{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,29]],"date-time":"2026-10-29T03:29:08Z","timestamp":1793244548760,"version":"4.5.0"},"reference-count":71,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2019,3,13]],"date-time":"2019-03-13T00:00:00Z","timestamp":1552435200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2019,3,13]],"date-time":"2019-03-13T00:00:00Z","timestamp":1552435200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    The oral cavity contains different types of microbial species that colonize human host via extensive cell-to-cell interactions and biofilm formation.\n                    <jats:italic>Candida albicans<\/jats:italic>\n                    <jats:bold>\u2014<\/jats:bold>\n                    a yeast-like fungus that inhabits mucosal surfaces\n                    <jats:bold>\u2014<\/jats:bold>\n                    is also a significant colonizer of subgingival sites in patients with chronic periodontitis. It is notable however that one of the main infectious agents that causes periodontal disease is an anaerobic bacterium\n                    <jats:bold>\u2014<\/jats:bold>\n                    <jats:italic>Porphyromonas gingivalis<\/jats:italic>\n                    . In our study, we evaluated the different strategies of both pathogens in the mutual colonization of an artificial surface and confirmed that a protective environment existed for\n                    <jats:italic>P. gingivalis<\/jats:italic>\n                    within developed fungal biofilm formed under oxic conditions where fungal cells grow mainly in their filamentous form i.e. hyphae. A direct physical contact between fungi and\n                    <jats:italic>P. gingivalis<\/jats:italic>\n                    was initiated via a modulation of gene expression for the major fungal cell surface adhesin Als3 and the aspartic proteases Sap6 and Sap9. Proteomic identification of the fungal surfaceome suggested also an involvement of the Mp65 adhesin and a \u201cmoonlighting\u201d protein, enolase, as partners for the interaction with\n                    <jats:italic>P. gingivalis<\/jats:italic>\n                    . Using mutant strains of these bacteria that are defective in the production of the gingipains\n                    <jats:bold>\u2014<\/jats:bold>\n                    the proteolytic enzymes that also harbor hemagglutinin domains\n                    <jats:bold>\u2014<\/jats:bold>\n                    significant roles of these proteins in the formation of bacteria-protecting biofilm were clearly demonstrated.\n                  <\/jats:p>","DOI":"10.1038\/s41598-019-40771-8","type":"journal-article","created":{"date-parts":[[2019,3,13]],"date-time":"2019-03-13T07:02:46Z","timestamp":1552460566000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":81,"title":["Adhesive protein-mediated cross-talk between Candida albicans and Porphyromonas gingivalis in dual species biofilm protects the anaerobic bacterium in unfavorable oxic environment"],"prefix":"10.1038","volume":"9","author":[{"given":"Dominika","family":"Bartnicka","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Justyna","family":"Karkowska-Kuleta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marcin","family":"Zawrotniak","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dorota","family":"Sata\u0142a","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kinga","family":"Michalik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gabriela","family":"Zielinska","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Oliwia","family":"Bochenska","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrzej","family":"Kozik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Izabela","family":"Ciaston","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joanna","family":"Koziel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lindsay C.","family":"Dutton","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3813-4410","authenticated-orcid":false,"given":"Angela