{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T01:25:14Z","timestamp":1783560314170,"version":"3.55.0"},"reference-count":88,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,2,8]],"date-time":"2021-02-08T00:00:00Z","timestamp":1612742400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,2,8]],"date-time":"2021-02-08T00:00:00Z","timestamp":1612742400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100000209","name":"National Academies of Sciences, Engineering, and Medicine | National Academy of Sciences","doi-asserted-by":"publisher","award":["AID-263-A-15-00002"],"award-info":[{"award-number":["AID-263-A-15-00002"]}],"id":[{"id":"10.13039\/100000209","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007917","name":"United States Department of Agriculture | Agricultural Research Service","doi-asserted-by":"publisher","award":["59-8072-6-001"],"award-info":[{"award-number":["59-8072-6-001"]}],"id":[{"id":"10.13039\/100007917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007917","name":"United States Department of Agriculture | Agricultural Research Service","doi-asserted-by":"publisher","award":["59-8072-6-001"],"award-info":[{"award-number":["59-8072-6-001"]}],"id":[{"id":"10.13039\/100007917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007917","name":"United States Department of Agriculture | Agricultural Research Service","doi-asserted-by":"publisher","award":["59-8072-6-001"],"award-info":[{"award-number":["59-8072-6-001"]}],"id":[{"id":"10.13039\/100007917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007917","name":"United States Department of Agriculture | Agricultural Research Service","doi-asserted-by":"publisher","award":["59-8072-6-001"],"award-info":[{"award-number":["59-8072-6-001"]}],"id":[{"id":"10.13039\/100007917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100007917","name":"United States Department of Agriculture | Agricultural Research Service","doi-asserted-by":"publisher","award":["59-8072-6-001"],"award-info":[{"award-number":["59-8072-6-001"]}],"id":[{"id":"10.13039\/100007917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003712","name":"ISTK | Korea Food Research Institute","doi-asserted-by":"publisher","award":["E0142102-03"],"award-info":[{"award-number":["E0142102-03"]}],"id":[{"id":"10.13039\/501100003712","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000199","name":"U.S. Department of Agriculture","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100000199","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100009644","name":"National Academies of Sciences, Engineering, and Medicine | National Academy of Engineering","doi-asserted-by":"publisher","award":["AID-263-A-15-00002"],"award-info":[{"award-number":["AID-263-A-15-00002"]}],"id":[{"id":"10.13039\/100009644","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["npj Biofilms Microbiomes"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Environmental cues promote microbial biofilm formation and physiological and genetic heterogeneity. In food production facilities, biofilms produced by pathogens are a major source for food contamination; however, the pathogenesis of biofilm-isolated sessile cells is not well understood. We investigated the pathogenesis of sessile <jats:italic>Listeria monocytogenes<\/jats:italic> (<jats:italic>Lm<\/jats:italic>) using cell culture and mouse models. <jats:italic>Lm<\/jats:italic> sessile cells express reduced levels of the <jats:italic>lap<\/jats:italic>, <jats:italic>inlA, hly<\/jats:italic>, <jats:italic>prfA<\/jats:italic>, and <jats:italic>sigB<\/jats:italic> and show reduced adhesion, invasion, translocation, and cytotoxicity in the cell culture model than the planktonic cells. Oral challenge of C57BL\/6 mice with food, clinical, or murinized-InlA (InlA<jats:sup>m<\/jats:sup>) strains reveals that at 12 and 24\u2009h post-infection (hpi), <jats:italic>Lm<\/jats:italic> burdens are lower in tissues of mice infected with sessile cells than those infected with planktonic cells. However, these differences are negligible at 48 hpi. Besides, the expressions of <jats:italic>inlA<\/jats:italic> and <jats:italic>lap<\/jats:italic> mRNA in sessile <jats:italic>Lm<\/jats:italic> from intestinal content are about 6.0- and 280-fold higher than the sessle inoculum, respectively, suggesting sessile <jats:italic>Lm<\/jats:italic> can still upregulate virulence genes shortly after ingestion (12\u2009h). Similarly, exposure to simulated gastric fluid (SGF, pH 3) and intestinal fluid (SIF, pH 7) for 13\u2009h shows equal reduction in sessile and planktonic cell counts, but induces LAP and InlA expression and pathogenic phenotypes. Our data show that the virulence of biofilm-isolated <jats:italic>Lm<\/jats:italic> is temporarily attenuated and can be upregulated in mice during the early stage (12\u201324 hpi) but fully restored at a later stage (48 hpi) of infection. Our study further demonstrates that in vitro cell culture assay is unreliable; therefore, an animal model is essential for studying the pathogenesis of biofilm-isolated bacteria.