O157:H7 and O104:H4 Vero/Shiga toxin-producing Escherichia coli outbreaks: respective role of cattle and humans

Kaper JB, Nataro JP, Mobley HL: Pathogenic Escherichia coli. Nat Rev Microbiol. 2004, 2: 123-140. 10.1038/nrmicro818. Gyles CL, Fairbrother JM: Escherichia coli. Pathogenesis of Bacterial Infections in Animals. Edited by: Gyles CL, Prescott JF, Songer JG, Thoen CO. 2010, Ames, IA: Wiley-Blackwell, 267-308. Nataro JP, Bopp CA, Fields PI, Kaper JB, Strockbine NA: Escherichia, Shigella, and Salmonella. Manual of Clinical Microbiology. Edited by: Versalovic J, Caroll KC, Funke G, Jorgensen JH, Landry ML, Warnock DW. 2011, Washington, DC: ASM Press, 603-626. 10 Nataro JP, Kaper JB: Diarrheagenic Escherichia coli. Clin Microbiol Rev. 1998, 11: 142-201. Griffin PM, Mead PM, Sivapalasingam S: Escherichia coli O157: H7 and other enterohemorrhagic E. coli. Infections of the Gastrointestinal Tract. Edited by: Blaser MJ, Smith PD, Ravdin JI, Greenberg HB, Guerrant RL. 2002, Philadelphia, PA: Lippincott Williams & Wilkins, 627-642. 2 Konowalchuk J, Speirs JI, Stavric S: Vero response to a cytotoxin of Escherichia coli. Infect Immun. 1977, 18: 775-779. O'Brien AD, LaVeck GD, Thompson MR, Formal SB: Production of Shigella dysenteriae type 1-like cytotoxin by Escherichia coli. J Infect Dis. 1982, 146: 763-769. 10.1093/infdis/146.6.763. Levine MM, Xu JG, Kaper JB, Lior H, Prado V, Tall B, Nataro J, Karch H, Wachsmuth K: A DNA probe to identify enterohemorrhagic Escherichia coli of O157:H7 and other serotypes that cause hemorrhagic colitis and hemolytic uremic syndrome. J Infect Dis. 1987, 156: 175-182. 10.1093/infdis/156.1.175. Coombes BK, Wickham ME, Mascarenhas M, Gruenheid S, Finlay BB, Karmali MA: Molecular analysis as an aid to assess the public health risk of non-O157 Shiga toxin-producing Escherichia col strains. Appl Environ Microbiol. 2008, 74: 2153-2160. 10.1128/AEM.02566-07. Mellmann A, Harmsen D, Cummings CA, Zentz EB, Leopold SR, Rico A, Prior K, Szczepanowski R, Ji Y, Zhang W, McLaughlin SF, Henkhaus JK, Leopold B, Bielaszewska M, Prager R, Brzoska PM, Moore RL, Guenther S, Rothberg JM, Karch H: Prospective genomic characterization of the German enterohemorrhagic Escherichia col O104:H4 outbreak by rapid next generation sequencing technology. PLoS One. 2011, 6: e22751-10.1371/journal.pone.0022751. Friedrich A: Enterohaemorrhagic Escherichia coli O104:H4: are we prepared now?. Euro Surveill. 2011, 16: pii = 19938. Available online: http://www.eurosurveillance.org/ViewArticle.aspx?ArticleId=19938 Mainil JG, Daube G: Verotoxigenic Escherichia col from animals, humans and foods: who's who?. J Appl Microbiol. 2005, 98: 1332-1344. 10.1111/j.1365-2672.2005.02653.x. Holland RE: Some infectious causes of diarrhea in young farm animals. Clin Microbiol Rev. 1990, 3: 345-375. Moxley RA, Smith DR: Attaching-effacing Escherichia coli infections in cattle. Vet Clin North Am Food Anim Pract. 2010, 26: 29-56. 10.1016/j.cvfa.2009.10.011. Uber AP, Trabulsi LR, Irino K, Beutin L, Ghilardi AC, Gomes TA, Liberatore AM, de Castro AF, Elias WP: Enteroaggregative Escherichia coli from humans and animals differ in major phenotypical traits and virulence genes. FEMS Microbiol Lett. 2006, 256: 251-257. 