{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T07:19:24Z","timestamp":1790752764138,"version":"4.1.0"},"reference-count":127,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2018,2,27]],"date-time":"2018-02-27T00:00:00Z","timestamp":1519689600000},"content-version":"unspecified","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100008747","name":"Center for Clinical and Translational Science, Ohio State University","doi-asserted-by":"publisher","award":["R21 AI113477"],"award-info":[{"award-number":["R21 AI113477"]}],"id":[{"id":"10.13039\/100008747","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008747","name":"Center for Clinical and Translational Science, Ohio State University","doi-asserted-by":"publisher","award":["R01 AI24121"],"award-info":[{"award-number":["R01 AI24121"]}],"id":[{"id":"10.13039\/100008747","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008747","name":"Center for Clinical and Translational Science, Ohio State University","doi-asserted-by":"publisher","award":["R01 HL127651"],"award-info":[{"award-number":["R01 HL127651"]}],"id":[{"id":"10.13039\/100008747","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell. Infect. Microbiol."],"DOI":"10.3389\/fcimb.2018.00038","type":"journal-article","created":{"date-parts":[[2018,2,27]],"date-time":"2018-02-27T04:45:42Z","timestamp":1519706742000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":101,"title":["Factors Mediating Environmental Biofilm Formation by Legionella pneumophila"],"prefix":"10.3389","volume":"8","author":[{"given":"Arwa","family":"Abu Khweek","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amal O.","family":"Amer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2018,2,27]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"18","DOI":"10.3389\/fcimb.2013.00018","article-title":"Biofilm-derived Legionella pneumophila evades the innate immune response in macrophages","volume":"3","author":"Abu Khweek","year":"2013","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1424","DOI":"10.1099\/mic.0.036640-0","article-title":"Analysis of HmsH and its role in plague biofilm formation","volume":"156","author":"Abu Khweek","year":"2010","journal-title":"Microbiology"},{"key":"B3","doi-asserted-by":"publisher","first-page":"e0146410","DOI":"10.1371\/journal.pone.0146410","article-title":"The sphingosine-1-phosphate lyase (LegS2) contributes to the restriction of Legionella pneumophila in murine macrophages","volume":"11","author":"Abu Khweek","year":"2016","journal-title":"PLoS ONE"},{"key":"B4","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1111\/j.1462-5822.2006.00785.x","article-title":"Anti-apoptotic signalling by the Dot\/Icm secretion system of L","volume":"9","author":"Abu-Zant","year":"2007","journal-title":"pneumophila. Cell Microbiol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"1222","DOI":"10.1128\/IAI.01077-13","article-title":"Three antagonistic cyclic di-GMP-catabolizing enzymes promote differential Dot\/Icm effector delivery and intracellular survival at the early steps of Legionella pneumophila infection","volume":"82","author":"Allombert","year":"2014","journal-title":"Infect. Immun."},{"key":"B6","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1007\/s00284-014-0648-y","article-title":"Role of biofilm in protection of the replicative form of Legionella pneumophila","volume":"69","author":"Andreozzi","year":"2014","journal-title":"Curr. Microbiol."},{"key":"B7","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1016\/S0168-6445(03)00055-X","article-title":"Bacterial iron homeostasis","volume":"27","author":"Andrews","year":"2003","journal-title":"FEMS Microbiol. Rev."},{"key":"B8","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1046\/j.1462-2920.1999.00046.x","article-title":"Legionella: from environmental habitats to disease pathology, detection and control","volume":"1","author":"Atlas","year":"1999","journal-title":"Environ. Microbiol."},{"key":"B9","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00248-007-9245-y","article-title":"Evaluation of adherence, hydrophobicity, aggregation, and biofilm development of Flavobacterium johnsoniae-like isolates","volume":"55","author":"Basson","year":"2008","journal-title":"Microb. Ecol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"486","DOI":"10.3389\/fmicb.2016.00486","article-title":"Legionella pneumophila: the paradox of a highly sensitive opportunistic waterborne pathogen able to persist in the environment","volume":"7","author":"Berjeaud","year":"2016","journal-title":"Front. Microbiol."},{"key":"B11","doi-asserted-by":"publisher","first-page":"2187","DOI":"10.1128\/AEM.01214-07","article-title":"Packaging of live Legionella pneumophila into pellets expelled by Tetrahymena spp. does not require bacterial replication and depends on a Dot\/Icm-mediated survival mechanism","volume":"74","author":"Berk","year":"2008","journal-title":"Appl. Environ. Microbiol."