H.","family":"Nobbs","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Barbara","family":"Potempa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5987-5712","authenticated-orcid":false,"given":"Zbigniew","family":"Baster","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zenon","family":"Rajfur","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"Potempa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Rapala-Kozik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2019,3,13]]},"reference":[{"key":"40771_CR1","doi-asserted-by":"crossref","first-page":"81","DOI":"10.4103\/0973-029X.141325","volume":"18","author":"A Agrawal","year":"2014","unstructured":"Agrawal, A., Singh, A., Verma, R. & Murari, A. Oral candidiasis: An overview. J. Oral Maxillofac. Pathol. 18, 81 (2014).","journal-title":"J. Oral Maxillofac. Pathol."},{"key":"40771_CR2","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1080\/13693780701218689","volume":"45","author":"J Perlroth","year":"2007","unstructured":"Perlroth, J., Choi, B. & Spellberg, B. Nosocomial fungal infections: epidemiology, diagnosis, and treatment. Med. Mycol. 45, 321\u2013346 (2007).","journal-title":"Med. Mycol."},{"key":"40771_CR3","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1016\/j.micinf.2016.01.002","volume":"18","author":"M Gulati","year":"2016","unstructured":"Gulati, M. & Nobile, C. J. Candida albicans biofilms: development, regulation and molecular mechanisms. Microbes Infect. 18, 310\u2013321 (2016).","journal-title":"Microbes Infect."},{"key":"40771_CR4","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1146\/annurev-micro-091014-104330","volume":"69","author":"CJ Nobile","year":"2015","unstructured":"Nobile, C. J. & Johnson, A. D. Candida albicans biofilms and human disease. Annu. Rev. Microbiol. 69, 71\u201392 (2015).","journal-title":"Annu. Rev. Microbiol."},{"key":"40771_CR5","doi-asserted-by":"crossref","first-page":"e34242","DOI":"10.1371\/journal.pone.0034242","volume":"7","author":"SM Huse","year":"2012","unstructured":"Huse, S. M., Ye, Y., Zhou, Y. & Fodor, A. A core human microbiome as viewed through 16S rRNA sequence clusters. PLoS One 7, e34242 (2012).","journal-title":"PLoS One"},{"key":"40771_CR6","doi-asserted-by":"crossref","first-page":"e1003616","DOI":"10.1371\/journal.ppat.1003616","volume":"9","author":"KH Metwalli","year":"2013","unstructured":"Metwalli, K. H., Khan, S. A., Krom, B. P. & Jabra-Rizk, M. A. Streptococcus mutans, Candida albicans, and the human mouth: a sticky situation. PLoS Pathog. 9, e1003616 (2013).","journal-title":"PLoS Pathog."},{"key":"40771_CR7","doi-asserted-by":"crossref","first-page":"532","DOI":"10.1007\/s00248-011-9989-2","volume":"63","author":"X He","year":"2012","unstructured":"He, X. et al. Adherence to Streptococci facilitates Fusobacterium nucleatum integration into an oral microbial community. Microb. Ecol. 63, 532\u2013542 (2012).","journal-title":"Microb. Ecol."},{"key":"40771_CR8","doi-asserted-by":"crossref","first-page":"3696","DOI":"10.1128\/IAI.00438-09","volume":"77","author":"CV Bamford","year":"2009","unstructured":"Bamford, C. V. et al. Streptococcus gordonii modulates Candida albicans biofilm formation through intergeneric communication. Infect. Immun. 77, 3696\u2013704 (2009).","journal-title":"Infect. Immun."},{"issue":"52","key":"40771_CR9","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1111\/j.1600-0757.2009.00311.x","volume":"2000","author":"M Kuboniwa","year":"2010","unstructured":"Kuboniwa, M. & Lamont, R. J. Subgingival biofilm formation. Periodontol. 2000(52), 38\u201352 (2010).","journal-title":"Periodontol."},{"key":"40771_CR10","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1038\/nri3785","volume":"15","author":"G Hajishengallis","year":"2015","unstructured":"Hajishengallis, G. Periodontitis: from microbial immune subversion to systemic inflammation. Nat. Rev. Immunol. 15, 30\u201344 (2015).","journal-title":"Nat. Rev. Immunol."