10.1111/j.1574-6968.2006.00124.x. Mainil J, Bez S, Jacquemin E, Kaeckenbeeck A: Les souches pathogènes d'Escherichia coli chez les chiens et les chats: I) Détection des souches entérotoxinogènes (ETEC), entéropathogènes (EPEC), vérotoxinogènes (VTEC), entérohémorragiques (EHEC) et nécrotoxinogènes (NTEC). Ann Med Vet. 1998, 142: 39-46. in French Garmendia J, Frankel G, Crepin VF: Enteropathogenic and enterohemorrhagic Escherichia coli infections: translocation, translocation, translocation. Infect Immun. 2005, 73: 2573-2585. 10.1128/IAI.73.5.2573-2585.2005. Schmidt MA: LEE ways: tales of EPEC, ATEC and EHEC. Cell Microbiol. 2010, 12: 1544-1552. 10.1111/j.1462-5822.2010.01518.x. Moon HW, Whipp SC, Argenzio RA, Levine MM, Giannella RA: Attaching and effacing activities of rabbit and human enteropathogenic Escherichia coli in pig and rabbit intestines. Infect Immun. 1983, 41: 1340-1351. Mainil J, Janssen L, Charlier G, Jacquemin E, China B, Goffaux F: Les souches pathogènes d'Escherichia coli chez les chiens et les chats: II) Données cliniques et bactériologiques sur les souches entéropathogènes. Ann Med Vet. 2000, 144: 335-343. in French Karmali MA, Gannon V, Sargeant JM: Verocytotoxin-producing Escherichia coli (VTEC). Vet Microbiol. 2010, 140: 360-370. 10.1016/j.vetmic.2009.04.011. Karmali MA, Mascarenhas M, Shen S, Ziebell K, Johnson S, Reid-Smith R, Isaac-Renton J, Clark C, Rahn K, Kaper JB: Association of genomic O island 122 of Escherichia coli EDL 933 with verocytotoxin-producing Escherichia col seropathotypes that are linked to epidemic and/or serious disease. J Clin Microbiol. 2003, 41: 4930-4940. 10.1128/JCM.41.11.4930-4940.2003. Pennington H: Escherichia coli O157. Lancet. 2010, 376: 1428-1435. 10.1016/S0140-6736(10)60963-4. Bolton DJ: Verocytotoxigenic (Shiga toxin-producing) Escherichia coli: virulence factors and pathogenicity in the farm to fork paradigm. Foodborne Pathog Dis. 2011, 8: 357-365. 10.1089/fpd.2010.0699. Bardiau M, Gregoire F, Muylaert A, Nahayo A, Duprez JN, Mainil J, Linden A: Enteropathogenic (EPEC), enterohaemorragic (EHEC) and verotoxigenic (VTEC) Escherichia coli in wild cervids. J Appl Microbiol. 2010, 109: 2214-2222. 10.1111/j.1365-2672.2010.04855.x. WHO Collaborating Centre for Reference and Research on Escherichia and Klebsiella, Statens Serum Institut: Identification of three vtx1 and seven vtx2 subtypes of Verocytotoxin encoding genes of Escherichia coli by conventional PCR amplification. [http://www.ssi.dk/English/PublicHealth/National%20Reference%20Laboratories/~/media/Indhold/EN%20-%20engelsk/Public%20Health/National%20Reference%20Laboratories/Vtx%20detection%20%20subtyping%20protocol_EQA-2010-11_rev4.ashx] Mainil JG, Jacquemin E, Pohl P, Fairbrother JM, Ansuini A, Le BC, Ball HJ, De Rycke J, Oswald E: Comparison of necrotoxigenic Escherichia coli isolates from farm animals and from humans. Vet Microbiol. 1999, 70: 123-135. 10.1016/S0378-1135(99)00134-0. Newton HJ, Sloan J, Bulach DM, Seemann T, Allison CC, Tauschek M, Robins-Browne RM, Paton JC, Whittam TS, Paton AW, Hartland EL: Shiga toxin-producing Escherichia coli strains negative for locus of enterocyte effacement. Emerg Infect Dis. 2009, 15: 372-380. 10.3201/eid1502.080631. Bardiau M, Szalo M, Mainil JG: Initial adherence of EPEC. EHEC and VTEC to host cells. Vet Res. 2010, 41: 57- Jenkins C, Perry NT, Cheasty T, Shaw DJ, Frankel G, Dougan G, Gunn GJ, Smith HR, Paton AW, Paton JC: Distribution of the saa gene in strains of Shiga toxin-producing Escherichia coli of human and bovine origins. J Clin Microbiol. 2003, 41: 1775-1778. 