},{"key":"B12","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1128\/AEM.64.1.279-286.1998","article-title":"Production of respirable vesicles containing live Legionella pneumophila cells by two Acanthamoeba spp","volume":"64","author":"Berk","year":"1998","journal-title":"Appl. Environ. Microbiol."},{"key":"B13","doi-asserted-by":"publisher","first-page":"328","DOI":"10.1139\/m92-055","article-title":"In situ colonization of polyvinyl chloride, brass, and copper by Legionella pneumophila","volume":"38","author":"Bezanson","year":"1992","journal-title":"Can. J. Microbiol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"e77875","DOI":"10.1371\/journal.pone.0077875","article-title":"Intra-amoeba multiplication induces chemotaxis and biofilm colonization and formation for Legionella","volume":"8","author":"Bigot","year":"2013","journal-title":"PLoS ONE"},{"key":"B15","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1111\/j.1365-2672.2008.03765.x","article-title":"Multiresistant waterborne pathogens isolated from water reservoirs and cooling systems","volume":"105","author":"Blasco","year":"2008","journal-title":"J. Appl. Microbiol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1111\/j.1365-2958.2010.07470.x","article-title":"Systematic analysis of cyclic di-GMP signalling enzymes and their role in biofilm formation and virulence in Yersinia pestis","volume":"79","author":"Bobrov","year":"2011","journal-title":"Mol. Microbiol."},{"key":"B17","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1016\/S1387-2656(05)11011-4","article-title":"Water ecology of Legionella and protozoan: environmental and public health perspectives","volume":"11","author":"Borella","year":"2005","journal-title":"Biotechnol. Annu. Rev."},{"key":"B18","doi-asserted-by":"publisher","first-page":"1341","DOI":"10.1111\/j.1462-2920.2006.01229.x","article-title":"Long-term survival of Legionella pneumophila associated with Acanthamoeba castellanii vesicles","volume":"9","author":"Bouyer","year":"2007","journal-title":"Environ. Microbiol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1111\/j.1462-5822.2009.01398.x","article-title":"Caenorhabditis is a metazoan host for Legionella","volume":"12","author":"Brassinga","year":"2010","journal-title":"Cell. Microbiol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"1228","DOI":"10.1111\/j.1462-5822.2006.00703.x","article-title":"Virulence strategies for infecting phagocytes deduced from the in vivo transcriptional program of Legionella pneumophila","volume":"8","author":"Br\u00fcggemann","year":"2006","journal-title":"Cell. Microbiol."},{"key":"B21","doi-asserted-by":"publisher","first-page":"280","DOI":"10.1111\/1574-6941.12294","article-title":"Microbial diversities (16S and 18S rRNA gene pyrosequencing) and environmental pathogens within drinking water biofilms grown on the common premise plumbing materials unplasticized polyvinylchloride and copper","volume":"88","author":"Buse","year":"2014","journal-title":"FEMS Microbiol. Ecol."},{"key":"B22","doi-asserted-by":"publisher","first-page":"930","DOI":"10.1111\/j.1365-2958.2010.07259.x","article-title":"H-NOX regulation of c-di-GMP metabolism and biofilm formation in Legionella pneumophila","volume":"77","author":"Carlson","year":"2010","journal-title":"Mol. Microbiol."},{"key":"B23","doi-asserted-by":"publisher","first-page":"5188","DOI":"10.1073\/pnas.89.11.5188","article-title":"Legionella pneumophila mip gene potentiates intracellular infection of protozoa and human macrophages","volume":"89","author":"Cianciotto","year":"1992","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B24","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1016\/j.mib.2017.01.002","article-title":"The ins and outs of cyclic di-GMP signaling in Vibrio cholerae","volume":"36","author":"Conner","year":"2017","journal-title":"Curr. Opin. Microbiol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1016\/j.jhin.2009.07.005","article-title":"Resistance of Legionella pneumophila serotype 1 biofilms to chlorine-based disinfection","volume":"74","author":"Cooper","year":"2010","journal-title":"J. Hosp. Infect."},{"key":"B26","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1007\/BF01569816","article-title":"Overview of microbial biofilms","volume":"15","author":"Costerton","year":"1995","journal-title":"J. Ind. Microbiol."},{"key":"B27","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1146\/annurev.mi.41.100187.002251","article-title":"Bacterial biofilms in nature and disease","volume":"41","author":"Costerton","year":"1987","journal-title":"Annu. Rev. Microbiol."