<\/jats:p>","DOI":"10.1038\/s41522-021-00189-5","type":"journal-article","created":{"date-parts":[[2021,2,8]],"date-time":"2021-02-08T21:31:35Z","timestamp":1612819895000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":37,"title":["Biofilm-isolated Listeria monocytogenes exhibits reduced systemic dissemination at the early (12\u201324\u2009h) stage of infection in a mouse model"],"prefix":"10.1038","volume":"7","author":[{"given":"Xingjian","family":"Bai","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dongqi","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Luping","family":"Xu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shivendra","family":"Tenguria","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8405-2961","authenticated-orcid":false,"given":"Rishi","family":"Drolia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicholas L. F.","family":"Gallina","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2262-8168","authenticated-orcid":false,"given":"Abigail D.","family":"Cox","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ok-Kyung","family":"Koo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3640-1554","authenticated-orcid":false,"given":"Arun K.","family":"Bhunia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,2,8]]},"reference":[{"key":"189_CR1","doi-asserted-by":"publisher","first-page":"1236","DOI":"10.1016\/j.micinf.2007.05.011","volume":"9","author":"B Swaminathan","year":"2007","unstructured":"Swaminathan, B. & Gerner-Smidt, P. The epidemiology of human listeriosis. Microbes Infect. 9, 1236\u20131243 (2007).","journal-title":"Microbes Infect."},{"key":"189_CR2","doi-asserted-by":"publisher","first-page":"1973","DOI":"10.1084\/jem.20170495","volume":"214","author":"S Becattini","year":"2017","unstructured":"Becattini, S. et al. Commensal microbes provide first line defense against Listeria monocytogenes infection. J. Exp. Med. 214, 1973\u20131989 (2017).","journal-title":"J. Exp. Med."},{"key":"189_CR3","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1016\/j.chom.2018.03.004","volume":"23","author":"R Drolia","year":"2018","unstructured":"Drolia, R., Tenguria, S., Durkes, A. C., Turner, J. R. & Bhunia, A. K. Listeria adhesion protein induces intestinal epithelial barrier dysfunction for bacterial translocation. Cell Host Microbe 23, 470\u2013484 (2018).","journal-title":"Cell Host Microbe"},{"key":"189_CR4","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1016\/j.tim.2018.12.007","volume":"27","author":"R Drolia","year":"2019","unstructured":"Drolia, R. & Bhunia, A. K. Crossing the intestinal barrier via Listeria adhesion protein and internalin A. Trends Microbiol. 27, 408\u2013425 (2019).","journal-title":"Trends Microbiol."},{"key":"189_CR5","doi-asserted-by":"publisher","first-page":"931","DOI":"10.1128\/IAI.72.2.931-936.2004","volume":"72","author":"JL Wampler","year":"2004","unstructured":"Wampler, J. L., Kim, K. P., Jaradat, Z. & Bhunia, A. K. Heat shock protein 60 acts as a receptor for the Listeria adhesion protein in Caco-2 cells. Infect. Immun. 72, 931\u2013936 (2004).","journal-title":"Infect. Immun."},{"key":"189_CR6","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0020694","volume":"6","author":"B Jagadeesan","year":"2011","unstructured":"Jagadeesan, B. et al. N-Terminal Gly224-Gly411 domain in Listeria adhesion protein interacts with host receptor Hsp60. PLoS ONE 6, e20694 (2011).","journal-title":"PLoS ONE"},{"key":"189_CR7","doi-asserted-by":"publisher","first-page":"324","DOI":"10.1111\/j.1574-6968.2006.00140.x","volume":"256","author":"KP Kim","year":"2006","unstructured":"Kim, K. P. et al. Adhesion characteristics of Listeria adhesion protein (LAP)-expressing Escherichia coli to Caco-2 cells and of recombinant LAP to eukaryotic receptor Hsp60 as examined in a surface plasmon resonance sensor. FEMS Microbiol. Lett. 256, 324\u2013332 (2006).","journal-title":"FEMS Microbiol. Lett."},{"key":"189_CR8","doi-asserted-by":"publisher","first-page":"6344","DOI":"10.1038\/s41467-020-20200-5","volume":"11","author":"R Drolia","year":"2020","unstructured":"Drolia, R. et al. Receptor-targeted engineered probiotics mitigate lethal Listeria infecion. Nat. Commun. 11, 6344 (2020).","journal-title":"Nat. Commun."},{"key":"189_CR9","doi-asserted-by":"publisher","first-page":"2263","DOI":"10.1084\/jem.20110560","volume":"208","author":"G Nikitas","year":"2011","unstructured":"Nikitas, G. et al. Transcytosis of Listeria monocytogenes across the intestinal barrier upon specific targeting of goblet cell accessible E-cadherin. J. Exp. Med. 208, 2263\u20132277 (2011).","journal-title":"J. Exp. Med."},{"key":"189_CR10","doi-asserted-by":"publisher","first-page":"891","DOI":"10.1016\/j.cell.2007.03.049","volume":"129","author":"T Wollert","year":"2007","unstructured":"Wollert, T. et al. Extending the host range of Listeria monocytogenes by rational protein design. Cell 129, 891\u2013902 (2007).","journal-title":"Cell"},{"key":"189_CR11","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1111\/j.1365-2958.1995.tb02297.x","volume":"16","author":"S Dramsi","year":"1995","unstructured":"Dramsi, S. et al. Entry of Listeria monocytogenes into hepatocytes requires expression of InlB, a surface protein of the internalin multigene family. Mol. Microbiol. 