10.1128/JCM.41.4.1775-1778.2003. Levine MM: Escherichia col that cause diarrhea: enterotoxigenic, enteropathogenic, enteroinvasive, enterohemorrhagic, and enteroadherent. J Infect Dis. 1987, 155: 377-389. 10.1093/infdis/155.3.377. Nataro JP, Kaper JB, Robins-Browne R, Prado V, Vial P, Levine MM: Patterns of adherence of diarrheagenic Escherichia coli to HEp-2 cells. Pediatr Infect Dis J. 1987, 6: 829-831. 10.1097/00006454-198709000-00008. Bielaszewska M, Mellmann A, Zhang W, Kock R, Fruth A, Bauwens A, Peters G, Karch H: Characterisation of the Escherichia coli strain associated with an outbreak of haemolytic uraemic syndrome in Germany, 2011: a microbiological study. Lancet Infect Dis. 2011, 11: 671-676. Flores J, Okhuysen PC: Enteroaggregative Escherichia coli infection. Curr Opin Gastroenterol. 2009, 25: 8-11. 10.1097/MOG.0b013e32831dac5e. Harrington SM, Dudley EG, Nataro JP: Pathogenesis of enteroaggregative Escherichia coli infection. FEMS Microbiol Lett. 2006, 254: 12-18. 10.1111/j.1574-6968.2005.00005.x. Huang DB, Mohanty A, DuPont HL, Okhuysen PC, Chiang T: A review of an emerging enteric pathogen: enteroaggregative Escherichia coli. J Med Microbiol. 2006, 55: 1303-1311. 10.1099/jmm.0.46674-0. Weintraub A: Enteroaggregative Escherichia coli: epidemiology, virulence and detection. J Med Microbiol. 2007, 56: 4-8. 10.1099/jmm.0.46930-0. Scheutz F, Moller NE, Frimodt-Moller J, Boisen N, Morabito S, Tozzoli R, Nataro J, Caprioli A: Characteristics of the enteroaggregative Shiga toxin/verotoxin-producing Escherichia coli O104:H4 strain causing the outbreak of haemolytic uraemic syndrome in Germany, May to June 2011. Euro Surveill. 2011, 16: pii = 19889. Available online: http://www.eurosurveillance.org/ViewArticle.aspx?ArticleId=19889 Morabito S, Karch H, Mariani-Kurkdjian P, Schmidt H, Minelli F, Bingen E, Caprioli A: Enteroaggregative, Shiga toxin-producing Escherichia coli O111:H2 associated with an outbreak of hemolytic-uremic syndrome. J Clin Microbiol. 1998, 36: 840-842. Iyoda S, Tamura K, Itoh K, Izumiya H, Ueno N, Nagata K, Togo M, Terajima J, Watanabe H: Inducible stx phages are lysogenized in the enteroaggregative and other phenotypic Escherichia coli O86:HNM isolated from patients. FEMS Microbiol Lett. 2000, 191: 7-10. 10.1111/j.1574-6968.2000.tb09311.x. EU Reference Laboratory for E. coli, Istituto Superiore di Sanità: Detection and identification of Verocytotoxin-producing Escherichia coli (VTEC) O104:H4 in food by Real Time PCR Laboratory procedure. [http://www.iss.it/binary/vtec/cont/Lab_proc_O104_rev2.pdf] Aldabe B, Delmas Y, Gault G, Vendrely B, Llanas B, Charron M, Castor C, Ong N, Weill F, Mariani-Kurkdjian P, Terrier F, Desjardin M, Simoes J, Le Bihan B, Combe C, Rolland P: Household transmission of haemolytic uraemic syndrome associated with Escherichia coli O104:H4, south-western France, June 2011. Euro Surveill. 2011, 16: pii = 19934. Available online: http://www.eurosurveillance.org/ViewArticle.aspx?ArticleId=19934 Karch H, Russmann H, Schmidt H, Schwarzkopf A, Heesemann J: Long-term shedding and clonal turnover of enterohemorrhagic Escherichia coli O157 in diarrheal diseases. J Clin Microbiol. 1995, 33: 1602-1605. Aidar L, Penteado AS, Trabulsi LR, Blanco JE, Blanco M, Blanco J, de Castro Pestana: Subtypes of intimin among non-toxigenic Escherichia coli from diarrheic calves in Brazil. Can J Vet Res. 2000, 64: 15-20. Penteado AS, Aidar L, de Castro Pestana, Yamada A, Andrade JR, Blanco J, Blanco M, Blanco JE: eae-negative attaching and effacing Escherichia coli from piglets with diarrhea. Res Microbiol. 