},{"key":"40771_CR11","doi-asserted-by":"crossref","first-page":"45","DOI":"10.3389\/fimmu.2015.00045","volume":"6","author":"M Hussain","year":"2015","unstructured":"Hussain, M., Stover, C. M. & Dupont, A. P. gingivalis in periodontal disease and atherosclerosis - scenes of action for antimicrobial peptides and complement. Front. Immunol. 6, 45 (2015).","journal-title":"Front. Immunol."},{"key":"40771_CR12","doi-asserted-by":"crossref","first-page":"783","DOI":"10.1080\/13693780802060899","volume":"46","author":"B Urz\u00faa","year":"2008","unstructured":"Urz\u00faa, B. et al. Yeast diversity in the oral microbiota of subjects with periodontitis: Candida albicans and Candida dubliniensis colonize the periodontal pockets. Med. Mycol. 46, 783\u2013793 (2008).","journal-title":"Med. Mycol."},{"key":"40771_CR13","doi-asserted-by":"crossref","first-page":"3335","DOI":"10.1128\/JCM.01532-12","volume":"50","author":"BA McManus","year":"2012","unstructured":"McManus, B. A. et al. Enrichment of multilocus sequence typing clade 1 with oral Candida albicans isolates in patients with untreated periodontitis. J. Clin. Microbiol. 50, 3335\u20133344 (2012).","journal-title":"J. Clin. Microbiol."},{"key":"40771_CR14","first-page":"230","volume":"30","author":"JJ Melton","year":"2010","unstructured":"Melton, J. J. et al. Recovery of Candida dubliniensis and other Candida species from the oral cavity of subjects with periodontitis who had well-controlled and poorly controlled type 2. diabetes: A pilot study. Spec. Care Dent. 30, 230\u2013234 (2010).","journal-title":"diabetes: A pilot study. Spec. Care Dent."},{"key":"40771_CR15","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1111\/j.1365-2591.2009.01601.x","volume":"42","author":"TT Miranda","year":"2009","unstructured":"Miranda, T. T. et al. Diversity and frequency of yeasts from the dorsum of the tongue and necrotic root canals associated with primary apical periodontitis. Int. Endod. J. 42, 839\u2013844 (2009).","journal-title":"Int. Endod. J."},{"key":"40771_CR16","doi-asserted-by":"crossref","first-page":"467","DOI":"10.3109\/13693786.2011.633233","volume":"50","author":"JCO Sardi","year":"2012","unstructured":"Sardi, J. C. O., Duque, C., H\u00f6fling, J. F. & Gon\u00e7alves, R. B. Genetic and phenotypic evaluation of Candida albicans strains isolated from subgingival biofilm of diabetic patients with chronic periodontitis. Med. Mycol. 50, 467\u201375 (2012).","journal-title":"Med. Mycol."},{"key":"40771_CR17","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/S0003-9969(01)00114-5","volume":"47","author":"G Pizzo","year":"2002","unstructured":"Pizzo, G. et al. Genotyping and antifungal susceptibility of human subgingival Candida albicans isolates. Arch. Oral Biol. 47, 189\u2013196 (2002).","journal-title":"Arch. Oral Biol."},{"key":"40771_CR18","doi-asserted-by":"crossref","first-page":"2411","DOI":"10.1016\/j.cub.2014.08.057","volume":"24","author":"EP Fox","year":"2014","unstructured":"Fox, E. P. et al. Anaerobic bacteria grow within Candida albicans biofilms and induce biofilm formation in suspension cultures. Curr. Biol. 24, 2411\u20136 (2014).","journal-title":"Curr. Biol."},{"key":"40771_CR19","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1046\/j.0902-0055.2003.00121.x","volume":"19","author":"PI Diaz","year":"2004","unstructured":"Diaz, P. I. & Rogers, A. H. The effect of oxygen on the growth and physiology of Porphyromonas gingivalis. Oral Microbiol. Immunol. 19, 88\u201394 (2004).","journal-title":"Oral Microbiol. Immunol."},{"key":"40771_CR20","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1128\/EC.00316-06","volume":"6","author":"R Dumitru","year":"2007","unstructured":"Dumitru, R. et al. In vivo and in vitro anaerobic mating in Candida albicans. Eukaryot. Cell 6, 465\u2013472 (2007).","journal-title":"Eukaryot. Cell"},{"issue":"6","key":"40771_CR21","doi-asserted-by":"crossref","first-page":"e00187","DOI":"10.1128\/mSphere.00187-16","volume":"1","author":"PT van Leeuwen","year":"2016","unstructured":"van Leeuwen, P. T. et al. Interspecies interactions between Clostridium difficile and Candida albicans. mSphere 1(6), e00187\u201316 (2016).","journal-title":"mSphere"},{"key":"40771_CR22","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1111\/omi.12154","volume":"32","author":"IMG Cavalcanti","year":"2017","unstructured":"Cavalcanti, I. M. G., Cury, D. B., Jenkinson, A. A. & Nobbs, H. F. A. H. Interactions between Streptococcus oralis, Actinomyces oris, and Candida albicans in the development of multispecies oral microbial biofilms on salivary pellicle. Mol. Oral Microbiol. 