16, 251\u2013261 (1995).","journal-title":"Mol. Microbiol."},{"key":"189_CR12","doi-asserted-by":"publisher","first-page":"379","DOI":"10.4161\/viru.2.5.17703","volume":"2","author":"A Camejo","year":"2011","unstructured":"Camejo, A. et al. The arsenal of virulence factors deployed by Listeria monocytogenes to promote its cell infection cycle. Virulence 2, 379\u2013394 (2011).","journal-title":"Virulence"},{"key":"189_CR13","doi-asserted-by":"publisher","first-page":"1263","DOI":"10.1128\/iai.60.4.1263-1267.1992","volume":"60","author":"DA Portnoy","year":"1992","unstructured":"Portnoy, D. A., Chakraborty, T., Goebel, W. & Cossart, P. Molecular determinants of Listeria monocytogenes pathogenesis. Infect. Immun. 60, 1263\u20131267 (1992).","journal-title":"Infect. Immun."},{"key":"189_CR14","doi-asserted-by":"publisher","first-page":"17333","DOI":"10.1073\/pnas.1007765107","volume":"107","author":"E Gouin","year":"2010","unstructured":"Gouin, E. et al. The Listeria monocytogenes InlC protein interferes with innate immune responses by targeting the I kappa B kinase subunit IKK alpha. Proc. Nat. Acad. Sci. USA 107, 17333\u201317338 (2010).","journal-title":"Proc. Nat. Acad. Sci. USA"},{"key":"189_CR15","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1006734","volume":"13","author":"M Kortebi","year":"2017","unstructured":"Kortebi, M. et al. Listeria monocytogenes switches from dissemination to persistence by adopting a vacuolar lifestyle in epithelial cells. PLoS Pathog. 13, e1006734 (2017).","journal-title":"PLoS Pathog."},{"key":"189_CR16","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1016\/j.mib.2011.01.005","volume":"14","author":"A de las Heras","year":"2011","unstructured":"de las Heras, A., Cain, R. J., Bielecka, M. K. & Vazquez-Boland, J. A. Regulation of Listeria virulence: PrfA master and commander. Curr. Opin. Microbiol. 14, 118\u2013127 (2011).","journal-title":"Curr. Opin. Microbiol."},{"key":"189_CR17","doi-asserted-by":"publisher","first-page":"1827","DOI":"10.1099\/mic.0.28758-0","volume":"152","author":"MJ Kazmierczak","year":"2006","unstructured":"Kazmierczak, M. J., Wiedmann, M. & Boor, K. J. Contributions of Listeria monocytogenes {sigma}B and PrfA to expression of virulence and stress response genes during extra- and intracellular growth. Microbiology 152, 1827\u20131838 (2006).","journal-title":"Microbiology"},{"key":"189_CR18","doi-asserted-by":"publisher","first-page":"1962","DOI":"10.3389\/fmicb.2018.01962","volume":"9","author":"N Horn","year":"2018","unstructured":"Horn, N. & Bhunia, A. K. Food-associated stress primes foodborne pathogens for the gastrointestinal phase of infection. Front. Microbiol. 9, 1962 (2018).","journal-title":"Front. Microbiol."},{"key":"189_CR19","doi-asserted-by":"crossref","unstructured":"Gali\u00e9, S., Garc\u00eda-Guti\u00e9rrez, C., Migu\u00e9lez, E. M., Villar, C. J. & Lomb\u00f3, F. Biofilms in the food industry: health aspects and control methods. Front. Microbiol. 9, 898 (2018).","DOI":"10.3389\/fmicb.2018.00898"},{"key":"189_CR20","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1111\/1751-7915.12186","volume":"7","author":"S Yaron","year":"2014","unstructured":"Yaron, S. & Romling, U. Biofilm formation by enteric pathogens and its role in plant colonization and persistence. Microb. Biotechnol. 7, 496\u2013516 (2014).","journal-title":"Microb. Biotechnol."},{"key":"189_CR21","doi-asserted-by":"publisher","first-page":"150","DOI":"10.4315\/0362-028X.JFP-13-150","volume":"77","author":"V Ferreira","year":"2014","unstructured":"Ferreira, V., Wiedmann, M., Teixeira, P. & Stasiewicz, M. J. Listeria monocytogenes persistence in food-associated environments: epidemiology, strain characteristics, and implications for public health. J. Food Prot. 77, 150\u2013170 (2014).","journal-title":"J. Food Prot."},{"key":"189_CR22","doi-asserted-by":"publisher","first-page":"92","DOI":"10.4315\/0362-028X-68.1.92","volume":"68","author":"AG Moltz","year":"2005","unstructured":"Moltz, A. G. & Martin, S. E. Formation of biofilms by Listeria monocytogenes under various growth conditions. J. Food Prot. 68, 92\u201397 (2005).","journal-title":"J. Food Prot."},{"key":"189_CR23","doi-asserted-by":"publisher","first-page":"85","DOI":"10.3390\/foods7060085","volume":"7","author":"P Rodr\u00edguez-L\u00f3pez","year":"2018","unstructured":"Rodr\u00edguez-L\u00f3pez, P., Rodr\u00edguez-Herrera, J., V\u00e1zquez-S\u00e1nchez, D. & Lopez Cabo, M. Current knowledge on Listeria monocytogenes biofilms in food-related environments: incidence, resistance to biocides, ecology and biocontrol. Foods 7, 85 (2018).","journal-title":"Foods"},{"key":"189_CR24","doi-asserted-by":"publisher","first-page":"623","DOI":"10.1038\/nrmicro2171","volume":"7","author":"NE Freitag","year":"2009","unstructured":"Freitag, N. E., Port, G. C. & Miner, M. D. Listeria monocytogenes from saprophyte to intracellular pathogen. Nat. Rev. Microbiol. 7, 623\u2013628 (2009).","journal-title":"Nat. Rev. Microbiol."