2001, 152: 75-81. 10.1016/S0923-2508(00)01170-0. Oundo JO, Kariuki SM, Boga HI, Muli FW, Iijima Y: High incidence of enteroaggregative Escherichia coli among food handlers in three areas of Kenya: a possible transmission route of travelers' diarrhea. J Travel Med. 2008, 15: 31-38. 10.1111/j.1708-8305.2007.00174.x. Law D, Chart H: Enteroaggregative Escherichia coli. J Appl Microbiol. 1998, 84: 685-697. 10.1046/j.1365-2672.1998.00372.x. Frank C, Faber M, Askar M, Bernard H, Fruth A, Gilsdorf A, Hohle M, Karch H, Krause G, Prager R, Spode A, Stark K, Werber D, HUS investigation team: Large and ongoing outbreak of haemolytic uraemic syndrome, Germany, May 2011. Euro Surveill. 2011, 16: pii = 19878. Available online: http://www.eurosurveillance.org/ViewArticle.aspx?ArticleId=19878 European Food Safety Authority: Tracing seeds, in particular fenugreek (Trigonella foenum-graecum) seeds, in relation to the Shiga toxin-producing E. coli (STEC) O104:H4 2011 Outbreaks in Germany and France. 2011. [http://www.efsa.europa.eu/en/supporting/doc/176e.pdf] Pennington H: Escherichia coli O104, Germany 2011. Lancet Infect Dis. 2011, 11: 652-653. 10.1016/S1473-3099(11)70166-9. Buchholz U, Bernard H, Werber D, Bohmer MM, Remschmidt C, Wilking H, Delere Y, an der HM, Adlhoch C, Dreesman J, Ehlers J, Ethelberg S, Faber M, Frank C, Fricke G, Greiner M, Höhle M, Ivarsson S, Jark U, Kirchner M, Koch J, Krause G, Luber P, Rosner B, Stark K, Kühne M: German outbreak of Escherichia coli O104:H4 associated with sprouts. N Engl J Med. 2011, 365: 1763-1770. 10.1056/NEJMoa1106482. Berger CN, Sodha SV, Shaw RK, Griffin PM, Pink D, Hand P, Frankel G: Fresh fruit and vegetables as vehicles for the transmission of human pathogens. Environ Microbiol. 2010, 12: 2385-2397. 10.1111/j.1462-2920.2010.02297.x. Ferens WA, Hovde CJ: Escherichia coli O157:H7: animal reservoir and sources of human infection. Foodborne Pathog Dis. 2011, 8: 465-487. 10.1089/fpd.2010.0673. Michino H, Araki K, Minami S, Takaya S, Sakai N, Miyazaki M, Ono A, Yanagawa H: Massive outbreak of Escherichia coli O157:H7 infection in schoolchildren in Sakai City, Japan, associated with consumption of white radish sprouts. Am J Epidemiol. 1999, 150: 787-796. 10.1093/oxfordjournals.aje.a010082. Robert Koch Institute: Report: Final presentation and evaluation of epidemiological findings in the EHEC O104:H4 outbreak, Germany 2011. Berlin. 2011, [http://www.rki.de/cln_151/nn_217400/EN/Home/EHEC__final__report,templateId=raw,property=publicationFile.pdf/EHEC_final_report.pdf] Gault G, Weill F, Mariani-Kurkdjian P, Jourdan-da SN, King L, Aldabe B, Charron M, Ong N, Castor C, Mace M, Bingen E, Noel H, Vaillant V, Bone A, Vendrely B, Delmas Y, Combe C, Bercion R, d'Andigne E, Desjardin M, de Valk H, Rolland P: Outbreak of haemolytic uraemic syndrome and bloody diarrhoea due to Escherichia coli O104:H4, south-west France, June 2011. Euro Surveill. 2011, 16: pii = 19905. Available online: http://www.eurosurveillance.org/ViewArticle.aspx?ArticleId=19905 Mariani-Kurkdjian P, Bingen E, Gault G, Silva NJ, Weill FX: Escherichia coli O104:H4 south-west France, June 2011. Lancet Infect Dis. 2011, 11: 732-733. 