{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T16:25:58Z","timestamp":1791822358508,"version":"4.3.4"},"reference-count":86,"publisher":"MDPI AG","issue":"4","license":[{"start":{"date-parts":[[2018,4,10]],"date-time":"2018-04-10T00:00:00Z","timestamp":1523318400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Toxins"],"abstract":"<jats:p>Cyclomodulins are bacterial toxins that interfere with the eukaryotic cell cycle. A new cyclomodulin called colibactin, which is synthetized by the pks genomic island, was discovered in 2006. Despite many efforts, colibactin has not yet been purified, and its structure remains elusive. Interestingly, the pks island is found in members of the family Enterobacteriaceae (mainly Escherichia coli and Klebsiella pneumoniae) isolated from different origins, including from intestinal microbiota, septicaemia, newborn meningitis, and urinary tract infections. Colibactin-producing bacteria induce chromosomal instability and DNA damage in eukaryotic cells, which leads to senescence of epithelial cells and apoptosis of immune cells. The pks island is mainly observed in B2 phylogroup E. coli strains, which include extra-intestinal pathogenic E. coli strains, and pks E. coli are over-represented in biopsies isolated from colorectal cancer. In addition, pks E. coli bacteria increase the number of tumours in diverse colorectal cancer mouse models. Thus, colibactin could have a major impact on human health. In the present review, we will focus on the biological effects of colibactin, the distribution of the pks island, and summarize what is currently known about its synthesis and its structure.<\/jats:p>","DOI":"10.3390\/toxins10040151","type":"journal-article","created":{"date-parts":[[2018,4,11]],"date-time":"2018-04-11T12:16:50Z","timestamp":1523449010000},"page":"151","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":233,"title":["Colibactin: More Than a New Bacterial Toxin"],"prefix":"10.3390","volume":"10","author":[{"given":"Tiphanie","family":"Fa\u00efs","sequence":"first","affiliation":[{"name":"Universit\u00e9 Clermont Auvergne, Inserm U1071, M2iSH, USC-INRA 2018, F-63000 Clermont-Ferrand, France"},{"name":"CHU Clermont-Ferrand, Laboratoire de Bact\u00e9riologie, Centre de Biologie, F-63003 Clermont-Ferrand, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9579-0660","authenticated-orcid":false,"given":"Julien","family":"Delmas","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Clermont Auvergne, Inserm U1071, M2iSH, USC-INRA 2018, F-63000 Clermont-Ferrand, France"},{"name":"CHU Clermont-Ferrand, Laboratoire de Bact\u00e9riologie, Centre de Biologie, F-63003 Clermont-Ferrand, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9465-7844","authenticated-orcid":false,"given":"Nicolas","family":"Barnich","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Clermont Auvergne, Inserm U1071, M2iSH, USC-INRA 2018, F-63000 Clermont-Ferrand, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard","family":"Bonnet","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Clermont Auvergne, Inserm U1071, M2iSH, USC-INRA 2018, F-63000 Clermont-Ferrand, France"},{"name":"CHU Clermont-Ferrand, Laboratoire de Bact\u00e9riologie, Centre de Biologie, F-63003 Clermont-Ferrand, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3433-870X","authenticated-orcid":false,"given":"Guillaume","family":"Dalmasso","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Clermont Auvergne, Inserm U1071, M2iSH, USC-INRA 2018, F-63000 Clermont-Ferrand, France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2018,4,10]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.tim.2005.01.002","article-title":"Cyclomodulins: bacterial effectors that modulate the eukaryotic cell cycle","volume":"13","author":"Taieb","year":"2005","journal-title":"Trends Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"848","DOI":"10.1126\/science.1127059","article-title":"Escherichia coli induces DNA double-strand breaks in eukaryotic cells","volume":"313","author":"Homburg","year":"2006","journal-title":"Science"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"11537","DOI":"10.1073\/pnas.1001261107","article-title":"Escherichia coli induces DNA damage in vivo and triggers genomic instability in mammalian cells","volume":"107","author":"Petit","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2122","DOI":"10.1128\/JCM.02365-09","article-title":"Cyclomodulins in urosepsis strains of Escherichia coli","volume":"48","author":"Dubois","year":"2010","journal-title":"J. Clin. Microbiol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"3906","DOI":"10.1128\/JCM.00949-08","article-title":"Molecular epidemiology and phylogenetic distribution of the Escherichia coli pks genomic island","volume":"46","author":"Johnson","year":"2008","journal-title":"J. Clin. Microbiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"4696","DOI":"10.1128\/IAI.00522-09","article-title":"Genetic structure and distribution of the colibactin genomic island among members of the family Enterobacteriaceae","volume":"77","author":"Putze","year":"2009","journal-title":"Infect. Immun."