"references-count":39,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["4246"],"URL":"https:\/\/doi.org\/10.1186\/s12866-025-04246-3","relation":{},"ISSN":["1471-2180"],"issn-type":[{"value":"1471-2180","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,16]]},"assertion":[{"value":"29 May 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 July 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 August 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Ethical permission for the meningococcal isolates collected during the carriage study [] had previously been given by the regional ethical review board in Uppsala, Sweden (reference number 2017\/499). The meningococcal isolates could not be linked to specific individuals.The study was carried out in strict accordance with the European Union Directive 2010\/63\/EU (and its revision 86\/609\/EEC) on the protection of animals used for scientific purposes and under the administrative authorization for animal experimentation (Permit Number 75-1554) and the approval of the Institut Pasteur Board (Dap210087\/N\u00b0APAFIS #34085-2021112212171298 v1). The protocol was approved by the Institut Pasteur Review Board, which is part of the Regional Ethical Committee for Animal Experiments of the Paris Region (Protocol 23.133_AC456). The ARRIVE guidelines were adhered to in this study [].","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"515"}}