Porphyromonas gingivalis FimA and Mfa1 fimbriae: Current insights on localization, function, biogenesis, and genotype

Socransky, 1998, Microbial complexes in subgingival plaque, J Clin Periodontol, 25, 134, 10.1111/j.1600-051X.1998.tb02419.x Hajishengallis, 2012, The keystone-pathogen hypothesis, Nat Rev Microbiol, 10, 717, 10.1038/nrmicro2873 Hajishengallis, 2011, Low-abundance biofilm species orchestrates inflammatory periodontal disease through the commensal microbiota and complement, Cell Host Microbe, 10, 497, 10.1016/j.chom.2011.10.006 Lamont, 2018, The oral microbiota: dynamic communities and host interactions, Nat Rev Microbiol, 16, 745, 10.1038/s41579-018-0089-x Kebschull, 2010, "Gum bug, leave my heart alone!"—epidemiologic and mechanistic evidence linking periodontal infections and atherosclerosis, J Dent Res, 89, 879, 10.1177/0022034510375281 Maresz, 2013, Porphyromonas gingivalis facilitates the development and progression of destructive arthritis through its unique bacterial peptidylarginine deiminase (PAD), PLoS Pathog, 9, 10.1371/journal.ppat.1003627 Madianos, 2013, Adverse pregnancy outcomes (APOs) and periodontal disease: pathogenic mechanisms, J Periodontol, 84, S170, 10.1902/jop.2013.1340015 Dominy, 2019, Porphyromonas gingivalis in Alzheimer’s disease brains: evidence for disease causation and treatment with small-molecule inhibitors, Sci Adv, 5, eaau3333, 10.1126/sciadv.aau3333 Lamont, 1998, Life below the gum line: pathogenic mechanisms of Porphyromonas gingivalis, Microbiol Mol Biol Rev, 62, 1244, 10.1128/MMBR.62.4.1244-1263.1998 Yoshimura, 2009, Surface components of Porphyromonas gingivalis, J Periodontal Res, 44, 1, 10.1111/j.1600-0765.2008.01135.x Enersen, 2013, Porphyromonas gingivalis fimbriae, J Oral Microbiol, 5, 10.3402/jom.v5i0.20265 Hamada, 1998, The importance of fimbriae in the virulence and ecology of some oral bacteria, Oral Microbiol Immunol, 13, 129, 10.1111/j.1399-302X.1998.tb00724.x Lamont, 2002, In or out: the invasiveness of oral bacteria, Periodontol 2000, 30, 61, 10.1034/j.1600-0757.2002.03006.x Hasegawa, 2016, Role of Mfa5 in Expression of Mfa1 Fimbriae in Porphyromonas gingivalis, J Dent Res, 95, 1291, 10.1177/0022034516655083 Hasegawa, 2009, Anchoring and length regulation of Porphyromonas gingivalis Mfa1 fimbriae by the downstream gene product Mfa2, Microbiology (Reading, Engl), 155, 3333, 10.1099/mic.0.028928-0 Hasegawa, 2013, Localization and function of the accessory protein Mfa3 in Porphyromonas gingivalis Mfa1 fimbriae, Mol Oral Microbiol, 28, 467, 10.1111/omi.12040 Ikai, 2015, Mfa4, an accessory protein of Mfa1 fimbriae, modulates fimbrial biogenesis, cell auto-aggregation, and biofilm formation in Porphyromonas gingivalis, PLoS One, 10, 10.1371/journal.pone.0139454 Nishiyama, 2007, Involvement of minor components associated with the FimA fimbriae of Porphyromonas gingivalis in adhesive functions, Microbiology (Reading, Engl), 153, 1916, 10.1099/mic.0.2006/005561-0 Nagano, 2010, FimB regulates FimA fimbriation in Porphyromonas gingivalis, J Dent Res, 89, 903, 10.1177/0022034510370089 Naito, 2008, Determination of the genome sequence of Porphyromonas gingivalis strain ATCC 33277 and genomic comparison with strain W83 revealed extensive genome rearrangements in P. gingivalis, DNA Res, 15, 215, 10.1093/dnares/dsn013 Nelson, 2003, Complete genome sequence of the oral pathogenic