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1016\/j.micpath.2012.05.011","article-title":"Escherichia coli strains with the capacity for long-term persistence in the bowel microbiota carry the potentially genotoxic pks island","volume":"53","author":"Nowrouzian","year":"2012","journal-title":"Microb. Pathog."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"313","DOI":"10.4161\/gmic.28932","article-title":"Maternally acquired genotoxic Escherichia coli alters offspring\u2019s intestinal homeostasis","volume":"5","author":"Payros","year":"2014","journal-title":"Gut Microbes"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1016\/j.ijmm.2017.02.003","article-title":"Human Escherichia coli isolates from hemocultures: Septicemia linked to urogenital tract infections is caused by isolates harboring more virulence genes than bacteraemia linked to other conditions","volume":"307","author":"Vrba","year":"2017","journal-title":"Int. J. Med. Microbiol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"3704","DOI":"10.1128\/IAI.00716-15","article-title":"The Genotoxin Colibactin Is a Determinant of Virulence in Escherichia coli K1 Experimental Neonatal Systemic Infection","volume":"83","author":"McCarthy","year":"2015","journal-title":"Infect. Immun."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1420","DOI":"10.1016\/j.urology.2010.12.059","article-title":"Acute Escherichia coli prostatitis in previously health young men: bacterial virulence factors, antimicrobial resistance, and clinical outcomes","volume":"77","author":"Krieger","year":"2011","journal-title":"Urology"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Buc, E., Dubois, D., Sauvanet, P., Raisch, J., Delmas, J., Darfeuille-Michaud, A., Pezet, D., and Bonnet, R. (2013). High prevalence of mucosa-associated E. coli producing cyclomodulin and genotoxin in colon cancer. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0056964"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Ekl\u00f6f, V., L\u00f6fgren-Burstr\u00f6m, A., Zingmark, C., Edin, S., Larsson, P., Karling, P., Alexeyev, O., Ruteg\u00e5rd, J., Wikberg, M.L., and Palmqvist, R. (2017). Cancer-associated fecal microbial markers in colorectal cancer detection. Int. J. Cancer.","DOI":"10.1002\/ijc.31011"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1126\/science.1224820","article-title":"Intestinal inflammation targets cancer-inducing activity of the microbiota","volume":"338","author":"Arthur","year":"2012","journal-title":"Science"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1136\/gutjnl-2013-304739","article-title":"Colonic mucosa-associated diffusely adherent afaC+ Escherichia coli expressing lpfA and pks are increased in inflammatory bowel disease and colon cancer","volume":"63","author":"Friswell","year":"2014","journal-title":"Gut"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"859","DOI":"10.1158\/1078-0432.CCR-13-1343","article-title":"Colonization of the human gut by E. coli and colorectal cancer risk","volume":"20","author":"Bonnet","year":"2014","journal-title":"Clin. Cancer Res."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1932","DOI":"10.1136\/gutjnl-2013-305257","article-title":"Bacterial genotoxin colibactin promotes colon tumour growth by inducing a senescence-associated secretory phenotype","volume":"63","author":"Cougnoux","year":"2014","journal-title":"Gut"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"2620","DOI":"10.1158\/0008-5472.CAN-16-3472","article-title":"Locoregional Effects of Microbiota in a Preclinical Model of Colon Carcinogenesis","volume":"77","author":"Tomkovich","year":"2017","journal-title":"Cancer Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"3468","DOI":"10.1021\/cr0503097","article-title":"Assembly-line enzymology for polyketide and nonribosomal Peptide antibiotics: logic, machinery, and mechanisms","volume":"106","author":"Fischbach","year":"2006","journal-title":"Chem. Rev."