},{"key":"B28","doi-asserted-by":"publisher","first-page":"1413","DOI":"10.1016\/j.bbrc.2005.04.060","article-title":"Legionella pneumophila Philadelphia-1 tatB and tatC affect intracellular replication and biofilm formation","volume":"331","author":"De Buck","year":"2005","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B29","doi-asserted-by":"publisher","first-page":"557","DOI":"10.1111\/j.1462-2920.2009.02025.x","article-title":"Biofilms: the environmental playground of Legionella pneumophila","volume":"12","author":"Declerck","year":"2010","journal-title":"Environ. Microbiol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1016\/j.micres.2007.06.001","article-title":"Replication of Legionella pneumophila in biofilms of water distribution pipes","volume":"164","author":"Declerck","year":"2009","journal-title":"Microbiol. Res."},{"key":"B31","doi-asserted-by":"publisher","first-page":"7716","DOI":"10.1128\/JB.187.22.7716-7726.2005","article-title":"Evidence for acquisition of Legionella type IV secretion substrates via interdomain horizontal gene transfer","volume":"187","author":"de Felipe","year":"2005","journal-title":"J. Bacteriol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1080\/08927010500044286","article-title":"Legionella pneumophila associated with the protozoan Hartmannella vermiformis in a model multi-species biofilm has reduced susceptibility to disinfectants","volume":"21","author":"Donlan","year":"2005","journal-title":"Biofouling"},{"key":"B33","doi-asserted-by":"publisher","first-page":"2168","DOI":"10.1128\/IAI.01304-10","article-title":"Lcl of Legionella pneumophila is an immunogenic GAG binding adhesin that promotes interactions with lung epithelial cells and plays a crucial role in biofilm formation","volume":"79","author":"Duncan","year":"2011","journal-title":"Infect. Immun."},{"key":"B34","doi-asserted-by":"publisher","first-page":"1087","DOI":"10.1016\/j.watres.2010.10.025","article-title":"Efficiency of water disinfectants against Legionella pneumophila and Acanthamoeba","volume":"45","author":"Dupuy","year":"2011","journal-title":"Water Res."},{"key":"B35","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1128\/JCM.16.4.697-699.1982","article-title":"Comparative study of selective media for isolation of Legionella pneumophila from potable water","volume":"16","author":"Edelstein","year":"1982","journal-title":"J. Clin. Microbiol."},{"key":"B36","doi-asserted-by":"publisher","first-page":"6850","DOI":"10.1128\/AEM.00831-12","article-title":"Effects of disinfection on Legionella spp., eukarya, and biofilms in a hot water system","volume":"78","author":"Farhat","year":"2012","journal-title":"Appl. Environ. Microbiol."},{"key":"B37","doi-asserted-by":"publisher","first-page":"7025","DOI":"10.1128\/JB.184.24.7025-7041.2002","article-title":"Ultrastructural analysis of differentiation in Legionella pneumophila","volume":"184","author":"Faulkner","year":"2002","journal-title":"J. Bacteriol."},{"key":"B38","doi-asserted-by":"publisher","first-page":"506","DOI":"10.1128\/CMR.15.3.506-526.2002","article-title":"Legionella and Legionnaires' disease: 25 years of investigation","volume":"15","author":"Fields","year":"2002","journal-title":"Clin. Microbiol. Rev."},{"key":"B39","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.1056\/NEJM197712012972201","article-title":"Legionnaires' disease: description of an epidemic of pneumonia","volume":"297","author":"Fraser","year":"1977","journal-title":"N. Engl. J. Med."},{"key":"B40","doi-asserted-by":"publisher","first-page":"1267","DOI":"10.1111\/j.1462-2920.2007.01245.x","article-title":"Acanthamoeba polyphaga resuscitates viable non-culturable Legionella pneumophila after disinfection","volume":"9","author":"Garc\u00eda","year":"2007","journal-title":"Environ. Microbiol."},{"key":"B41","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1128\/JCM.11.3.286-291.1980","article-title":"Amino acid requirements of Legionella pneumophila","volume":"11","author":"George","year":"1980","journal-title":"J. Clin. Microbiol."},{"key":"B42","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1080\/08927010902802232","article-title":"Incorporation of natural uncultivable Legionella pneumophila into potable water biofilms provides a protective niche against chlorination stress","volume":"25","author":"Giao","year":"2009","journal-title":"Biofouling"},{"key":"B43","doi-asserted-by":"publisher","first-page":"532","DOI":"10.1007\/s00284-008-9237-2","article-title":"Effect of bacterial interference on biofilm development by Legionella pneumophila","volume":"57","author":"Guerrieri","year":"2008","journal-title":"Curr. Microbiol."},{"key":"B44","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1046\/j.1462-5822.2000.00044.x","article-title":"Dictyostelium discoideum: a new host model system for intracellular pathogens of the genus Legionella","volume":"2","author":"H\u00e4gele","year":"2000","journal-title":"Cell. Microbiol."},{"key":"B45","doi-asserted-by":"publisher","first-page":"660","DOI":"10.1002\/mbo3.271","article-title":"Identification of vacuoles containing extraintestinal differentiated forms of Legionella pneumophila in colonized Caenorhabditis elegans soil nematodes","volume":"4","author":"Hellinga","year":"2015","journal-title":"Microbiologyopen"},{"key":"B46","doi-asserted-by":"publisher","first-page":"1604","DOI":"10.1128\/IAI.70.3.1604-1608.2002","article-title":"Influence of the alternative sigma(28) factor on virulence and flagellum expression of Legionella pneumophila","volume":"70","author":"Heuner","year":"2002","journal-title":"Infect. Immun."