Bacterium Porphyromonas gingivalis strain W83, J Bacteriol, 185, 5591, 10.1128/JB.185.18.5591-5601.2003 Watanabe, 2011, Complete genome sequence of the bacterium Porphyromonas gingivalis TDC60, which causes periodontal disease, J Bacteriol, 193, 4259, 10.1128/JB.05269-11 Xu, 2016, A distinct type of pilus from the human microbiome, Cell, 165, 690, 10.1016/j.cell.2016.03.016 Kloppsteck, 2016, Structure of the fimbrial protein Mfa4 from Porphyromonas gingivalis in its precursor form: implications for a donor-strand complementation mechanism, Sci Rep, 6, 22945, 10.1038/srep22945 Hall, 2018, Structural and functional characterization of shaft, anchor, and tip proteins of the Mfa1 fimbria from the periodontal pathogen Porphyromonas gingivalis, Sci Rep, 8, 1793, 10.1038/s41598-018-20067-z Shoji, 2020, Biogenesis of type V pili, Microbiol Immunol, 64, 643, 10.1111/1348-0421.12838 Shibata, 2020, Structure of polymerized type V pilin reveals assembly mechanism involving protease-mediated strand exchange, Nat Microbiol, 5, 830, 10.1038/s41564-020-0705-1 Hospenthal, 2017, A comprehensive guide to pilus biogenesis in Gram-negative bacteria, Nat Rev Microbiol, 15, 365, 10.1038/nrmicro.2017.40 Heidler, 2021, Porphyromonas gingivalis fimbrial protein Mfa5 contains a von Willebrand factor domain and an intramolecular isopeptide, Commun Biol, 4, 106, 10.1038/s42003-020-01621-w Nakayama, 2015, Porphyromonas gingivalis and related bacteria: from colonial pigmentation to the type IX secretion system and gliding motility, J Periodontal Res, 50, 1, 10.1111/jre.12255 Veith, 2017, Type IX secretion: the generation of bacterial cell surface coatings involved in virulence, gliding motility and the degradation of complex biopolymers, Mol Microbiol, 106, 35, 10.1111/mmi.13752 Lasica, 2017, The type IX secretion system (T9SS): highlights and recent insights into its structure and function, Front Cell Infect Microbiol, 7, 215, 10.3389/fcimb.2017.00215 Hasegawa, 2014, Porphyromonas gingivalis fimbriae: recent developments describing the function and localization of mfa1 gene cluster proteins, J Oral Biosci, 56, 86, 10.1016/j.job.2014.04.002 Amano, 2004, Variations of Porphyromonas gingivalis fimbriae in relation to microbial pathogenesis, J Periodontal Res, 39, 136, 10.1111/j.1600-0765.2004.00719.x Fujiwara-Takahashi, 2020, Phylogenetic diversity in fim and mfa gene clusters between Porphyromonas gingivalis and Porphyromonas gulae, as a potential cause of host specificity, J Oral Microbiol, 12, 10.1080/20002297.2020.1775333 Nagano, 2015, A major fimbrilin variant of Mfa1 fimbriae in Porphyromonas gingivalis, J Dent Res, 94, 1143, 10.1177/0022034515588275 Sakae, 2021, Diversity analysis of genes encoding Mfa1-fimbrial components in Porphyromonas gingivalis strains, PLoS One, 16, e0255111, 10.1371/journal.pone.0255111 Nagano, 2018, Distribution of Porphyromonas gingivalis fimA and mfa1 fimbrial genotypes in subgingival plaques, PeerJ, 6, e5581, 10.7717/peerj.5581 Yoshimura, 1984, Purification and characterization of a novel type of fimbriae from the oral anaerobe Bacteroides gingivalis, J Bacteriol, 160, 949, 10.1128/jb.160.3.949-957.1984 Dickinson, 1988, Molecular cloning and sequencing of the gene encoding the fimbrial subunit protein of Bacteroides gingivalis, J Bacteriol, 170, 1658, 10.1128/jb.170.4.1658-1665.1988 Yoshimura, 1989, Purification and