32, 60\u201373 (2017).","journal-title":"Mol. Oral Microbiol."},{"key":"40771_CR23","doi-asserted-by":"crossref","unstructured":"Dutton, L. C., Jenkinson, H. F., Lamont, R. J. & Nobbs, A. H. Role of Candida albicans secreted aspartyl protease Sap9 in interkingdom biofilm formation. Pathog. Dis. 74 (2016).","DOI":"10.1093\/femspd\/ftw005"},{"key":"40771_CR24","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1080\/13693780701299333","volume":"45","author":"SA Klotz","year":"2007","unstructured":"Klotz, S. A. et al. Candida albicans Als proteins mediate aggregation with bacteria and yeasts. Med. Mycol. 45, 363\u2013370 (2007).","journal-title":"Med. Mycol."},{"key":"40771_CR25","doi-asserted-by":"crossref","first-page":"66","DOI":"10.3389\/fcimb.2014.00066","volume":"4","author":"RC Silva","year":"2014","unstructured":"Silva, R. C. et al. Extracellular enolase of Candida albicans is involved in colonization of mammalian intestinal epithelium. Front. Cell. Infect. Microbiol. 4, 66 (2014).","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"40771_CR26","doi-asserted-by":"crossref","first-page":"5795","DOI":"10.1002\/pmic.200600332","volume":"6","author":"DP Thomas","year":"2006","unstructured":"Thomas, D. P., Bachmann, S. P. & Lopez-Ribot, J. L. Proteomics for the analysis of the Candida albicans biofilm lifestyle. Proteomics 6, 5795\u2013804 (2006).","journal-title":"Proteomics"},{"key":"40771_CR27","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1186\/1471-2180-11-106","volume":"11","author":"S Sandini","year":"2011","unstructured":"Sandini, S. et al. The MP65 gene is required for cell wall integrity, adherence to epithelial cells and biofilm formation in Candida albicans. BMC Microbiol. 11, 106 (2011).","journal-title":"BMC Microbiol."},{"key":"40771_CR28","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1002\/yea.1775","volume":"27","author":"AG Sorgo","year":"2010","unstructured":"Sorgo, A. G. et al. Mass spectrometric analysis of the secretome of Candida albicans. Yeast 27, 661\u2013672 (2010).","journal-title":"Yeast"},{"key":"40771_CR29","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1186\/s12866-014-0303-6","volume":"14","author":"R Rajendran","year":"2014","unstructured":"Rajendran, R. et al. Extracellular DNA release confers heterogeneity in Candida albicans biofilm formation. BMC Microbiol. 14, 303 (2014).","journal-title":"BMC Microbiol."},{"key":"40771_CR30","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1091\/mbc.e03-03-0179","volume":"15","author":"M Martchenko","year":"2004","unstructured":"Martchenko, M., Alarco, A.-M., Harcus, D. & Whiteway, M. Superoxide dismutases in Candida albicans: transcriptional regulation and functional characterization of the hyphal-induced SOD5 gene. Mol. Biol. Cell 15, 456\u201367 (2004).","journal-title":"Mol. Biol. Cell"},{"key":"40771_CR31","doi-asserted-by":"crossref","first-page":"1622","DOI":"10.1128\/EC.00103-10","volume":"9","author":"AH Nobbs","year":"2010","unstructured":"Nobbs, A. H., Margaret Vickerman, M. & Jenkinson, H. F. Heterologous expression of Candida albicans cell wall-associated adhesins in Saccharomyces cerevisiae reveals differential specificities in adherence and biofilm formation and in binding oral Streptococcus gordonii. Eukaryot. Cell 9, 1622\u20131634 (2010).","journal-title":"Eukaryot. Cell"},{"key":"40771_CR32","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1556\/EuJMI.1.2011.1.7","volume":"1","author":"N Li","year":"2011","unstructured":"Li, N. & Collyer, C. A. Gingipains from Porphyromonas gingivalis \u2014 complex domain structures confer diverse functions. Eur. J. Microbiol. Immunol. 