},{"key":"B47","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/j.watres.2005.10.030","article-title":"Inactivation credit of UV radiation for viruses, bacteria and protozoan (oo)cysts in water: a review","volume":"40","author":"Hijnen","year":"2006","journal-title":"Water Res."},{"key":"B48","doi-asserted-by":"publisher","first-page":"286","DOI":"10.1111\/j.1758-2229.2011.00247.x","article-title":"Legionella spp. outdoors: colonization, communication and persistence","volume":"3","author":"Hilbi","year":"2011","journal-title":"Environ. Microbiol. Rep."},{"key":"B49","doi-asserted-by":"publisher","first-page":"30","DOI":"10.1099\/mic.0.2007\/008698-0","article-title":"Transcriptional profiling of Legionella pneumophila biofilm cells and the influence of iron on biofilm formation","volume":"154","author":"Hindr\u00e9","year":"2008","journal-title":"Microbiology"},{"key":"B50","doi-asserted-by":"publisher","first-page":"1319","DOI":"10.1084\/jem.158.4.1319","article-title":"Formation of a novel phagosome by the Legionnaires' disease bacterium (Legionella pneumophila) in human monocytes","volume":"158","author":"Horwitz","year":"1983","journal-title":"J. Exp. Med."},{"key":"B51","doi-asserted-by":"crossref","first-page":"3039","DOI":"10.1099\/00221287-144-11-3039","article-title":"Biofilm susceptibility to bacteriophage attack: the role of phage-borne polysaccharide depolymerase","volume":"144","author":"Hughes","year":"1998","journal-title":"Microbiology"},{"key":"B52","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1038\/nrmicro1967","article-title":"The Legionella pneumophila replication vacuole: making a cosy niche inside host cells","volume":"7","author":"Isberg","year":"2009","journal-title":"Nat. Rev. Microbiol."},{"key":"B53","doi-asserted-by":"publisher","first-page":"1333","DOI":"10.1002\/eji.201242835","article-title":"A bacterial protein promotes the recognition of the Legionella pneumophila vacuole by autophagy","volume":"43","author":"Khweek","year":"2013","journal-title":"Eur. J. Immunol."},{"key":"B54","doi-asserted-by":"publisher","first-page":"263","DOI":"10.7888\/juoeh.16.263","article-title":"Intracellular multiplication of Legionella pneumophila in Tetrahymena thermophila","volume":"16","author":"Kikuhara","year":"1994","journal-title":"J. UOEH"},{"key":"B55","doi-asserted-by":"publisher","first-page":"4433","DOI":"10.1016\/S0043-1354(02)00188-4","article-title":"Literature review\u2013efficacy of various disinfectants against Legionella in water systems","volume":"36","author":"Kim","year":"2002","journal-title":"Water Res."},{"key":"B56","doi-asserted-by":"publisher","first-page":"1934","DOI":"10.1099\/mic.0.026641-0","article-title":"Pseudomonas aeruginosa Las quorum sensing autoinducer suppresses growth and biofilm production in Legionella species","volume":"155","author":"Kimura","year":"2009","journal-title":"Microbiology"},{"key":"B57","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1016\/j.colsurfb.2006.04.018","article-title":"Polysaccharide differences between planktonic and biofilm-associated EPS from Pseudomonas fluorescens B52","volume":"52","author":"Kives","year":"2006","journal-title":"Colloids Surf. B Biointerfaces"},{"key":"B58","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1007\/s00248-007-9325-z","article-title":"Evidence for the presence of Legionella bacteriophages in environmental water samples","volume":"56","author":"Lammertyn","year":"2008","journal-title":"Microb. Ecol."},{"key":"B59","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1111\/j.1365-2672.2009.04208.x","article-title":"The role of biofilms and protozoa in Legionella pathogenesis: implications for drinking water","volume":"107","author":"Lau","year":"2009","journal-title":"J. Appl. Microbiol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"6235","DOI":"10.1128\/JB.00635-06","article-title":"Compensatory functions of two alkyl hydroperoxide reductases in the oxidative defense system of Legionella pneumophila","volume":"188","author":"LeBlanc","year":"2006","journal-title":"J. Bacteriol."},{"key":"B61","doi-asserted-by":"publisher","first-page":"2854","DOI":"10.1128\/AEM.02916-06","article-title":"Survival of Mycobacterium avium, Legionella pneumophila, Escherichia coli, and caliciviruses in drinking water-associated biofilms grown under high-shear turbulent flow","volume":"73","author":"Lehtola","year":"2007","journal-title":"Appl. Environ. Microbiol."},{"key":"B62","doi-asserted-by":"publisher","first-page":"371","DOI":"10.1038\/nrmicro3028","article-title":"Antimicrobial activity of metals: mechanisms, molecular targets and applications","volume":"11","author":"Lemire","year":"2013","journal-title":"Nat. Rev. Microbiol."