properties of a 75-kilodalton major protein, an immunodominant surface antigen, from the oral anaerobe Bacteroides gingivalis, Infect Immun, 57, 3646, 10.1128/iai.57.11.3646-3652.1989 Hamada, 1996, Isolation and characterization of a minor fimbria from Porphyromonas gingivalis, Infect Immun, 64, 4788, 10.1128/iai.64.11.4788-4794.1996 Ogawa, 1994, Molecular cloning and characterization of the genes encoding the immunoreactive major cell-surface proteins of Porphyromonas gingivalis, FEMS Microbiol Lett, 120, 23, 10.1111/j.1574-6968.1994.tb07002.x Ogawa, 1995, Immunochemical characterisation and epitope mapping of a novel fimbrial protein (Pg-II fimbria) of Porphyromonas gingivalis, FEMS Immunol Med Microbiol, 11, 247, 10.1111/j.1574-695X.1995.tb00122.x Hamada, 2002, Cytokine production induced by a 67-kDa fimbrial protein from Porphyromonas gingivalis, Oral Microbiol Immunol, 17, 197, 10.1034/j.1399-302X.2002.170311.x Watanabe, 1992, Molecular cloning and expression of a major surface protein (the 75-kDa protein) of Porphyromonas (Bacteroides) gingivalis in Escherichia coli, FEMS Microbiol Lett, 71, 47, 10.1111/j.1574-6968.1992.tb05233.x Park, 2005, Short fimbriae of Porphyromonas gingivalis and their role in coadhesion with Streptococcus gordonii, Infect Immun, 73, 3983, 10.1128/IAI.73.7.3983-3989.2005 Shoji, 2004, The major structural components of two cell surface filaments of Porphyromonas gingivalis are matured through lipoprotein precursors, Mol Microbiol, 52, 1513, 10.1111/j.1365-2958.2004.04105.x Shoji, 2010, Recombinant Porphyromonas gingivalis FimA preproprotein expressed in Escherichia coli is lipidated and the mature or processed recombinant FimA protein forms a short filament in vitro, Can J Microbiol, 56, 959, 10.1139/W10-084 Kadowaki, 1998, Arg-gingipain acts as a major processing enzyme for various cell surface proteins in Porphyromonas gingivalis, J Biol Chem, 273, 29072, 10.1074/jbc.273.44.29072 Onoe, 1995, Identification of Porphyromonas gingivalis prefimbrilin possessing a long leader peptide: possible involvement of trypsin-like protease in fimbrilin maturation, Microb Pathog, 19, 351, 10.1016/S0882-4010(96)80006-4 Nagano, 2012, Porphyromonas gingivalis FimA fimbriae: roles of the fim gene cluster in the fimbrial assembly and antigenic heterogeneity among fimA genotypes, J Oral Biosci, 54, 160, 10.1016/j.job.2012.07.002 Lee, 2018, Maturation of the Mfa1 fimbriae in the oral pathogen Porphyromonas gingivalis, Front Cell Infect Microbiol, 8, 137, 10.3389/fcimb.2018.00137 Alaei, 2019, Peptide-based inhibitors of fimbrial biogenesis in Porphyromonas gingivalis, Infect Immun, 87, 10.1128/IAI.00750-18 Chung, 2000, Identification of a Porphyromonas gingivalis receptor for the Streptococcus gordonii SspB protein, Infect Immun, 68, 6758, 10.1128/IAI.68.12.6758-6762.2000 Watanabe, 1996, Sequence and product analyses of the four genes downstream from the fimbrilin gene (fimA) of the oral anaerobe Porphyromonas gingivalis, Microbiol Immunol, 40, 725, 10.1111/j.1348-0421.1996.tb01133.x Whittaker, 2002, Distribution and evolution of von Willebrand/integrin A domains: widely dispersed domains with roles in cell adhesion and elsewhere, Mol Biol Cell, 13, 3369, 10.1091/mbc.e02-05-0259 Glew, 2012, PG0026 is the C-terminal signal peptidase of a novel secretion system of Porphyromonas gingivalis, J Biol Chem, 287, 24605, 