},{"key":"189_CR25","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.ijfoodmicro.2011.01.005","volume":"145","author":"B Carpentier","year":"2011","unstructured":"Carpentier, B. & Cerf, O. Review\u2014Persistence of Listeria monocytogenes in food industry equipment and premises. Int. J. Food Microbiol. 145, 1\u20138 (2011).","journal-title":"Int. J. Food Microbiol."},{"key":"189_CR26","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1128\/AAC.00576-06","volume":"51","author":"LM Weigel","year":"2007","unstructured":"Weigel, L. M. et al. High-level vancomycin-resistant Staphylococcus aureus isolates associated with a polymicrobial biofilm. Antimicrob. Agents Chemother. 51, 231 (2007).","journal-title":"Antimicrob. Agents Chemother."},{"key":"189_CR27","doi-asserted-by":"publisher","first-page":"835","DOI":"10.1111\/j.1462-2920.2010.02378.x","volume":"13","author":"S Renier","year":"2011","unstructured":"Renier, S., H\u00e9braud, M. & Desvaux, M. Molecular biology of surface colonization by Listeria monocytogenes: an additional facet of an opportunistic Gram\u2010positive foodborne pathogen. Envviron. Microbiol. 13, 835\u2013850 (2011).","journal-title":"Envviron. Microbiol"},{"key":"189_CR28","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1038\/nrmicro821","volume":"2","author":"L Hall-Stoodley","year":"2004","unstructured":"Hall-Stoodley, L., Costerton, J. W. & Stoodley, P. Bacterial biofilms: from the natural environment to infectious diseases. Nat. Rev. Microbiol. 2, 95\u2013108 (2004).","journal-title":"Nat. Rev. Microbiol."},{"key":"189_CR29","doi-asserted-by":"publisher","first-page":"1318","DOI":"10.1126\/science.284.5418.1318","volume":"284","author":"JW Costerton","year":"1999","unstructured":"Costerton, J. W., Stewart, P. S. & Greenberg, E. P. Bacterial biofilms: a common cause of persistent infections. Science 284, 1318\u20131322 (1999).","journal-title":"Science"},{"key":"189_CR30","doi-asserted-by":"publisher","first-page":"881","DOI":"10.3201\/eid0809.020063","volume":"8","author":"RM Donlan","year":"2002","unstructured":"Donlan, R. M. Biofilms: microbial life on surfaces. Emerg. Infect. Dis. 8, 881 (2002).","journal-title":"Emerg. Infect. Dis."},{"key":"189_CR31","doi-asserted-by":"publisher","first-page":"2271","DOI":"10.1128\/AEM.02361-09","volume":"76","author":"M Harmsen","year":"2010","unstructured":"Harmsen, M., Lappann, M., Kn\u00f8chel, S. & Molin, S. Role of extracellular DNA during biofilm formation by Listeria monocytogenes. Appl. Environ. Microbiol. 76, 2271\u20132279 (2010).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR32","first-page":"159","volume":"8","author":"C Kokare","year":"2009","unstructured":"Kokare, C., Chakraborty, S., Khopade, A. & Mahadik, K. R. Biofilm: importance and applications. Ind. J. Biotechnol. 8, 159\u2013168 (2009).","journal-title":"Ind. J. Biotechnol."},{"key":"189_CR33","doi-asserted-by":"publisher","first-page":"1481","DOI":"10.4315\/0362-028X.JFP-17-466","volume":"81","author":"L Luque-Sastre","year":"2018","unstructured":"Luque-Sastre, L., Fox, E. M., Jordan, K. & Fanning, S. A comparative study of the susceptibility of Listeria species to sanitizer treatments when grown under planktonic and biofilm conditions. J. Food Prot. 81, 1481\u20131490 (2018).","journal-title":"J. Food Prot."},{"key":"189_CR34","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1016\/S0378-1097(02)00571-2","volume":"210","author":"F Tremoulet","year":"2002","unstructured":"Tremoulet, F. et al. Comparison of protein patterns of Listeria monocytogenes grown in biofilm or in planktonic mode by proteomic analysis. FEMS Microbiol. Lett. 210, 25\u201331 (2002).","journal-title":"FEMS Microbiol. Lett."},{"key":"189_CR35","doi-asserted-by":"publisher","first-page":"6075","DOI":"10.1128\/AEM.01592-13","volume":"79","author":"A Louren\u00e7o","year":"2013","unstructured":"Louren\u00e7o, A. et al. Comparison of Listeria monocytogenes exoproteomes from biofilm and planktonic state: Lmo2504, a protein associated with biofilms. Appl. Environ. Microbiol. 79, 6075\u20136082 (2013).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR36","doi-asserted-by":"publisher","first-page":"1161","DOI":"10.1021\/pr501114e","volume":"14","author":"MM Mata","year":"2015","unstructured":"Mata, M. M., da Silva, W. P., Wilson, R., Lowe, E. & Bowman, J. P. Attached and planktonic Listeria monocytogenes global proteomic responses and associated influence of strain genetics and temperature. J. Proteome Res. 14, 1161\u20131173 (2015).","journal-title":"J. Proteome Res."},{"key":"189_CR37","doi-asserted-by":"publisher","first-page":"50","DOI":"10.1016\/j.ijfoodmicro.2015.12.004","volume":"219","author":"N Gilmartin","year":"2016","unstructured":"Gilmartin, N., Gi\u00e3o, M. S., Keevil, C. W. & O\u2019Kennedy, R. Differential internalin A levels in biofilms of Listeria monocytogenes grown on different surfaces and nutrient conditions. Int. J. Food Microbiol. 219, 50\u201355 (2016).","journal-title":"Int. J. Food Microbiol."