},{"key":"B63","doi-asserted-by":"publisher","first-page":"e00316","DOI":"10.1128\/mBio.00316-10","article-title":"Cyclic diguanylate signaling proteins control intracellular growth of Legionella pneumophila","volume":"2","author":"Levi","year":"2011","journal-title":"MBio"},{"key":"B64","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1111\/j.1365-2672.2006.02970.x","article-title":"Effect of flow regimes on the presence of Legionella within the biofilm of a model plumbing system","volume":"101","author":"Liu","year":"2006","journal-title":"J. Appl. Microbiol."},{"key":"B65","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1016\/j.ijheh.2010.03.004","article-title":"Free-living amoebae: biological by-passes in water treatment","volume":"213","author":"Loret","year":"2010","journal-title":"Int. J. Hyg. Environ. Health"},{"key":"B66","doi-asserted-by":"publisher","first-page":"2177","DOI":"10.1084\/jem.20060766","article-title":"NF-kappaB translocation prevents host cell death after low-dose challenge by Legionella pneumophila","volume":"203","author":"Losick","year":"2006","journal-title":"J. Exp. Med."},{"key":"B67","doi-asserted-by":"publisher","first-page":"e46462","DOI":"10.1371\/journal.pone.0046462","article-title":"Essential roles and regulation of the Legionella pneumophila collagen-like adhesin during biofilm formation","volume":"7","author":"Mallegol","year":"2012","journal-title":"PLoS ONE"},{"key":"B68","doi-asserted-by":"publisher","first-page":"2885","DOI":"10.1128\/AEM.72.4.2885-2895.2006","article-title":"Planktonic replication is essential for biofilm formation by Legionella pneumophila in a complex medium under static and dynamic flow conditions","volume":"72","author":"Mampel","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"B69","doi-asserted-by":"publisher","first-page":"199","DOI":"10.21775\/cimb.025.199","article-title":"Role of Cyclic di-GMP in the bacterial virulence and evasion of the plant immunity","volume":"25","author":"Martinez-Gil","year":"2017","journal-title":"Curr. Issues Mol. Biol."},{"key":"B70","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1016\/S0092-8674(02)00829-2","article-title":"Parallel quorum sensing systems converge to regulate virulence in Vibrio cholerae","volume":"110","author":"Miller","year":"2002","journal-title":"Cell"},{"key":"B71","doi-asserted-by":"publisher","first-page":"5720","DOI":"10.1128\/IAI.73.9.5720-5734.2005","article-title":"Components of the Legionella pneumophila flagellar regulon contribute to multiple virulence traits, including lysosome avoidance and macrophage death","volume":"73","author":"Molofsky","year":"2005","journal-title":"Infect. Immun."},{"key":"B72","doi-asserted-by":"publisher","first-page":"3121","DOI":"10.1099\/00221287-147-11-3121","article-title":"Role of biofilms in the survival of Legionella pneumophila in a model potable-water system","volume":"147","author":"Murga","year":"2001","journal-title":"Microbiology"},{"key":"B73","doi-asserted-by":"publisher","first-page":"789","DOI":"10.1016\/j.chembiol.2005.05.007","article-title":"Iron salts perturb biofilm formation and disrupt existing biofilms of Pseudomonas aeruginosa","volume":"12","author":"Musk","year":"2005","journal-title":"Chem. Biol."},{"key":"B74","doi-asserted-by":"crossref","first-page":"1688","DOI":"10.1128\/AEM.64.5.1688-1693.1998","article-title":"Isolation of an amoeba naturally harboring a distinctive Legionella species","volume":"64","author":"Newsome","year":"1998","journal-title":"Appl. Environ. Microbiol."},{"key":"B75","doi-asserted-by":"publisher","first-page":"274","DOI":"10.1128\/CMR.00052-09","article-title":"Molecular pathogenesis of infections caused by Legionella pneumophila","volume":"23","author":"Newton","year":"2010","journal-title":"Clin. Microbiol. Rev."},{"key":"B76","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1146\/annurev-genet-102108-134304","article-title":"Bacterial quorum-sensing network architectures","volume":"43","author":"Ng","year":"2009","journal-title":"Annu. Rev. Genet."},{"key":"B77","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1146\/annurev.micro.54.1.49","article-title":"Biofilm formation as microbial development","volume":"54","author":"O'Toole","year":"2000","journal-title":"Annu. Rev. Microbiol."