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"501","DOI":"10.4161\/gmic.21737","article-title":"Genotoxicity of Escherichia coli Nissle 1917 strain cannot be dissociated from its probiotic activity","volume":"3","author":"Olier","year":"2012","journal-title":"Gut Microbes"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"6986","DOI":"10.1128\/AAC.00130-16","article-title":"Antibiotic Activity of Escherichia coli against Multiresistant Staphylococcus aureus","volume":"60","author":"Cougnoux","year":"2016","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"9244","DOI":"10.1021\/ja503450q","article-title":"Comparative metabolomics and structural characterizations illuminate colibactin pathway-dependent small molecules","volume":"136","author":"Vizcaino","year":"2014","journal-title":"J. Am. Chem. Soc."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1314","DOI":"10.1038\/s41467-017-01403-9","article-title":"Identification of an analgesic lipopeptide produced by the probiotic Escherichia coli strain Nissle 1917","volume":"8","author":"Pujo","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1038\/nrmicro3432","article-title":"Urinary tract infections: epidemiology, mechanisms of infection and treatment options","volume":"13","author":"Walker","year":"2015","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1016\/S0140-6736(75)92496-4","article-title":"Epidemiology of Escherichia coli K1 in healthy and diseased newborns","volume":"1","author":"Sarff","year":"1975","journal-title":"Lancet"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1111\/1758-2229.12019","article-title":"The Clermont Escherichia coli phylo-typing method revisited: improvement of specificity and detection of new phylo-groups","volume":"5","author":"Clermont","year":"2013","journal-title":"Environ. Microbiol. Rep."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1085","DOI":"10.1093\/molbev\/msh118","article-title":"A specific genetic background is required for acquisition and expression of virulence factors in Escherichia coli","volume":"21","author":"Clermont","year":"2004","journal-title":"Mol. Biol. Evol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"6560","DOI":"10.3748\/wjg.v20.i21.6560","article-title":"Colon cancer-associated B2 Escherichia coli colonize gut mucosa and promote cell proliferation","volume":"20","author":"Raisch","year":"2014","journal-title":"World J. Gastroenterol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1038\/nrmicro2298","article-title":"The population genetics of commensal Escherichia coli","volume":"8","author":"Tenaillon","year":"2010","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Lai, Y.-C., Lin, A.-C., Chiang, M.-K., Dai, Y.-H., Hsu, C.-C., Lu, M.-C., Liau, C.-Y., and Chen, Y.-T. (2014). Genotoxic Klebsiella pneumoniae in Taiwan. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0096292"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2075","DOI":"10.1086\/315488","article-title":"A 5-year study of the seroepidemiology of Klebsiella pneumoniae: high prevalence of capsular serotype K1 in Taiwan and implication for vaccine efficacy","volume":"181","author":"Fung","year":"2000","journal-title":"J. Infect. Dis."},{"key":"ref_32","first-page":"203","article-title":"Klebsiella pneumoniae infection on a rehabilitation unit: comparison of epidemiologic typing methods","volume":"14","author":"Thompson","year":"1993","journal-title":"Infect. Control Hosp. Epidemiol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"43120","DOI":"10.1038\/srep43120","article-title":"Prevalence and characteristics of pks genotoxin gene cluster-positive clinical Klebsiella pneumoniae isolates in Taiwan","volume":"7","author":"Chen","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1194","DOI":"10.1002\/cbic.201300208","article-title":"In vivo evidence for a prodrug activation mechanism during colibactin maturation","volume":"14","author":"Bian","year":"2013","journal-title":"Chembiochem"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"3359","DOI":"10.1021\/ja312154m","article-title":"A prodrug resistance mechanism is involved in colibactin biosynthesis and cytotoxicity","volume":"135","author":"Brotherton","year":"2013","journal-title":"J. Am. Chem. Soc."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Martin, P., Marcq, I., Magistro, G., Penary, M., Garcie, C., Payros, D., Boury, M., Olier, M., Nougayr\u00e8de, J.