10.1074/jbc.M112.369223 Nagano, 2013, FimA fimbriae of the periodontal disease-associated bacterium Porphyromonas gingivalis, Yakugaku Zasshi, 133, 963, 10.1248/yakushi.13-00177 Amano, 1996, Structural domains of Porphyromonas gingivalis recombinant fimbrillin that mediate binding to salivary proline-rich protein and statherin, Infect Immun, 64, 1631, 10.1128/iai.64.5.1631-1637.1996 Kataoka, 1997, Active sites of salivary proline-rich protein for binding to Porphyromonas gingivalis fimbriae, Infect Immun, 65, 3159, 10.1128/iai.65.8.3159-3164.1997 Nagata, 1997, Role of the carboxyl-terminal region of Porphyromonas gingivalis fimbrillin in binding to salivary proteins, Infect Immun, 65, 422, 10.1128/iai.65.2.422-427.1997 Amano, 1999, Molecular interactions of Porphyromonas gingivalis fimbriae with host proteins: kinetic analyses based on surface plasmon resonance, Infect Immun, 67, 2399, 10.1128/IAI.67.5.2399-2405.1999 Kataoka, 1999, Secretion of functional salivary peptide by Streptococcus gordonii which inhibits fimbria-mediated adhesion of Porphyromonas gingivalis, Infect Immun, 67, 3780, 10.1128/IAI.67.8.3780-3785.1999 Amano, 1998, Binding of Porphyromonas gingivalis fimbriae to proline-rich glycoproteins in parotid saliva via a domain shared by major salivary components, Infect Immun, 66, 2072, 10.1128/IAI.66.5.2072-2077.1998 Amano, 1994, Salivary receptors for recombinant fimbrillin of Porphyromonas gingivalis, Infect Immun, 62, 3372, 10.1128/iai.62.8.3372-3380.1994 Amano, 1996, Binding sites of salivary statherin for Porphyromonas gingivalis recombinant fimbrillin, Infect Immun, 64, 4249, 10.1128/iai.64.10.4249-4254.1996 Zhang, 2013, Integrin alpha5beta1-fimbriae binding and actin rearrangement are essential for Porphyromonas gingivalis invasion of osteoblasts and subsequent activation of the JNK pathway, BMC Microbiol, 13, 5, 10.1186/1471-2180-13-5 Amano, 2007, Disruption of epithelial barrier and impairment of cellular function by Porphyromonas gingivalis, Front Biosci, 12, 3965, 10.2741/2363 Nakagawa, 2005, Inhibitory effects of Porphyromonas gingivalis fimbriae on interactions between extracellular matrix proteins and cellular integrins, Microbes Infect, 7, 157, 10.1016/j.micinf.2004.10.007 Yilmaz, 2002, Involvement of integrins in fimbriae-mediated binding and invasion by Porphyromonas gingivalis, Cell Microbiol, 4, 305, 10.1046/j.1462-5822.2002.00192.x Tamai, 2005, Requirement for intercellular adhesion molecule 1 and caveolae in invasion of human oral epithelial cells by Porphyromonas gingivalis, Infect Immun, 73, 6290, 10.1128/IAI.73.10.6290-6298.2005 Sojar, 2005, Identification of glyceraldehyde-3-phosphate dehydrogenase of epithelial cells as a second molecule that binds to Porphyromonas gingivalis fimbriae, FEMS Immunol Med Microbiol, 45, 25, 10.1016/j.femsim.2005.01.006 Sojar, 2002, Porphyromonas gingivalis fimbriae bind to cytokeratin of epithelial cells, Infect Immun, 70, 96, 10.1128/IAI.70.1.96-101.2002 Nakamura, 1999, Specific interactions between Porphyromonas gingivalis fimbriae and human extracellular matrix proteins, FEMS Microbiol Lett, 175, 267, 10.1111/j.1574-6968.1999.tb13630.x Kontani, 1996, Cysteine protease of Porphyromonas gingivalis 381 enhances binding of fimbriae to cultured human fibroblasts and matrix proteins, Infect Immun, 64, 756, 10.1128/iai.64.3.756-762.1996 Naito, 1993, Adherence and hydrophobicity