},{"key":"B78","doi-asserted-by":"publisher","first-page":"5702","DOI":"10.1128\/AEM.02982-05","article-title":"Biological filtration limits carbon availability and affects downstream biofilm formation and community structure","volume":"72","author":"Pang","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"B79","doi-asserted-by":"publisher","first-page":"935","DOI":"10.1080\/08927014.2016.1212988","article-title":"New insights into Legionella pneumophila biofilm regulation by c-di-GMP signaling","volume":"32","author":"P\u00e9castaings","year":"2016","journal-title":"Biofouling"},{"key":"B80","doi-asserted-by":"publisher","first-page":"809","DOI":"10.1080\/08927014.2010.520159","article-title":"Sessile Legionella pneumophila is able to grow on surfaces and generate structured monospecies biofilms","volume":"26","author":"P\u00e9castaings","year":"2010","journal-title":"Biofouling"},{"key":"B81","doi-asserted-by":"publisher","first-page":"1613","DOI":"10.1128\/AEM.72.2.1613-1622.2006","article-title":"Temperature-regulated formation of mycelial mat-like biofilms by Legionella pneumophila","volume":"72","author":"Piao","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"B82","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1264\/jsme2.ME16010","article-title":"Iron Availability modulates the persistence of Legionella pneumophila in complex biofilms","volume":"31","author":"Portier","year":"2016","journal-title":"Microb. Environ."},{"key":"B83","doi-asserted-by":"publisher","first-page":"1553","DOI":"10.1126\/science.1212868","article-title":"Host proteasomal degradation generates amino acids essential for intracellular bacterial growth","volume":"334","author":"Price","year":"2011","journal-title":"Science"},{"key":"B84","doi-asserted-by":"publisher","first-page":"1720","DOI":"10.1111\/j.1462-5822.2006.00795.x","article-title":"Intracellular innate resistance to bacterial pathogens","volume":"8","author":"Radtke","year":"2006","journal-title":"Cell. Microbiol."},{"key":"B85","doi-asserted-by":"publisher","first-page":"477","DOI":"10.3109\/17435390.2013.796537","article-title":"Discrete nanoparticles induce loss of Legionella pneumophila biofilms from surfaces","volume":"8","author":"Raftery","year":"2014","journal-title":"Nanotoxicology"},{"key":"B86","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1016\/j.ijmm.2016.05.012","article-title":"Legionella-protozoa-nematode interactions in aquatic biofilms and influence of Mip on Caenorhabditis elegans colonization","volume":"306","author":"Rasch","year":"2016","journal-title":"Int. J. Med. Microbiol."},{"key":"B87","doi-asserted-by":"crossref","first-page":"688","DOI":"10.1128\/JCM.13.4.688-695.1981","article-title":"Metal requirements of Legionella pneumophila","volume":"13","author":"Reeves","year":"1981","journal-title":"J. Clin. Microbiol."},{"key":"B88","doi-asserted-by":"crossref","first-page":"5701","DOI":"10.1128\/JB.181.18.5701-5710.1999","article-title":"Role of the alkyl hydroperoxide reductase (ahpCF) gene in oxidative stress defense of the obligate Anaerobe bacteroides fragilis","volume":"181","author":"Rocha","year":"1999","journal-title":"J. Bacteriol."},{"key":"B89","doi-asserted-by":"crossref","first-page":"1842","DOI":"10.1128\/AEM.60.6.1842-1851.1994","article-title":"Influence of plumbing materials on biofilm formation and growth of Legionella pneumophila in potable water systems","volume":"60","author":"Rogers","year":"1994","journal-title":"Appl. Environ. Microbiol."},{"key":"B90","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1128\/MMBR.00043-12","article-title":"Cyclic di-GMP: the first 25 years of a universal bacterial second messenger","volume":"77","author":"R\u00f6mling","year":"2013","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"B91","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1136\/jcp.33.12.1179","article-title":"Preliminary report on the pathogenicity of Legionella pneumophila for freshwater and soil amoebae","volume":"33","author":"Rowbotham","year":"1980","journal-title":"J. Clin. Pathol."},{"key":"B92","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1016\/S0140-6736(81)90141-0","article-title":"Pontiac fever, amoebae, and legionellae","volume":"1","author":"Rowbotham","year":"1981","journal-title":"Lancet"},{"key":"B93","doi-asserted-by":"publisher","first-page":"946","DOI":"10.1038\/nrmicro1046","article-title":"Iron and microbial infection","volume":"2","author":"Schaible","year":"2004","journal-title":"Nat. Rev. Microbiol."},{"key":"B94","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1007\/978-3-319-41171-2_8","article-title":"Inflammasome recognition and regulation of the Legionella Flagellum","volume":"397","author":"Schell","year":"2016","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"B95","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1046\/j.1365-2672.2003.02019.x","article-title":"Formation of natural biofilms during chlorine dioxide and u.v. disinfection in a public drinking water distribution system","volume":"95","author":"Schwartz","year":"2003","journal-title":"J. Appl. Microbiol."},{"key":"B96","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1016\/S1074-5521(02)00198-9","article-title":"Molecular interactions in biofilms","volume":"9","author":"Shirtliff","year":"2002","journal-title":"Chem. Biol."},{"key":"B97","doi-asserted-by":"publisher","first-page":"1995","DOI":"10.1021\/es203933h","article-title":"Monitoring bacterial twitter: does quorum sensing determine the behavior of water and wastewater treatment biofilms?","volume":"46","author":"Shrout","year":"2012","journal-title":"Environ. Sci. Technol."