-P., and Audebert, M. (2013). Interplay between siderophores and colibactin genotoxin biosynthetic pathways in Escherichia coli. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003437"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1021\/acschembio.6b00014","article-title":"Characterization of Polyketide Synthase Machinery from the pks Island Facilitates Isolation of a Candidate Precolibactin","volume":"11","author":"Zha","year":"2016","journal-title":"ACS Chem. Biol."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Zha, L., Jiang, Y., Henke, M.T., Wilson, M.R., Wang, J.X., Kelleher, N.L., and Balskus, E.P. (2017). Colibactin assembly line enzymes use S-adenosylmethionine to build a cyclopropane ring. Nat. Chem. Biol.","DOI":"10.1038\/nchembio.2448"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"4195","DOI":"10.1021\/jacs.7b00659","article-title":"Domain-Targeted Metabolomics Delineates the Heterocycle Assembly Steps of Colibactin Biosynthesis","volume":"139","author":"Trautman","year":"2017","journal-title":"J. Am. Chem. Soc."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"5426","DOI":"10.1021\/jacs.6b02276","article-title":"Convergent and Modular Synthesis of Candidate Precolibactins. Structural Revision of Precolibactin A","volume":"138","author":"Healy","year":"2016","journal-title":"J. Am. Chem. Soc."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"13138","DOI":"10.1039\/C5CC02718G","article-title":"Colibactin biosynthesis and biological activity depend on the rare aminomalonyl polyketide precursor","volume":"51","author":"Brachmann","year":"2015","journal-title":"Chem. Commun. (Camb.)"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1038\/nchembio.2157","article-title":"Divergent biosynthesis yields a cytotoxic aminomalonate-containing precolibactin","volume":"12","author":"Li","year":"2016","journal-title":"Nat. Chem. Biol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1038\/nchem.2221","article-title":"The colibactin warhead crosslinks DNA","volume":"7","author":"Vizcaino","year":"2015","journal-title":"Nat. Chem."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Guntaka, N.S., Healy, A.R., Crawford, J.M., Herzon, S.B., and Bruner, S.D. (2017). Structure and Functional Analysis of ClbQ, an Unusual Intermediate-Releasing Thioesterase from the Colibactin Biosynthetic Pathway. ACS Chem. Biol.","DOI":"10.1021\/acschembio.7b00479"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"15009","DOI":"10.1038\/nmicrobiol.2015.9","article-title":"MATE transport of the E. coli-derived genotoxin colibactin","volume":"1","author":"Mousa","year":"2016","journal-title":"Nat. Microbiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1715","DOI":"10.1002\/cbic.201500239","article-title":"Critical Intermediates Reveal New Biosynthetic Events in the Enigmatic Colibactin Pathway","volume":"16","author":"Li","year":"2015","journal-title":"Chembiochem"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.jmb.2012.09.017","article-title":"Analysis of structure-function relationships in the colibactin-maturating enzyme ClbP","volume":"424","author":"Cougnoux","year":"2012","journal-title":"J. Mol. Biol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"35562","DOI":"10.1074\/jbc.M111.221960","article-title":"ClbP is a prototype of a peptidase subgroup involved in biosynthesis of nonribosomal peptides","volume":"286","author":"Dubois","year":"2011","journal-title":"J. Biol. Chem."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1111\/mmi.13272","article-title":"Escherichia coli\u2005ClbS is a colibactin resistance protein","volume":"99","author":"Dubois","year":"2016","journal-title":"Mol. Microbiol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Tripathi, P., Shine, E.E., Healy, A.R., Kim, C.S., Herzon, S.B., Bruner, S.D., and Crawford, J.M. (2017). ClbS is a cyclopropane hydrolase that confers colibactin resistance. J. Am. Chem. Soc.","DOI":"10.1021\/jacs.7b09971"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"3154","DOI":"10.1039\/C5SC00101C","article-title":"Two more pieces of the colibactin genotoxin puzzle from Escherichia coli show incorporation of an unusual 1-aminocyclopropanecarboxylic acid moiety","volume":"6","author":"Bian","year":"2015","journal-title":"Chem. Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1545","DOI":"10.1021\/acs.orglett.5b00432","article-title":"Isolation of a metabolite from the pks island provides insights into colibactin biosynthesis and activity","volume":"17","author":"Brotherton","year":"2015","journal-title":"Org. Lett."