10.1016/S1473-3099(11)70266-3. Frank C, Werber D, Cramer JP, Askar M, Faber M, Heiden MA, Bernard H, Fruth A, Prager R, Spode A, Wadl M, Zoufaly A, Jordan S, Kemper MJ, Follin P, Müller L, King LA, Rosner B, Buchholz U, Stark K, Krause G, HUS Investigation Team: Epidemic profile of Shiga-toxin-producing Escherichia coli O104:H4 outbreak in Germany. N Engl J Med. 2011, 365: 1771-1780. 10.1056/NEJMoa1106483. Rasko DA, Webster DR, Sahl JW, Bashir A, Boisen N, Scheutz F, Paxinos EE, Sebra R, Chin CS, Iliopoulos D, Klammer A, Peluso P, Lee L, Kislyuk AO, Bullard J, Kasarskis A, Wang S, Eid J, Rank D, Redman JC, Steyert SR, Frimodt-Møller J, Struve C, Petersen AM, Krogfelt KA, Nataro JP, Schadt EE, Waldor MK: Origins of the Escherichia coli strain causing an outbreak of hemolytic-uremic syndrome in Germany. N Engl J Med. 2011, 365: 709-717. 10.1056/NEJMoa1106920. European Centre for Disease Prevention and Control: Annual Epidemiological Report on Communicable Diseases in Europe 2010. 2010, Stockholm: ECDC, [http://ecdc.europa.eu/en/publications/publications/1011_sur_annual_epidemiological_report_on_communicable_diseases_in_europe.pdf] Wendel AM, Johnson DH, Sharapov U, Grant J, Archer JR, Monson T, Koschmann C, Davis JP: Multistate outbreak of Escherichia coli O157:H7 infection associated with consumption of packaged spinach, August-September 2006: the Wisconsin investigation. Clin Infect Dis. 2009, 48: 1079-1086. 10.1086/597399. Karmali MA, Steele BT, Petric M, Lim C: Sporadic cases of haemolytic-uraemic syndrome associated with faecal cytotoxin and cytotoxin-producing Escherichia coli in stools. Lancet. 1983, 1: 619-620. Riley LW, Remis RS, Helgerson SD, McGee HB, Wells JG, Davis BR, Hebert RJ, Olcott ES, Johnson LM, Hargrett NT, Blake PA, Cohen ML: Hemorrhagic colitis associated with a rare Escherichia coli serotype. N Engl J Med. 1983, 308: 681-685. 10.1056/NEJM198303243081203. Persson S, Olsen KE, Scheutz F, Krogfelt KA, Gerner-Smidt P: A method for fast and simple detection of major diarrhoeagenic Escherichia coli in the routine diagnostic laboratory. Clin Microbiol Infect. 2007, 13: 516-524. 10.1111/j.1469-0691.2007.01692.x. Weinstein DL, Jackson MP, Samuel JE, Holmes RK, O'Brien AD: Cloning and sequencing of a Shiga-like toxin type II variant from Escherichia coli strain responsible for edema disease of swine. J Bacteriol. 1988, 170: 4223-4230. Karch H, Meyer T, Russmann H, Heesemann J: Frequent loss of Shiga-like toxin genes in clinical isolates of Escherichia coli upon subcultivation. Infect Immun. 1992, 60: 3464-3467. Bielaszewska M, Prager R, Kock R, Mellmann A, Zhang W, Tschape H, Tarr PI, Karch H: Shiga toxin gene loss and transfer in vitro and in vivo during enterohemorrhagic Escherichia coli O26 infection in humans. Appl Environ Microbiol. 2007, 73: 3144-3150. 10.1128/AEM.02937-06. Toth I, Schmidt H, Dow M, Malik A, Oswald E, Nagy B: Transduction of porcine enteropathogenic Escherichia coli with a derivative of a Shiga toxin 2-encoding bacteriophage in a porcine ligated ileal loop system. Appl Environ Microbiol. 2003, 69: 7242-7247. 10.1128/AEM.69.12.7242-7247.2003. Johannes L, Romer W: Shiga toxins-from cell biology to biomedical applications. Nat Rev Microbiol. 2010, 8: 105-116. Sandvig K, Bergan J, Dyve AB, Skotland T, Torgersen ML: Endocytosis and retrograde transport of Shiga toxin. Toxicon. 2010, 56: 1181-1185. 10.1016/j.toxicon.2009.11.021. Pruimboom-Brees IM, Morgan TW, Ackermann MR, Nystrom ED, Samuel JE, Cornick NA, Moon HW: Cattle lack vascular receptors for Escherichia coli O157:H7 Shiga toxins. Proc Natl Acad Sci USA. 2000, 97: 10325-10329. 