1, 41\u201358 (2011).","journal-title":"Eur. J. Microbiol. Immunol."},{"key":"40771_CR33","doi-asserted-by":"crossref","first-page":"3077","DOI":"10.1111\/j.1462-2920.2011.02573.x","volume":"13","author":"HF Jenkinson","year":"2011","unstructured":"Jenkinson, H. F. Beyond the oral microbiome. Environ. Microbiol. 13, 3077\u20133087 (2011).","journal-title":"Environ. Microbiol."},{"key":"40771_CR34","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1038\/nrmicro.2017.107","volume":"16","author":"MB Lohse","year":"2018","unstructured":"Lohse, M. B., Gulati, M., Johnson, A. D. & Nobile, C. J. Development and regulation of single- and multi-species Candida albicans biofilms. Nat. Rev. Microbiol. 16, 19\u201331 (2018).","journal-title":"Nat. Rev. Microbiol."},{"key":"40771_CR35","doi-asserted-by":"crossref","first-page":"e1005447","DOI":"10.1371\/journal.pgen.1005447","volume":"11","author":"PR Desai","year":"2015","unstructured":"Desai, P. R., van Wijlick, L., Kurtz, D., Juchimiuk, M. & Ernst, J. F. Hypoxia and temperature regulated morphogenesis in Candida albicans. PLoS Genet. 11, e1005447 (2015).","journal-title":"PLoS Genet."},{"key":"40771_CR36","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1128\/EC.00031-12","volume":"11","author":"N Grahl","year":"2012","unstructured":"Grahl, N., Shepardson, K. M., Chung, D. & Cramer, R. A. Hypoxia and fungal pathogenesis: to air or not to air? Eukaryot. Cell 11, 560\u201370 (2012).","journal-title":"Eukaryot. Cell"},{"key":"40771_CR37","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1574-6968.2009.01668.x","volume":"299","author":"ME Shirtliff","year":"2009","unstructured":"Shirtliff, M. E., Peters, B. M. & Jabra-Rizk, M. A. Cross-kingdom interactions: Candida albicans and bacteria. FEMS Microbiol. Lett. 299, 1\u20138 (2009).","journal-title":"FEMS Microbiol. Lett."},{"key":"40771_CR38","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3923\/mj.2019.1.7","volume":"9","author":"MM Janus","year":"2017","unstructured":"Janus, M. M. et al. Candida albicans alters the bacterial microbiome of early in vitro oral biofilms. J. Oral Microbiol. 9, 1\u201310 (2017).","journal-title":"J. Oral Microbiol."},{"key":"40771_CR39","doi-asserted-by":"crossref","first-page":"1731","DOI":"10.1007\/s00216-008-1947-6","volume":"391","author":"J Wesolowski","year":"2008","unstructured":"Wesolowski, J., Hassan, R. Y. A., Hodde, S., Bardroff, C. & Bilitewski, U. Sensing of oxygen in microtiter plates: a novel tool for screening drugs against pathogenic yeasts. Anal. Bioanal. Chem. 391, 1731\u20137 (2008).","journal-title":"Anal. Bioanal. Chem."},{"key":"40771_CR40","doi-asserted-by":"crossref","first-page":"1172","DOI":"10.1016\/j.archoralbio.2008.06.003","volume":"53","author":"LM Barros","year":"2008","unstructured":"Barros, L. M. et al. Genetic diversity and exoenzyme activities of Candida albicans and Candida dubliniensis isolated from the oral cavity of Brazilian periodontal patients. Arch. Oral Biol. 53, 1172\u20131178 (2008).","journal-title":"Arch. Oral Biol."},{"key":"40771_CR41","doi-asserted-by":"crossref","unstructured":"Winter, M. B. et al. Global identification of biofilm-specific proteolysis in Candida albicans. MBio 7 (2016).","DOI":"10.1128\/mBio.01514-16"},{"key":"40771_CR42","doi-asserted-by":"crossref","first-page":"1300366","DOI":"10.1080\/20002297.2017.1300366","volume":"9","author":"E Gerits","year":"2017","unstructured":"Gerits, E., Verstraeten, N. & Michiels, J. New approaches to combat Porphyromonas gingivalis. biofilms. J. Oral Microbiol. 9, 1300366 (2017).","journal-title":"biofilms. J. Oral Microbiol."},{"key":"40771_CR43","doi-asserted-by":"crossref","first-page":"1225","DOI":"10.1099\/jmm.0.021832-0","volume":"59","author":"HMHN Bandara","year":"2010","unstructured":"Bandara, H. M. H. N., Lam, O. L. T., Watt, R. M., Jin, L. J. & Samaranayake, L. P. Bacterial lipopolysaccharides variably modulate in vitro biofilm formation of Candida species. J. Med. Microbiol. 