},{"key":"189_CR38","doi-asserted-by":"crossref","unstructured":"Bhunia, A. K. Animal and cell culture models to study foodborne pathogens. in Foodborne Microbial Pathogens 117\u2013132 (Springer, 2018).","DOI":"10.1007\/978-1-4939-7349-1_5"},{"key":"189_CR39","doi-asserted-by":"publisher","first-page":"7854","DOI":"10.1128\/AEM.01519-10","volume":"76","author":"S van der Veen","year":"2010","unstructured":"van der Veen, S. & Abee, T. Importance of SigB for Listeria monocytogenes static and continuous-flow biofilm formation and disinfectant resistance. Appl. Environ. Microbiol. 76, 7854\u20137860 (2010).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR40","doi-asserted-by":"publisher","first-page":"418","DOI":"10.1016\/j.fm.2010.09.014","volume":"28","author":"PS Ibusquiza","year":"2011","unstructured":"Ibusquiza, P. S., Herrera, J. J. R. & Cabo, M. L. Resistance to benzalkonium chloride, peracetic acid and nisin during formation of mature biofilms by Listeria monocytogenes. Food Microbiol. 28, 418\u2013425 (2011).","journal-title":"Food Microbiol."},{"key":"189_CR41","doi-asserted-by":"publisher","first-page":"2950","DOI":"10.1128\/AEM.68.6.2950-2958.2002","volume":"68","author":"D Djordjevic","year":"2002","unstructured":"Djordjevic, D., Wiedmann, M. & McLandsborough, L. A. Microtiter plate assay for assessment of Listeria monocytogenes biofilm formation. Appl. Environ. Microbiol. 68, 2950\u20132958 (2002).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR42","doi-asserted-by":"publisher","first-page":"3819","DOI":"10.1128\/JCM.42.8.3819-3822.2004","volume":"42","author":"M Doumith","year":"2004","unstructured":"Doumith, M., Buchrieser, C., Glaser, P., Jacquet, C. & Martin, P. Differentiation of the major Listeria monocytogenes serovars by multiplex PCR. J. Clin. Microbiol. 42, 3819\u20133822 (2004).","journal-title":"J. Clin. Microbiol."},{"key":"189_CR43","doi-asserted-by":"publisher","first-page":"1813","DOI":"10.1128\/AEM.03173-14","volume":"81","author":"M Guilbaud","year":"2015","unstructured":"Guilbaud, M., Piveteau, P., Desvaux, M., Brisse, S. & Briandet, R. Exploring the diversity of Listeria monocytogenes biofilm architecture by high-throughput confocal laser scanning microscopy and the predominance of the honeycomb-like morphotype. Appl. Environ. Microbiol. 81, 1813\u20131819 (2015).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR44","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1006\/mpat.1994.1011","volume":"16","author":"A Bhunia","year":"1994","unstructured":"Bhunia, A. et al. A six-hour in vitro virulence assay for Listeria monocytogenes using myeloma and hybridoma cells from murine and human sources. Microb. Pathog. 16, 99\u2013110 (1994).","journal-title":"Microb. Pathog."},{"key":"189_CR45","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1016\/j.bios.2010.05.020","volume":"26","author":"P Banerjee","year":"2010","unstructured":"Banerjee, P. & Bhunia, A. K. Cell-based biosensor for rapid screening of pathogens and toxins. Biosens. Bioelectron. 26, 99\u2013106 (2010).","journal-title":"Biosens. Bioelectron."},{"key":"189_CR46","first-page":"398","volume":"9","author":"AK Bhunia","year":"1999","unstructured":"Bhunia, A. K. & Feng, X. Examination of cytopathic effect and apoptosis in Listeria monocytogenes-infected hybridoma B-lymphocyte (Ped-2E9) line in vitro. J. Microbiol. Biotechnol. 9, 398\u2013403 (1999).","journal-title":"J. Microbiol. Biotechnol."},{"key":"189_CR47","doi-asserted-by":"publisher","first-page":"5865","DOI":"10.1128\/JCM.43.12.5865-5872.2005","volume":"43","author":"KM Gray","year":"2005","unstructured":"Gray, K. M., Banada, P. P., O\u2019Neal, E. & Bhunia, A. K. Rapid Ped-2E9 cell-based cytotoxicity analysis and genotyping of Bacillus species. J. Clin. Microbiol. 43, 5865\u20135872 (2005).","journal-title":"J. Clin. Microbiol."},{"key":"189_CR48","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1080\/09603120903281283","volume":"20","author":"VF Bueno","year":"2010","unstructured":"Bueno, V. F. et al. Characterization of Listeria monocytogenes isolates of food and human origins from Brazil using molecular typing procedures and in vitro cell culture assays. Int. J. Environ. Health Res. 20, 43\u201359 (2010).","journal-title":"Int. J. Environ. Health Res."},{"key":"189_CR49","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1016\/S0147-9571(02)00039-5","volume":"26","author":"A Menon","year":"2003","unstructured":"Menon, A., Shroyer, M. L., Wampler, J. L., Chawan, C. B. & Bhunia, A. K. In vitro study of Listeria monocytogenes infection to murine primary and human transformed B cells. Comp. Immunol. Microbiol. Infect. Dis. 26, 157\u2013174 (2003).","journal-title":"Comp. Immunol. Microbiol. Infect. Dis."},{"key":"189_CR50","doi-asserted-by":"crossref","unstructured":"Louie, A., Zhang, T., Becattini, S., Waldor, M. K. & Portnoy, D. A. A multiorgan trafficking circuit provides purifying selection of Listeria monocytogenes virulence genes. mBio 10, e02948 (2019).","DOI":"10.1128\/mBio.02948-19"},{"key":"189_CR51","doi-asserted-by":"publisher","first-page":"198","DOI":"10.4315\/0362-028X-53.3.198","volume":"53","author":"BR Charlton","year":"1990","unstructured":"Charlton, B. R., Kinde, H. & Jensen, L. H. Environmental survey for Listeria species in California milk processing plants. J. Food Prot. 53, 198\u2013201 (1990).","journal-title":"J. Food Prot."