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"3642","DOI":"10.1073\/pnas.92.9.3642","article-title":"CC-1065 and the duocarmycins: unraveling the keys to a new class of naturally derived DNA alkylating agents","volume":"92","author":"Boger","year":"1995","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"3578","DOI":"10.1021\/cr2001367","article-title":"Chemistry and biology of acylfulvenes: sesquiterpene-derived antitumor agents","volume":"112","author":"Tanasova","year":"2012","journal-title":"Chem. Rev."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Bossuet-Greif, N., Vignard, J., Taieb, F., Mirey, G., Dubois, D., Petit, C., Oswald, E., and Nougayr\u00e8de, J.-P. (2018). The Colibactin Genotoxin Generates DNA Interstrand Cross-Links in Infected Cells. MBio, 9.","DOI":"10.1128\/mBio.02393-17"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1038\/nrc3088","article-title":"DNA interstrand crosslink repair and cancer","volume":"11","author":"Deans","year":"2011","journal-title":"Nat. Rev. Cancer"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1039\/a806930a","article-title":"Thiazole and oxazole peptides: biosynthesis and molecular machinery","volume":"16","author":"Roy","year":"1999","journal-title":"Nat. Prod. Rep."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1021\/bi00568a015","article-title":"Binding of bleomycin to DNA: intercalation of the bithiazole rings","volume":"18","author":"Povirk","year":"1979","journal-title":"Biochemistry"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"15563","DOI":"10.1021\/jacs.6b10354","article-title":"A Mechanistic Model for Colibactin-Induced Genotoxicity","volume":"138","author":"Healy","year":"2016","journal-title":"J. Am. Chem. Soc."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1111\/j.1574-6968.2007.00889.x","article-title":"Expression analysis of the colibactin gene cluster coding for a novel polyketide in Escherichia coli","volume":"275","author":"Homburg","year":"2007","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"3358","DOI":"10.1128\/IAI.00659-16","article-title":"Iron Homeostasis Regulates the Genotoxicity of Escherichia coli That Produces Colibactin","volume":"84","author":"Tronnet","year":"2016","journal-title":"Infect. Immun."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Tronnet, S., Garcie, C., Brachmann, A.O., Piel, J., Oswald, E., and Martin, P. (2017). High iron supply inhibits the synthesis of the genotoxin colibactin by pathogenic Escherichia coli through a non-canonical Fur\/RyhB-mediated pathway. Pathog. Dis.","DOI":"10.1093\/femspd\/ftx066"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"916","DOI":"10.1093\/infdis\/jiw294","article-title":"The Bacterial Stress-Responsive Hsp90 Chaperone (HtpG) Is Required for the Production of the Genotoxin Colibactin and the Siderophore Yersiniabactin in Escherichia coli","volume":"214","author":"Garcie","year":"2016","journal-title":"J. Infect. Dis."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"523","DOI":"10.1007\/s00438-008-0330-9","article-title":"Transcriptomics and adaptive genomics of the asymptomatic bacteriuria Escherichia coli strain 83972","volume":"279","author":"Hancock","year":"2008","journal-title":"Mol. Genet. Genom."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"4724","DOI":"10.1038\/ncomms5724","article-title":"Microbial genomic analysis reveals the essential role of inflammation in bacteria-induced colorectal cancer","volume":"5","author":"Arthur","year":"2014","journal-title":"Nat. Commun."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Payros, D., Dobrindt, U., Martin, P., Secher, T., Bracarense, A.P.F.L., Boury, M., Laffitte, J., Pinton, P., Oswald, E., and Oswald, I.P. (2017). The Food Contaminant Deoxynivalenol Exacerbates the Genotoxicity of Gut Microbiota. MBio, 8.","DOI":"10.1128\/mBio.00007-17"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1136\/gutjnl-2014-307241","article-title":"Small-molecule inhibitors prevent the genotoxic and protumoural effects induced by colibactin-producing bacteria","volume":"65","author":"Cougnoux","year":"2016","journal-title":"Gut"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Reuter, C., Alzheimer, M., Walles, H., and Oelschlaeger, T.A. (2017). An Adherent Mucus Layer Attenuates the Genotoxic Effect of Colibactin. Cell. Microbiol.","DOI":"10.1111\/cmi.12812"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"2420","DOI":"10.1128\/IAI.00064-15","article-title":"Oral tolerance failure upon neonatal gut colonization with Escherichia coli producing the genotoxin colibactin","volume":"83","author":"Secher","year":"2015","journal-title":"Infect. Immun."