59, 1225\u201334 (2010).","journal-title":"J. Med. Microbiol."},{"key":"40771_CR44","doi-asserted-by":"crossref","first-page":"5266","DOI":"10.1128\/jb.173.17.5266-5274.1991","volume":"173","author":"PA Goulbourne","year":"1991","unstructured":"Goulbourne, P. A. & Ellen, R. P. Evidence that Porphyromonas (Bacteroides) gingivalis fimbriae function in adhesion to Actinomyces viscosus. J. Bacteriol. 173, 5266\u201374 (1991).","journal-title":"J. Bacteriol."},{"key":"40771_CR45","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1016\/S0378-1097(03)00615-3","volume":"226","author":"M Hashimoto","year":"2003","unstructured":"Hashimoto, M., Ogawa, S., Asai, Y., Takai, Y. & Ogawa, T. Binding of Porphyromonas gingivalis fimbriae to Treponema denticola dentilisin. FEMS Microbiol. Lett. 226, 267\u201371 (2003).","journal-title":"FEMS Microbiol. Lett."},{"key":"40771_CR46","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1111\/j.1399-302X.1993.tb00573.x","volume":"8","author":"RJ Lamont","year":"1993","unstructured":"Lamont, R. J., Bevan, C. A., Gil, S., Persson, R. E. & Rosan, B. Involvement of Porphyromonas gingivalis fimbriae in adherence to Streptococcus gordonii. Oral Microbiol. Immunol. 8, 272\u20136 (1993).","journal-title":"Oral Microbiol. Immunol."},{"key":"40771_CR47","doi-asserted-by":"crossref","first-page":"1341","DOI":"10.1128\/IAI.72.3.1341-1348.2004","volume":"72","author":"K Maeda","year":"2004","unstructured":"Maeda, K. et al. Glyceraldehyde-3-phosphate dehydrogenase of Streptococcus oralis functions as a coadhesin for Porphyromonas gingivalis major fimbriae. Infect. Immun. 72, 1341\u20138 (2004).","journal-title":"Infect. Immun."},{"key":"40771_CR48","doi-asserted-by":"crossref","unstructured":"Sztukowska, M. N. et al. Community Development between Porphyromonas gingivalis and Candida albicans Mediated by InlJ and Als3. MBio 9 (2018).","DOI":"10.1128\/mBio.00202-18"},{"key":"40771_CR49","doi-asserted-by":"crossref","unstructured":"Kamaguchi, A. et al. Effect of Porphyromonas gingivalis vesicles on coaggregation of Staphylococcus aureus to oral Microorganisms. Curr. Microbiol. 47 (2003).","DOI":"10.1007\/s00284-003-4069-6"},{"key":"40771_CR50","doi-asserted-by":"crossref","first-page":"17955","DOI":"10.1074\/jbc.274.25.17955","volume":"274","author":"Y Shi","year":"1999","unstructured":"Shi, Y. et al. Genetic analyses of proteolysis, hemoglobin binding, and hemagglutination of Porphyromonas gingivalis. Construction of mutants with a combination of rgpA, rgpB, kgp, and hagA. J. Biol. Chem. 274, 17955\u201360 (1999).","journal-title":"J. Biol. Chem."},{"key":"40771_CR51","doi-asserted-by":"crossref","first-page":"397","DOI":"10.2174\/1389203033487036","volume":"4","author":"J Potempa","year":"2003","unstructured":"Potempa, J., Sroka, A., Imamura, T. & Travis, J. Gingipains, the major cysteine proteinases and virulence factors of Porphyromonas gingivalis: structure, function and assembly of multidomain protein complexes. Curr. Protein Pept. Sci. 4, 397\u2013407 (2003).","journal-title":"Curr. Protein Pept. Sci."},{"key":"40771_CR52","doi-asserted-by":"crossref","first-page":"3980","DOI":"10.4049\/jimmunol.175.6.3980","volume":"175","author":"NM O\u2019Brien-Simpson","year":"2005","unstructured":"O\u2019Brien-Simpson, N. M. et al. An immune response directed to proteinase and adhesin functional epitopes protects against Porphyromonas gingivalis-induced periodontal bone loss. J. Immunol. 175, 3980\u20139 (2005).","journal-title":"J. Immunol."},{"key":"40771_CR53","doi-asserted-by":"crossref","first-page":"797","DOI":"10.1128\/IAI.00966-10","volume":"79","author":"M Rapala-Kozik","year":"2011","unstructured":"Rapala-Kozik, M. et al. Adsorption of components of the plasma kinin-forming system on the surface of Porphyromonas gingivalis involves gingipains as the major docking platforms. Infect. Immun. 79, 797\u2013805 (2011).","journal-title":"Infect. Immun."