},{"key":"189_CR52","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1038\/nrmicro1838","volume":"6","author":"PS Stewart","year":"2008","unstructured":"Stewart, P. S. & Franklin, M. J. Physiological heterogeneity in biofilms. Nat. Rev. Microbiol. 6, 199 (2008).","journal-title":"Nat. Rev. Microbiol."},{"key":"189_CR53","doi-asserted-by":"publisher","first-page":"372","DOI":"10.1007\/s00284-013-0377-7","volume":"67","author":"Q Luo","year":"2013","unstructured":"Luo, Q. et al. PrfA led to reduced biofilm formation and contributed to altered gene expression patterns in biofilm-forming Listeria monocytogenes. Curr. Microbiol. 67, 372\u2013378 (2013).","journal-title":"Curr. Microbiol."},{"key":"189_CR54","doi-asserted-by":"crossref","unstructured":"Barbosa, J. et al. Biofilm formation among clinical and food isolates of Listeria monocytogenes. Int. J. Microbiol. 2013, 524975 (2013).","DOI":"10.1155\/2013\/524975"},{"key":"189_CR55","doi-asserted-by":"publisher","first-page":"7336","DOI":"10.1128\/AEM.69.12.7336-7342.2003","volume":"69","author":"MK Borucki","year":"2003","unstructured":"Borucki, M. K., Peppin, J. D., White, D., Loge, F. & Call, D. R. Variation in biofilm formation among strains of Listeria monocytogenes. Appl. Environ. Microbiol. 69, 7336\u20137342 (2003).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR56","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/j.ijfoodmicro.2004.03.031","volume":"97","author":"B Carpentier","year":"2004","unstructured":"Carpentier, B. & Chassaing, D. Interactions in biofilms between Listeria monocytogenes and resident microorganisms from food industry premises. Int. J. Food Microbiol. 97, 111\u2013122 (2004).","journal-title":"Int. J. Food Microbiol"},{"key":"189_CR57","doi-asserted-by":"publisher","first-page":"5846","DOI":"10.1128\/AEM.00816-09","volume":"75","author":"Y Pan","year":"2009","unstructured":"Pan, Y., Breidt, F. & Kathariou, S. Competition of Listeria monocytogenes serotype 1\/2a and 4b strains in mixed-culture biofilms. Appl. Environ. Microbiol. 75, 5846\u20135852 (2009).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR58","doi-asserted-by":"publisher","first-page":"909","DOI":"10.1111\/mmi.12140","volume":"87","author":"T Tiensuu","year":"2013","unstructured":"Tiensuu, T., Andersson, C., Ryd\u00e9n, P. & Johansson, J. Cycles of light and dark co\u2010ordinate reversible colony differentiation in Listeria monocytogenes. Mol. Microbiol. 87, 909\u2013924 (2013).","journal-title":"Mol. Microbiol."},{"key":"189_CR59","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1003131","volume":"9","author":"L Travier","year":"2013","unstructured":"Travier, L. et al. ActA promotes Listeria monocytogenes aggregation, intestinal colonization and carriage. PLoS Pathog. 9, e1003131 (2013).","journal-title":"PLoS Pathog."},{"key":"189_CR60","doi-asserted-by":"publisher","first-page":"2919","DOI":"10.1128\/AEM.02664-06","volume":"73","author":"P McGann","year":"2007","unstructured":"McGann, P., Wiedmann, M. & Boor, K. J. The alternative sigma factor {sigma}B and the virulence gene regulator PrfA both regulate transcription of Listeria monocytogenes internalins. Appl. Environ. Microbiol. 73, 2919\u20132930 (2007).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR61","doi-asserted-by":"publisher","first-page":"3969","DOI":"10.1128\/JB.00179-10","volume":"192","author":"KP Lemon","year":"2010","unstructured":"Lemon, K. P., Freitag, N. E. & Kolter, R. The virulence regulator PrfA promotes biofilm formation by Listeria monocytogenes. J. Bacteriol. 192, 3969\u20133976 (2010).","journal-title":"J. Bacteriol."},{"key":"189_CR62","doi-asserted-by":"publisher","first-page":"1196","DOI":"10.1016\/j.micinf.2007.05.007","volume":"9","author":"M Scortti","year":"2007","unstructured":"Scortti, M., Monz\u00f3, H. J., Lacharme-Lora, L., Lewis, D. A. & V\u00e1zquez-Boland, J. A. The PrfA virulence regulon. Microbes Infect. 9, 1196\u20131207 (2007).","journal-title":"Microbes Infect."},{"key":"189_CR63","doi-asserted-by":"publisher","first-page":"3896","DOI":"10.1128\/iai.63.10.3896-3903.1995","volume":"63","author":"A Lingnau","year":"1995","unstructured":"Lingnau, A. et al. Expression of the Listeria monocytogenes EGD inlA and inlB genes, whose products mediate bacterial entry into tissue culture cell lines, by PrfA-dependent and PrfA-independent mechanisms. Infect. Immun. 63, 3896\u20133903 (1995).","journal-title":"Infect. Immun."},{"key":"189_CR64","doi-asserted-by":"publisher","DOI":"10.1186\/s13099-015-0053-5","volume":"7","author":"MG Mathipa","year":"2015","unstructured":"Mathipa, M. G. & Thantsha, M. S. Cocktails of probiotics pre-adapted to multiple stress factors are more robust under simulated gastrointestinal conditions than their parental counterparts and exhibit enhanced antagonistic capabilities against Escherichia coli and Staphylococcus aureus. Gut Pathog. 7, 5 (2015).","journal-title":"Gut Pathog."