of invasive and noninvasive strains of Porphyromonas gingivalis, Oral Microbiol Immunol, 8, 195, 10.1111/j.1399-302X.1993.tb00559.x Kontani, 1999, Inhibitory effects of protamines on proteolytic and adhesive activities of Porphyromonas gingivalis, Infect Immun, 67, 4917, 10.1128/IAI.67.9.4917-4920.1999 Murakami, 1998, Fibronectin in saliva inhibits Porphyromonas gingivalis fimbria-induced expression of inflammatory cytokine gene in mouse macrophages, FEMS Immunol Med Microbiol, 22, 257, 10.1111/j.1574-695X.1998.tb01214.x Kontani, 1997, Adherence of Porphyromonas gingivalis to matrix proteins via a fimbrial cryptic receptor exposed by its own arginine-specific protease, Mol Microbiol, 24, 1179, 10.1046/j.1365-2958.1997.4321788.x Murakami, 1996, Porphyromonas gingivalis fimbrillin is one of the fibronectin-binding proteins, Infect Immun, 64, 2571, 10.1128/iai.64.7.2571-2576.1996 Sojar, 1995, Fibronectin binding domain of P. gingivalis fimbriae, Biochem Biophys Res Commun, 216, 785, 10.1006/bbrc.1995.2690 Maeda, 2004, Characterization of binding of Streptococcus oralis glyceraldehyde-3-phosphate dehydrogenase to Porphyromonas gingivalis major fimbriae, Infect Immun, 72, 5475, 10.1128/IAI.72.9.5475-5477.2004 Maeda, 2013, Identification and characterization of Porphyromonas gingivalis client proteins that bind to Streptococcus oralis glyceraldehyde-3-phosphate dehydrogenase, Infect Immun, 81, 753, 10.1128/IAI.00875-12 Maeda, 2004, Oral streptococcal glyceraldehyde-3-phosphate dehydrogenase mediates interaction with Porphyromonas gingivalis fimbriae, Microbes Infect, 6, 1163, 10.1016/j.micinf.2004.06.005 Maeda, 2004, Glyceraldehyde-3-phosphate dehydrogenase of Streptococcus oralis functions as a coadhesin for Porphyromonas gingivalis major fimbriae, Infect Immun, 72, 1341, 10.1128/IAI.72.3.1341-1348.2004 Nagata, 2009, Identification of the binding domain of Streptococcus oralis glyceraldehyde-3-phosphate dehydrogenase for Porphyromonas gingivalis major fimbriae, Infect Immun, 77, 5130, 10.1128/IAI.00439-09 Ellen, 1992, Inhibition of Actinomyces viscosus–Porphyromonas gingivalis coadhesion by trypsin and other proteins, Oral Microbiol Immunol, 7, 198, 10.1111/j.1399-302X.1992.tb00025.x Goulbourne, 1991, Evidence that Porphyromonas (Bacteroides) gingivalis fimbriae function in adhesion to Actinomyces viscosus, J Bacteriol, 173, 5266, 10.1128/jb.173.17.5266-5274.1991 Yamada, 2005, Synergistic biofilm formation by Treponema denticola and Porphyromonas gingivalis, FEMS Microbiol Lett, 250, 271, 10.1016/j.femsle.2005.07.019 Hashimoto, 2003, Binding of Porphyromonas gingivalis fimbriae to Treponema denticola dentilisin, FEMS Microbiol Lett, 226, 267, 10.1016/S0378-1097(03)00615-3 Umemoto, 1999, Fimbria-mediated coaggregation between human oral anaerobes Treponema medium and Porphyromonas gingivalis, Microbiol Immunol, 43, 837, 10.1111/j.1348-0421.1999.tb01218.x Hajishengallis, 2006, Porphyromonas gingivalis fimbriae proactively modulate beta2 integrin adhesive activity and promote binding to and internalization by macrophages, Infect Immun, 74, 5658, 10.1128/IAI.00784-06 Harokopakis, 2006, TLR2 transmodulates monocyte adhesion and transmigration via Rac1- and PI3K-mediated inside-out signaling in response to Porphyromonas gingivalis fimbriae, J Immunol, 176, 7645, 10.4049/jimmunol.176.12.7645 Harokopakis, 2005, Integrin activation by