},{"key":"B98","doi-asserted-by":"publisher","first-page":"1123","DOI":"10.1111\/j.1365-2958.2004.04206.x","article-title":"GGDEF and EAL domains inversely regulate cyclic di-GMP levels and transition from sessility to motility","volume":"53","author":"Simm","year":"2004","journal-title":"Mol. Microbiol."},{"key":"B99","doi-asserted-by":"publisher","first-page":"18113","DOI":"10.1074\/jbc.M801929200","article-title":"The Legionella autoinducer synthase LqsA produces an alpha-hydroxyketone signaling molecule","volume":"283","author":"Spirig","year":"2008","journal-title":"J. Biol. Chem."},{"key":"B100","doi-asserted-by":"crossref","first-page":"2047","DOI":"10.1128\/AEM.63.5.2047-2053.1997","article-title":"Resuscitation of viable but nonculturable Legionella pneumophila Philadelphia JR32 by Acanthamoeba castellanii","volume":"63","author":"Steinert","year":"1997","journal-title":"Appl. Environ. Microbiol."},{"key":"B101","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1111\/j.1574-6976.2002.tb00607.x","article-title":"Legionella pneumophila: an aquatic microbe goes astray","volume":"26","author":"Steinert","year":"2002","journal-title":"FEMS Microbiol. Rev."},{"key":"B102","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1016\/S0934-8840(98)80005-4","article-title":"Regrowth of Legionella pneumophila in a heat-disinfected plumbing system","volume":"288","author":"Steinert","year":"1998","journal-title":"Zentralbl. Bakteriol."},{"key":"B103","doi-asserted-by":"publisher","first-page":"e50560","DOI":"10.1371\/journal.pone.0050560","article-title":"Legionella pneumophila persists within biofilms formed by Klebsiella pneumoniae, Flavobacterium sp., and Pseudomonas fluorescens under dynamic flow conditions","volume":"7","author":"Stewart","year":"2012","journal-title":"PLoS ONE"},{"key":"B104","doi-asserted-by":"publisher","first-page":"656","DOI":"10.1080\/00365540410020785","article-title":"The efficacy of heat and chlorine treatment against thermotolerant Acanthamoebae and Legionellae","volume":"36","author":"Storey","year":"2004","journal-title":"Scand. J. Infect. Dis."},{"key":"B105","doi-asserted-by":"publisher","first-page":"2047","DOI":"10.1016\/j.actbio.2012.03.002","article-title":"Magnetic targeting of surface-modified superparamagnetic iron oxide nanoparticles yields antibacterial efficacy against biofilms of gentamicin-resistant staphylococci","volume":"8","author":"Subbiahdoss","year":"2012","journal-title":"Acta Biomater."},{"key":"B106","doi-asserted-by":"publisher","first-page":"222","DOI":"10.1016\/S0966-842X(01)02012-1","article-title":"The biofilm matrix\u2013an immobilized but dynamic microbial environment","volume":"9","author":"Sutherland","year":"2001","journal-title":"Trends Microbiol."},{"key":"B107","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1146\/annurev.micro.61.080706.093426","article-title":"Roles of cyclic diguanylate in the regulation of bacterial pathogenesis","volume":"61","author":"Tamayo","year":"2007","journal-title":"Annu. Rev. Microbiol."},{"key":"B108","doi-asserted-by":"publisher","first-page":"3016","DOI":"10.1002\/smll.201200575","article-title":"Superparamagnetic iron oxide nanoparticles (SPION) for the treatment of antibiotic-resistant biofilms","volume":"8","author":"Taylor","year":"2012","journal-title":"Small"},{"key":"B109","doi-asserted-by":"publisher","first-page":"538","DOI":"10.1007\/s00248-009-9514-z","article-title":"Legionella, protozoa, and biofilms: interactions within complex microbial systems","volume":"58","author":"Taylor","year":"2009","journal-title":"Microb. Ecol."},{"key":"B110","doi-asserted-by":"publisher","first-page":"4323","DOI":"10.1128\/AEM.00070-06","article-title":"Necrotrophic growth of Legionella pneumophila","volume":"72","author":"Temmerman","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"B111","doi-asserted-by":"publisher","first-page":"7532","DOI":"10.1128\/JB.01002-08","article-title":"Synergistic contribution of the Legionella pneumophila lqs genes to pathogen-host interactions","volume":"190","author":"Tiaden","year":"2008","journal-title":"J. Bacteriol."},{"key":"B112","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.1111\/j.1462-2920.2010.02167.x","article-title":"The autoinducer synthase LqsA and putative sensor kinase LqsS regulate phagocyte interactions, extracellular filaments and a genomic island of Legionella pneumophila","volume":"12","author":"Tiaden","year":"2010","journal-title":"Environ. Microbiol."},{"key":"B113","doi-asserted-by":"publisher","first-page":"2903","DOI":"10.1111\/j.1462-5822.2007.01005.x","article-title":"The Legionella pneumophila response regulator LqsR promotes host cell interactions as an element of the virulence regulatory network controlled by RpoS and LetA","volume":"9","author":"Tiaden","year":"2007","journal-title":"Cell. Microbiol."