},{"key":"189_CR65","doi-asserted-by":"publisher","first-page":"950","DOI":"10.1038\/nature08080","volume":"459","author":"A Toledo-Arana","year":"2009","unstructured":"Toledo-Arana, A. et al. The Listeria transcriptional landscape from saprophytism to virulence. Nature 459, 950\u2013956 (2009).","journal-title":"Nature"},{"key":"189_CR66","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1016\/j.micinf.2009.05.006","volume":"11","author":"KM Burkholder","year":"2009","unstructured":"Burkholder, K. M. et al. Expression of LAP, a SecA2-dependent secretory protein, is induced under anaerobic environment. Microbes Infect. 11, 859\u2013867 (2009).","journal-title":"Microbes Infect."},{"key":"189_CR67","doi-asserted-by":"publisher","first-page":"4876","DOI":"10.1128\/AEM.68.10.4876-4883.2002","volume":"68","author":"ZW Jaradat","year":"2002","unstructured":"Jaradat, Z. W. & Bhunia, A. K. Glucose and nutrient concentrations affect the expression of a 104-kilodalton Listeria adhesion protein in Listeria monocytogenes. Appl. Environ. Microbiol. 68, 4876\u20134883 (2002).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR68","doi-asserted-by":"publisher","first-page":"2765","DOI":"10.1128\/AEM.65.6.2765-2769.1999","volume":"65","author":"NI Santiago","year":"1999","unstructured":"Santiago, N. I., Zipf, A. & Bhunia, A. K. Influence of temperature and growth phase on expression of a 104-kilodalton Listeria adhesion protein in Listeria monocytogenes. Appl. Environ. Microbiol. 65, 2765\u20132769 (1999).","journal-title":"Appl. Environ. Microbiol."},{"key":"189_CR69","doi-asserted-by":"publisher","first-page":"3843","DOI":"10.1099\/mic.0.27257-0","volume":"150","author":"D Sue","year":"2004","unstructured":"Sue, D., Fink, D., Wiedmann, M. & Boor, K. J. {sigma}B-dependent gene induction and expression in Listeria monocytogenes during osmotic and acid stress conditions simulating the intestinal environment. Microbiology 150, 3843\u20133855 (2004).","journal-title":"Microbiology"},{"key":"189_CR70","doi-asserted-by":"publisher","first-page":"3956","DOI":"10.1093\/emboj\/18.14.3956","volume":"18","author":"M Lecuit","year":"1999","unstructured":"Lecuit, M. et al. A single amino acid in E-cadherin responsible for host specificity towards the human pathogen Listeria monocytogenes. EMBO J. 18, 3956\u20133963 (1999).","journal-title":"EMBO J."},{"key":"189_CR71","doi-asserted-by":"publisher","first-page":"318","DOI":"10.1186\/1471-2180-10-318","volume":"10","author":"IR Monk","year":"2010","unstructured":"Monk, I. R., Casey, P. G., Hill, C. & Gahan, C. G. M. Directed evolution and targeted mutagenesis to murinize Listeria monocytogenes internalin A for enhanced infectivity in the murine oral infection model. BMC Microbiol. 10, 318 (2010).","journal-title":"BMC Microbiol."},{"key":"189_CR72","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1017\/S0022172400022464","volume":"70","author":"DA van der Waaij","year":"1972","unstructured":"van der Waaij, D. A., Berghuis, J. & Lekkerkerk, J. Colonization resistance of the digestive tract of mice during systemic antibiotic treatment. Epidemiol. Infect. 70, 605\u2013610 (1972).","journal-title":"Epidemiol. Infect."},{"key":"189_CR73","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1128\/iai.47.1.118-122.1985","volume":"47","author":"DJ Hentges","year":"1985","unstructured":"Hentges, D. J., Stein, A. J., Casey, S. W. & Que, J. U. Protective role of intestinal flora against infection with Pseudomonas aeruginosa in mice: influence of antibiotics on colonization resistance. Infect. Immun. 47, 118\u2013122 (1985).","journal-title":"Infect. Immun."},{"key":"189_CR74","doi-asserted-by":"publisher","first-page":"1176","DOI":"10.1016\/j.micinf.2007.05.005","volume":"9","author":"P Schnupf","year":"2007","unstructured":"Schnupf, P. & Portnoy, D. A. Listeriolysin O: a phagosome-specific lysin. Microbes Infect. 9, 1176\u20131187 (2007).","journal-title":"Microbes Infect."},{"key":"189_CR75","doi-asserted-by":"publisher","first-page":"679","DOI":"10.4049\/jimmunol.156.2.679","volume":"156","author":"HW Rogers","year":"1996","unstructured":"Rogers, H. W., Callery, M. P., Deck, B. & Unanue, E. R. Listeria monocytogenes induces apoptosis of infected hepatocytes. J. Immunol. 156, 679\u2013684 (1996).","journal-title":"J. Immunol."},{"key":"189_CR76","doi-asserted-by":"publisher","first-page":"3147","DOI":"10.1128\/iai.58.9.3147-3150.1990","volume":"58","author":"J Roll","year":"1990","unstructured":"Roll, J. & Czuprynski, C. Hemolysin is required for extraintestinal dissemination of Listeria monocytogenes in intragastrically inoculated mice. Infect. Immun. 58, 3147\u20133150 (1990).","journal-title":"Infect. Immun."