},{"key":"40771_CR54","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1111\/j.1574-695X.2010.00737.x","volume":"60","author":"R Ito","year":"2010","unstructured":"Ito, R., Ishihara, K., Shoji, M., Nakayama, K. & Okuda, K. Hemagglutinin\/Adhesin domains of Porphyromonas gingivalis play key roles in coaggregation with Treponema denticola. FEMS Immunol. Med. Microbiol. 60, 251\u201360 (2010).","journal-title":"FEMS Immunol. Med. Microbiol."},{"key":"40771_CR55","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1186\/s12866-014-0258-7","volume":"14","author":"K Bao","year":"2014","unstructured":"Bao, K. et al. Role of Porphyromonas gingivalis gingipains in multi-species biofilm formation. BMC Microbiol. 14, 258 (2014).","journal-title":"BMC Microbiol."},{"key":"40771_CR56","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1111\/omi.12051","volume":"29","author":"A Haraguchi","year":"2014","unstructured":"Haraguchi, A., Miura, M., Fujise, O., Hamachi, T. & Nishimura, F. Porphyromonas gingivalis gingipain is involved in the detachment and aggregation of Aggregatibacter actinomycetemcomitans biofilm. Mol. Oral Microbiol. 29, 131\u201343 (2014).","journal-title":"Mol. Oral Microbiol."},{"key":"40771_CR57","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1186\/1471-2180-12-281","volume":"12","author":"ES Ovchinnikova","year":"2012","unstructured":"Ovchinnikova, E. S., Krom, B. P., Busscher, H. J. & van der Mei, H. C. Evaluation of adhesion forces of Staphylococcus aureus along the length of Candida albicans hyphae. BMC Microbiol. 12, 281 (2012).","journal-title":"BMC Microbiol."},{"key":"40771_CR58","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1099\/mic.0.083378-0","volume":"161","author":"CV Bamford","year":"2015","unstructured":"Bamford, C. V., Nobbs, A. H., Barbour, M. E., Lamont, R. J. & Jenkinson, H. F. Functional regions of Candida albicans hyphal cell wall protein Als3 that determine interaction with the oral bacterium Streptococcus gordonii. Microbiology 161, 18\u201329 (2015).","journal-title":"Microbiology"},{"key":"40771_CR59","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1099\/mic.0.083485-0","volume":"161","author":"LM Schlecht","year":"2015","unstructured":"Schlecht, L. M. et al. Systemic Staphylococcus aureus infection mediated by Candida albicans hyphal invasion of mucosal tissue. Microbiology 161, 168\u2013181 (2015).","journal-title":"Microbiology"},{"key":"40771_CR60","doi-asserted-by":"crossref","first-page":"3476","DOI":"10.1128\/IAI.00179-11","volume":"79","author":"B Henderson","year":"2011","unstructured":"Henderson, B. & Martin, A. Bacterial Virulence in the Moonlight: Multitasking Bacterial Moonlighting Proteins Are Virulence Determinants in Infectious Disease. Infect. Immun. 79, 3476\u20133491 (2011).","journal-title":"Infect. Immun."},{"key":"40771_CR61","doi-asserted-by":"crossref","first-page":"825","DOI":"10.18388\/abp.2015_1142","volume":"62","author":"K Seweryn","year":"2015","unstructured":"Seweryn, K. et al. Kinetic and thermodynamic characterization of the interactions between the components of human plasma kinin-forming system and isolated and purified cell wall proteins of Candida albicans. Acta Biochim. Pol. 62, 825\u201335 (2015).","journal-title":"Acta Biochim. Pol."},{"key":"40771_CR62","doi-asserted-by":"crossref","first-page":"4304","DOI":"10.1074\/jbc.M117.810440","volume":"293","author":"E Reyna-Beltr\u00e1n","year":"2018","unstructured":"Reyna-Beltr\u00e1n, E. et al. The Candida albicans ENO1 gene encodes a transglutaminase involved in growth, cell division, morphogenesis, and osmotic protection. J. Biol. Chem. 293, 4304\u20134323 (2018).","journal-title":"J. Biol. Chem."},{"key":"40771_CR63","doi-asserted-by":"crossref","first-page":"3103","DOI":"10.1128\/IAI.01200-09","volume":"78","author":"RR Spurbeck","year":"2010","unstructured":"Spurbeck, R. R. & Arvidson, C. G. Lactobacillus jensenii surface-associated proteins inhibit Neisseria gonorrhoeae adherence to epithelial cells. Infect. Immun. 