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"501","DOI":"10.3748\/wjg.v22.i2.501","article-title":"Gut microbiota imbalance and colorectal cancer","volume":"22","author":"Raisch","year":"2016","journal-title":"World J. Gastroenterol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Lucas, C., Barnich, N., and Nguyen, H.T.T. (2017). Microbiota, Inflammation and Colorectal Cancer. Int. J. Mol. Sci., 18.","DOI":"10.3390\/ijms18061310"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1051\/medsci\/20163202011","article-title":"[How some commensal bacteria would exacerbate colorectal carcinogenesis?]","volume":"32","author":"Raisch","year":"2016","journal-title":"Med. Sci. (Paris)"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/S0140-6736(05)17706-X","article-title":"Colorectal cancer","volume":"365","author":"Weitz","year":"2005","journal-title":"Lancet"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/S0016-5085(98)70194-5","article-title":"Association between intraepithelial Escherichia coli and colorectal cancer","volume":"115","author":"Swidsinski","year":"1998","journal-title":"Gastroenterology"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1126\/science.1651562","article-title":"Identification of FAP locus genes from chromosome 5q21","volume":"253","author":"Kinzler","year":"1991","journal-title":"Science"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"592","DOI":"10.1126\/science.aah3648","article-title":"Patients with familial adenomatous polyposis harbor colonic biofilms containing tumorigenic bacteria","volume":"359","author":"Dejea","year":"2018","journal-title":"Science"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1186\/s13099-017-0185-x","article-title":"Prevalence of pks-positive Escherichia coli in Japanese patients with or without colorectal cancer","volume":"9","author":"Shimpoh","year":"2017","journal-title":"Gut Pathog."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1038\/nature11550","article-title":"Diversity, stability and resilience of the human gut microbiota","volume":"489","author":"Lozupone","year":"2012","journal-title":"Nature"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1038\/nrmicro2819","article-title":"A bacterial driver-passenger model for colorectal cancer: beyond the usual suspects","volume":"10","author":"Tjalsma","year":"2012","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1042\/CS20160876","article-title":"Interactions between microsatellite instability and human gut colonization by Escherichia coli in colorectal cancer","volume":"131","author":"Bonnin","year":"2017","journal-title":"Clin. Sci."},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Secher, T., Samba-Louaka, A., Oswald, E., and Nougayr\u00e8de, J.-P. (2013). Escherichia coli producing colibactin triggers premature and transmissible senescence in mammalian cells. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0077157"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"675","DOI":"10.4161\/19490976.2014.969989","article-title":"The bacterial genotoxin colibactin promotes colon tumor growth by modifying the tumor microenvironment","volume":"5","author":"Dalmasso","year":"2014","journal-title":"Gut Microbes"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1080\/19490976.2016.1155020","article-title":"Targeting colorectal cancer-associated bacteria: A new area of research for personalized treatments","volume":"7","author":"Delmas","year":"2016","journal-title":"Gut Microbes"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1093\/infdis\/jiu071","article-title":"The genotoxin colibactin exacerbates lymphopenia and decreases survival rate in mice infected with septicemic Escherichia coli","volume":"210","author":"Marcq","year":"2014","journal-title":"J. Infect. Dis."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"103","DOI":"10.3389\/fcimb.2017.00103","article-title":"Colibactin Contributes to the Hypervirulence of pks(+) K1 CC23 Klebsiella pneumoniae in Mouse Meningitis Infections","volume":"7","author":"Lu","year":"2017","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1556\/1886.2016.00029","article-title":"Insights from 100 Years of Research with Probiotic E. coli","volume":"6","author":"Wassenaar","year":"2016","journal-title":"Eur. J. Microbiol. Immunol. 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