bacterial fimbriae through a pathway involving CD14, Toll-like receptor 2, and phosphatidylinositol-3-kinase, Eur J Immunol, 35, 1201, 10.1002/eji.200425883 Kanaya, 2009, Porphyromonas gingivalis fimbriae induce unique dendritic cell subsets via Toll-like receptor 2, J Periodontal Res, 44, 543, 10.1111/j.1600-0765.2008.01149.x Hajishengallis, 2009, Induction of distinct TLR2-mediated proinflammatory and proadhesive signaling pathways in response to Porphyromonas gingivalis fimbriae, J Immunol, 182, 6690, 10.4049/jimmunol.0900524 Davey, 2008, Bacterial fimbriae stimulate proinflammatory activation in the endothelium through distinct TLRs, J Immunol, 180, 2187, 10.4049/jimmunol.180.4.2187 Zhou, 2007, Identification of signaling pathways in macrophage exposed to Porphyromonas gingivalis or to its purified cell wall components, J Immunol, 179, 7777, 10.4049/jimmunol.179.11.7777 Hajishengallis, 2007, Porphyromonas gingivalis interactions with complement receptor 3 (CR3): innate immunity or immune evasion?, Front Biosci, 12, 4547, 10.2741/2409 Eskan, 2007, Differential activation of human gingival epithelial cells and monocytes by Porphyromonas gingivalis fimbriae, Infect Immun, 75, 892, 10.1128/IAI.01604-06 Hajishengallis, 2006, Differential interactions of fimbriae and lipopolysaccharide from Porphyromonas gingivalis with the Toll-like receptor 2-centred pattern recognition apparatus, Cell Microbiol, 8, 1557, 10.1111/j.1462-5822.2006.00730.x Coats, 2019, The distinct immune-stimulatory capacities of Porphyromonas gingivalis strains 381 and ATCC 33277 are determined by the fimB allele and gingipain activity, Infect Immun, 87, 10.1128/IAI.00319-19 Hajishengallis, 2007, Complement receptor 3 blockade promotes IL-12-mediated clearance of Porphyromonas gingivalis and negates its virulence in vivo, J Immunol, 179, 2359, 10.4049/jimmunol.179.4.2359 Hajishengallis, 2008, Subversion of innate immunity by periodontopathic bacteria via exploitation of complement receptor-3, Adv Exp Med Biol, 632, 203 Hajishengallis, 2005, Peptide mapping of bacterial fimbrial epitopes interacting with pattern recognition receptors, J Biol Chem, 280, 38902, 10.1074/jbc.M507326200 Takeshita, 1998, Porphyromonas gingivalis fimbriae use beta2 integrin (CD11/CD18) on mouse peritoneal macrophages as a cellular receptor, and the CD18 beta chain plays a functional role in fimbrial signaling, Infect Immun, 66, 4056, 10.1128/IAI.66.9.4056-4060.1998 Hajishengallis, 2008, Pathogen induction of CXCR4/TLR2 cross-talk impairs host defense function, Proc Natl Acad Sci U S A, 105, 13532, 10.1073/pnas.0803852105 Hajishengallis, 2013, Mechanism and implications of CXCR4-mediated integrin activation by Porphyromonas gingivalis, Mol Oral Microbiol, 28, 239, 10.1111/omi.12021 Lin, 2006, Porphyromonas gingivalis minor fimbriae are required for cell-cell interactions, Infect Immun, 74, 6011, 10.1128/IAI.00797-06 Kuboniwa, 2009, Distinct roles of long/short fimbriae and gingipains in homotypic biofilm development by Porphyromonas gingivalis, BMC Microbiol, 9, 105, 10.1186/1471-2180-9-105 Zeituni, 2010, The native 67-kilodalton minor fimbria of Porphyromonas gingivalis is a novel glycoprotein with DC-SIGN-targeting motifs, J Bacteriol, 192, 4103, 10.1128/JB.00275-10 Zeituni, 2009, Targeting of DC-SIGN on human dendritic cells by minor fimbriated Porphyromonas gingivalis strains elicits