},{"key":"189_CR77","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1016\/j.foodcont.2011.09.012","volume":"24","author":"M Arevalos-S\u00e1nchez","year":"2012","unstructured":"Arevalos-S\u00e1nchez, M., Regalado, C., Martin, S. E., Dom\u00ednguez-Dom\u00ednguez, J. & Garc\u00eda-Almend\u00e1rez, B. E. Effect of neutral electrolyzed water and nisin on Listeria monocytogenes biofilms, and on listeriolysin O activity. Food Control 24, 116\u2013122 (2012).","journal-title":"Food Control"},{"key":"189_CR78","doi-asserted-by":"publisher","DOI":"10.1038\/s41522-019-0112-7","volume":"6","author":"J Caro-Astorga","year":"2020","unstructured":"Caro-Astorga, J. et al. Biofilm formation displays intrinsic offensive and defensive features of Bacillus cereus. npj Biofilms Microbiomes 6, 3 (2020).","journal-title":"npj Biofilms Microbiomes"},{"key":"189_CR79","doi-asserted-by":"publisher","first-page":"2782","DOI":"10.1099\/mic.0.036509-0","volume":"156","author":"B Jagadeesan","year":"2010","unstructured":"Jagadeesan, B. et al. LAP, an alcohol acetaldehyde dehydrogenase enzyme in Listeria promotes bacterial adhesion to enterocyte-like Caco-2 cells only in pathogenic species. Microbiology 156, 2782\u20132795 (2010).","journal-title":"Microbiology"},{"key":"189_CR80","doi-asserted-by":"publisher","first-page":"e01324","DOI":"10.1128\/genomeA.01324-17","volume":"5","author":"TW Bailey","year":"2017","unstructured":"Bailey, T. W., do Nascimento, N. C. & Bhunia, A. K. Genome sequence of Listeria monocytogenes strain F4244, a 4b serotype. Genome Announc. 5, e01324\u201301317 (2017).","journal-title":"Genome Announc."},{"key":"189_CR81","doi-asserted-by":"crossref","unstructured":"Rychli, K., Guinane, C. M., Daly, K., Hill, C. & Cotter, P. D. G. Generation of nonpolar deletion mutants in Kieran Jordan, Edward M. Fox, and Martin Wagner (Eds) Listeria monocytogenes using the \u201cSOEing\u201d method. In Listeria monocytogenes 187\u2013200 (Springer, 2014).","DOI":"10.1007\/978-1-4939-0703-8_16"},{"key":"189_CR82","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.plasmid.2015.05.003","volume":"81","author":"H Abdelhamed","year":"2015","unstructured":"Abdelhamed, H., Lawrence, M. L. & Karsi, A. A novel suicide plasmid for efficient gene mutation in Listeria monocytogenes. Plasmid 81, 1\u20138 (2015).","journal-title":"Plasmid"},{"key":"189_CR83","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1186\/1471-2180-12-275","volume":"12","author":"M Mendonca","year":"2012","unstructured":"Mendonca, M. et al. Highly specific fiber optic immunosensor coupled with immunomagnetic separation for detection of low levels of Listeria monocytogenes and L. ivanovii. BMC Microbiol 12, 275 (2012).","journal-title":"BMC Microbiol"},{"key":"189_CR84","doi-asserted-by":"publisher","first-page":"5062","DOI":"10.1128\/IAI.00516-10","volume":"78","author":"KM Burkholder","year":"2010","unstructured":"Burkholder, K. M. & Bhunia, A. K. Listeria monocytogenes uses Listeria adhesion protein (LAP) to promote bacterial transepithelial translocation, and induces expression of LAP receptor Hsp60. Infect. Immun. 78, 5062\u20135073 (2010).","journal-title":"Infect. Immun."},{"key":"189_CR85","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1016\/j.mimet.2005.10.017","volume":"66","author":"F Yamada","year":"2006","unstructured":"Yamada, F. et al. Invasion assay of Listeria monocytogenes using Vero and Caco-2 cells. J. Microbiol. Methods 66, 96\u2013103 (2006).","journal-title":"J. Microbiol. Methods"},{"key":"189_CR86","doi-asserted-by":"publisher","first-page":"1375","DOI":"10.1016\/j.biopha.2017.09.051","volume":"95","author":"P Mannarreddy","year":"2017","unstructured":"Mannarreddy, P. et al. Cytotoxic effect of Cyperus rotundus rhizome extract on human cancer cell lines. Biomed. Pharmacother. 95, 1375\u20131387 (2017).","journal-title":"Biomed. Pharmacother."},{"key":"189_CR87","doi-asserted-by":"publisher","first-page":"74","DOI":"10.3389\/fsufs.2018.00074","volume":"2","author":"AK Singh","year":"2018","unstructured":"Singh, A. K., Bai, X., Amalaradjou, M. A. R. & Bhunia, A. K. Antilisterial and antibiofilm activities of pediocin and LAP functionalized gold nanoparticles. Front. Sustain. Food Syst. 2, 74 (2018).","journal-title":"Front. Sustain. Food Syst."},{"key":"189_CR88","doi-asserted-by":"publisher","first-page":"1356","DOI":"10.1128\/AEM.71.3.1356-1363.2005","volume":"71","author":"R Oozeer","year":"2005","unstructured":"Oozeer, R. et al. Differential activities of four Lactobacillus casei promoters during bacterial transit through the gastrointestinal tracts of human-microbiota-associated mice. Appl. Environ. Microbiol. 71, 1356\u20131363 (2005).","journal-title":"Appl. Environ. Microbiol."}],"container-title":["npj Biofilms and Microbiomes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41522-021-00189-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41522-021-00189-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41522-021-00189-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,1,28]],"date-time":"2023-01-28T23:08:07Z","timestamp":1674947287000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41522-021-00189-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,2,8]]},"references-count":88,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["189"],"URL":"https:\/\/doi.org\/10.1038\/s41522-021-00189-5","relation":{},"ISSN":["2055-5008"],"issn-type":[{"value":"2055-5008","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,2,8]]},"assertion":[{"value":"3 February 2020","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 January 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 February 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"18"}}