78, 3103\u201311 (2010).","journal-title":"Infect. Immun."},{"key":"40771_CR64","doi-asserted-by":"crossref","unstructured":"Karkowska-Kuleta, J. et al. The activity of bacterial peptidylarginine deiminase is important during formation of dual-species biofilm by periodontal pathogen Porphyromonas gingivalis and opportunistic fungus Candida albicans. Pathog. Dis. 76 (2018).","DOI":"10.1093\/femspd\/fty033"},{"key":"40771_CR65","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1159\/000441724","volume":"8","author":"M Benedyk","year":"2016","unstructured":"Benedyk, M. et al. Gingipains: Critical Factors in the Development of Aspiration Pneumonia Caused by Porphyromonas gingivalis. J. Innate Immun. 8, 185\u201398 (2016).","journal-title":"J. Innate Immun."},{"key":"40771_CR66","doi-asserted-by":"crossref","first-page":"807","DOI":"10.18388\/abp.2015_1140","volume":"62","author":"J Karkowska-Kuleta","year":"2015","unstructured":"Karkowska-Kuleta, J., Zajac, D., Bochenska, O. & Kozik, A. Surfaceome of pathogenic yeasts, Candida parapsilosis and Candida tropicalis, revealed with the use of cell surface shaving method and shotgun proteomic approach. Acta Biochim. Pol. 62, 807\u201319 (2015).","journal-title":"Acta Biochim. Pol."},{"key":"40771_CR67","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1080\/00362177585190271","volume":"13","author":"KL Lee","year":"1975","unstructured":"Lee, K. L., Buckley, H. R. & Campbell, C. C. An amino acid liquid synthetic medium for the development of mycelial and yeast forms of Candida albicans. Sabouraudia 13, 148\u201353 (1975).","journal-title":"Sabouraudia"},{"key":"40771_CR68","doi-asserted-by":"publisher","unstructured":"Potempa, J. & Nguyen, K.-A. Purification and characterization of gingipains. In Current Protocols in Protein Science 21.20.1\u201321.20.27, https:\/\/doi.org\/10.1002\/0471140864.ps2120s49 (John Wiley & Sons, Inc., 2007).","DOI":"10.1002\/0471140864.ps2120s49"},{"key":"40771_CR69","doi-asserted-by":"crossref","first-page":"391","DOI":"10.18388\/abp.2017_1609","volume":"64","author":"J Karkowska-Kuleta","year":"2017","unstructured":"Karkowska-Kuleta, J. et al. Binding of human plasminogen and high-molecular-mass kininogen by cell surface-exposed proteins of Candida parapsilosis. Acta Biochim. Pol. 64, 391\u2013400 (2017).","journal-title":"Acta Biochim. Pol."},{"key":"40771_CR70","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1515\/bc.2009.145","volume":"391","author":"J Karkowska-Kuleta","year":"2010","unstructured":"Karkowska-Kuleta, J., Kozik, A. & Rapala-Kozik, M. Binding and activation of the human plasma kinin-forming system on the cell walls of Candida albicans and Candida tropicalis. Biol. Chem. 391, 97\u2013103 (2010).","journal-title":"Biol. Chem."},{"key":"40771_CR71","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","volume":"25","author":"KJ Livak","year":"2001","unstructured":"Livak, K. J. & Schmittgen, T. D. Analysis of relative gene expression data using real-time quantitative PCR and the 2(\u2212Delta Delta C(T)) Method. Methods 25, 402\u20138 (2001).","journal-title":"Methods"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-40771-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-40771-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-40771-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,17]],"date-time":"2022-12-17T02:31:59Z","timestamp":1671244319000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-019-40771-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,3,13]]},"references-count":71,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2019,12]]}},"alternative-id":["40771"],"URL":"https:\/\/doi.org\/10.1038\/s41598-019-40771-8","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,3,13]]},"assertion":[{"value":"24 September 2018","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 January 2019","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"13 March 2019","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":"4376"}}