a distinct effector T cell response, J Immunol, 183, 5694, 10.4049/jimmunol.0901030 El-Awady, 2015, Porphyromonas gingivalis evasion of autophagy and intracellular killing by human myeloid dendritic cells involves DC-SIGN-TLR2 crosstalk, PLoS Pathog, 10 Daep, 2011, Structural dissection and in vivo effectiveness of a peptide inhibitor of Porphyromonas gingivalis adherence to Streptococcus gordonii, Infect Immun, 79, 67, 10.1128/IAI.00361-10 Daep, 2008, Interaction of Porphyromonas gingivalis with oral streptococci requires a motif that resembles the eukaryotic nuclear receptor box protein-protein interaction domain, Infect Immun, 76, 3273, 10.1128/IAI.00366-08 Daep, 2006, Structural characterization of peptide-mediated inhibition of Porphyromonas gingivalis biofilm formation, Infect Immun, 74, 5756, 10.1128/IAI.00813-06 Roky, 2020, Identification of functional domains of the minor fimbrial antigen involved in the interaction of Porphyromonas gingivalis with oral streptococci, Mol Oral Microbiol, 35, 66, 10.1111/omi.12280 Meghil, 2019, Disruption of immune homeostasis in human dendritic cells via regulation of autophagy and apoptosis by Porphyromonas gingivalis, Front Immunol, 10, 2286, 10.3389/fimmu.2019.02286 El-Awady, 2019, Polymicrobial synergy within oral biofilm promotes invasion of dendritic cells and survival of consortia members, NPJ Biofilms Microbiomes, 5, 11, 10.1038/s41522-019-0084-7 Hiramine, 2003, Porphyromonas gingivalis 67-kDa fimbriae induced cytokine production and osteoclast differentiation utilizing TLR2, FEMS Microbiol Lett, 229, 49, 10.1016/S0378-1097(03)00788-2 Takayanagi, 2020, Porphyromonas gingivalis Mfa1 induces chemokine and cell adhesion molecules in mouse gingival fibroblasts via toll-like receptors, J Clin Med, 9, 10.3390/jcm9124004 Jones, 1995, FimH adhesin of type 1 pili is assembled into a fibrillar tip structure in the Enterobacteriaceae, Proc Natl Acad Sci U S A, 92, 2081, 10.1073/pnas.92.6.2081 Kuehn, 1992, P pili in uropathogenic E. coli are composite fibres with distinct fibrillar adhesive tips, Nature, 356, 252, 10.1038/356252a0 Nagano, 2017, Novel fimbrilin PGN_1808 in Porphyromonas gingivalis, PLoS One, 12, 10.1371/journal.pone.0173541 Kuboniwa, 2010, Subgingival biofilm formation, Periodontol 2000, 52, 38, 10.1111/j.1600-0757.2009.00311.x Amano, 1999, Distribution of Porphyromonas gingivalis strains with fimA genotypes in periodontitis patients, J Clin Microbiol, 37, 1426, 10.1128/JCM.37.5.1426-1430.1999 Zhao, 2007, Prevalence of fimA genotypes of Porphyromonas gingivalis and periodontal health status in Chinese adults, J Periodontal Res Suppl, 42, 511, 10.1111/j.1600-0765.2007.00975.x Wang, 2020, The prevalence of fimA genotypes of Porphyromonas gingivalis in patients with chronic periodontitis: a meta-analysis, PLoS One, 15 Beikler, 2003, Prevalence of Porphyromonas gingivalis fimA genotypes in Caucasians, Eur J Oral Sci, 111, 390, 10.1034/j.1600-0722.2003.00065.x Nagano, 2013, Genetic and antigenic analyses of Porphyromonas gingivalis FimA fimbriae, Mol Oral Microbiol, 28, 392, 10.1111/omi.12032 Curtis, 1999, The rag locus of Porphyromonas gingivalis: a novel pathogenicity island, J Periodontal Res, 34, 400, 10.1111/j.1600-0765.1999.tb02273.x Hasegawa, 2021, Purification of native Mfa1 fimbriae from Porphyromonas gingivalis, Methods Mol Biol, 2210, 75, 10.1007/978-1-0716-0939-2_8