},{"key":"B114","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1128\/AEM.39.2.456-459.1980","article-title":"Growth of Legionella pneumophila in association with blue-green algae (cyanobacteria)","volume":"39","author":"Tison","year":"1980","journal-title":"Appl. Environ. Microbiol."},{"key":"B115","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1007\/s10661-006-9519-8","article-title":"Monitoring of biofilm-associated Legionella pneumophila on different substrata in model cooling tower system","volume":"125","author":"T\u00fcretgen","year":"2007","journal-title":"Environ. Monit. Assess."},{"key":"B116","doi-asserted-by":"crossref","first-page":"954","DOI":"10.1128\/AEM.44.4.954-959.1982","article-title":"Cocultivation of Legionella pneumophila and free-living amoebae","volume":"44","author":"Tyndall","year":"1982","journal-title":"Appl. Environ. Microbiol."},{"key":"B117","doi-asserted-by":"publisher","first-page":"12547","DOI":"10.1074\/jbc.R115.711507","article-title":"Biofilms and Cyclic di-GMP (c-di-GMP) signaling: lessons from Pseudomonas aeruginosa and Other bacteria","volume":"291","author":"Valentini","year":"2016","journal-title":"J. Biol. Chem."},{"key":"B118","doi-asserted-by":"publisher","first-page":"7144","DOI":"10.1128\/AEM.00926-10","article-title":"Detection of protozoan hosts for Legionella pneumophila in engineered water systems by using a biofilm batch test","volume":"76","author":"Valster","year":"2010","journal-title":"Appl. Environ. Microbiol."},{"key":"B119","doi-asserted-by":"publisher","first-page":"2789","DOI":"10.1016\/j.watres.2005.04.075","article-title":"Biofilm formation and multiplication of Legionella in a model warm water system with pipes of copper, stainless steel and cross-linked polyethylene","volume":"39","author":"van der Kooij","year":"2005","journal-title":"Water Res."},{"key":"B120","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1111\/j.1574-6941.2006.00181.x","article-title":"Introduction of a boost of Legionella pneumophila into a stagnant-water model by heat treatment","volume":"58","author":"Vervaeren","year":"2006","journal-title":"FEMS Microbiol. Ecol."},{"key":"B121","doi-asserted-by":"crossref","first-page":"1447","DOI":"10.1128\/AEM.46.6.1447-1449.1983","article-title":"Satellite growth of Legionella pneumophila with an environmental isolate of Flavobacterium breve","volume":"46","author":"Wadowsky","year":"1983","journal-title":"Appl. Environ. Microbiol."},{"key":"B122","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1111\/j.1462-5822.2006.00802.x","article-title":"Collagen binding protein Mip enables Legionella pneumophila to transmigrate through a barrier of NCI-H292 lung epithelial cells and extracellular matrix","volume":"9","author":"Wagner","year":"2007","journal-title":"Cell. Microbiol."},{"key":"B123","doi-asserted-by":"publisher","first-page":"2675","DOI":"10.1128\/JB.182.10.2675-2679.2000","article-title":"Biofilm, city of microbes","volume":"182","author":"Watnick","year":"2000","journal-title":"J. Bacteriol."},{"key":"B124","doi-asserted-by":"publisher","first-page":"1528","DOI":"10.1111\/j.1365-2958.2005.04490.x","article-title":"Intracellular multiplication of Legionella pneumophila depends on host cell amino acid transporter SLC1A5","volume":"55","author":"Wieland","year":"2005","journal-title":"Mol. Microbiol."},{"key":"B125","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1111\/j.1365-2672.1991.tb03828.x","article-title":"Decreased biocide susceptibility of adherent Legionella pneumophila","volume":"71","author":"Wright","year":"1991","journal-title":"J. Appl. Bacteriol."},{"key":"B126","doi-asserted-by":"publisher","first-page":"e23500","DOI":"10.1371\/journal.pone.0023500","article-title":"Isolation of genes involved in biofilm formation of a Klebsiella pneumoniae strain causing pyogenic liver abscess","volume":"6","author":"Wu","year":"2011","journal-title":"PLoS ONE"},{"key":"B127","doi-asserted-by":"publisher","first-page":"3129","DOI":"10.1073\/pnas.052694299","article-title":"Quorum-sensing regulators control virulence gene expression in Vibrio cholerae","volume":"99","author":"Zhu","year":"2002","journal-title":"Proc. Natl. Acad. Sci. U.S.A."}],"container-title":["Frontiers in Cellular and Infection Microbiology"],"original-title":[],"link":[{"URL":"http:\/\/journal.frontiersin.org\/article\/10.3389\/fcimb.2018.00038\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,10,28]],"date-time":"2020-10-28T19:11:04Z","timestamp":1603912264000},"score":1,"resource":{"primary":{"URL":"http:\/\/journal.frontiersin.org\/article\/10.3389\/fcimb.2018.00038\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,2,27]]},"references-count":127,"alternative-id":["10.3389\/fcimb.2018.00038"],"URL":"https:\/\/doi.org\/10.3389\/fcimb.2018.00038","relation":{},"ISSN":["2235-2988"],"issn-type":[{"value":"2235-2988","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,2,27]]},"article-number":"38"}}