10.1073/pnas.190329997. Hoey DE, Currie C, Else RW, Nutikka A, Lingwood CA, Gally DL, Smith DG: Expression of receptors for verotoxin 1 from Escherichia coli O157 on bovine intestinal epithelium. J Med Microbiol. 2002, 51: 143-149. Hoey DE, Sharp L, Currie C, Lingwood CA, Gally DL, Smith DG: Verotoxin 1 binding to intestinal crypt epithelial cells results in localization to lysosomes and abrogation of toxicity. Cell Microbiol. 2003, 5: 85-97. 10.1046/j.1462-5822.2003.00254.x. Oswald E, Schmidt H, Morabito S, Karch H, Marches O, Caprioli A: Typing of intimin genes in human and animal enterohemorrhagic and enteropathogenic Escherichia coli: characterization of a new intimin variant. Infect Immun. 2000, 68: 64-71. 10.1128/IAI.68.1.64-71.2000. Hartland EL, Batchelor M, Delahay RM, Hale C, Matthews S, Dougan G, Knutton S, Connerton I, Frankel G: Binding of intimin from enteropathogenic Escherichia coli to Tir and to host cells. Mol Microbiol. 1999, 32: 151-158. 10.1046/j.1365-2958.1999.01338.x. Luo Y, Frey EA, Pfuetzner RA, Creagh AL, Knoechel DG, Haynes CA, Finlay BB, Strynadka NC: Crystal structure of enteropathogenic Escherichia coli intimin-receptor complex. Nature. 2000, 405: 1073-1077. 10.1038/35016618. Hayashi T, Makino K, Ohnishi M, Kurokawa K, Ishii K, Yokoyama K, Han CG, Ohtsubo E, Nakayama K, Murata T, Tanaka M, Tobe T, Iida T, Takami H, Honda T, Sasakawa C, Ogasawara N, Yasunaga T, Kuhara S, Shiba T, Hattori M, Shinagawa H: Complete genome sequence of enterohemorrhagic Escherichia coli O157:H7 and genomic comparison with a laboratory strain K-12. DNA Res. 2001, 8: 11-22. 10.1093/dnares/8.1.11. Perna NT, Plunkett G, Burland V, Mau B, Glasner JD, Rose DJ, Mayhew GF, Evans PS, Gregor J, Kirkpatrick HA, Pósfai G, Hackett J, Klink S, Boutin A, Shao Y, Miller L, Grotbeck EJ, Davis NW, Lim A, Dimalanta ET, Potamousis KD, Apodaca J, Anantharaman TS, Lin J, Yen G, Schwartz DC, Welch RA, Blattner FR: Genome sequence of enterohaemorrhagic Escherichia coli O157:H7. Nature. 2001, 409: 529-533. 10.1038/35054089. Kulasekara BR, Jacobs M, Zhou Y, Wu Z, Sims E, Saenphimmachak C, Rohmer L, Ritchie JM, Radey M, McKevitt M, Freeman TL, Hayden H, Haugen E, Gillett W, Fong C, Chang J, Beskhlebnaya V, Waldor MK, Samadpour M, Whittam TS, Kaul R, Brittnacher M, Miller SI: Analysis of the genome of the Escherichia coli O157:H7 2006 spinach-associated outbreak isolate indicates candidate genes that may enhance virulence. Infect Immun. 2009, 77: 3713-3721. 10.1128/IAI.00198-09. Ogura Y, Ooka T, Iguchi A, Toh H, Asadulghani M, Oshima K, Kodama T, Abe H, Nakayama K, Kurokawa K, Tobe T, Hattori M, Hayashi T: Comparative genomics reveal the mechanism of the parallel evolution of O157 and non-O157 enterohemorrhagic Escherichia coli. Proc Natl Acad Sci USA. 2009, 106: 17939-17944. 10.1073/pnas.0903585106. Bardiau M, Labrozzo S, Mainil JG: Putative adhesins of enteropathogenic and enterohemorrhagic Escherichia coli of serogroup O26 isolated from humans and cattle. J Clin Microbiol. 2009, 47: 2090-2096. 10.1128/JCM.02048-08. Czeczulin JR, Balepur S, Hicks S, Phillips A, Hall R, Kothary MH, Navarro-Garcia F, Nataro JP: Aggregative adherence fimbria II, a second fimbrial antigen mediating aggregative adherence in enteroaggregative Escherichia coli. Infect Immun. 1997, 65: 4135-4145. Boisen N, Struve C, Scheutz F, Krogfelt KA, Nataro JP: New adhesin of enteroaggregative Escherichia coli related to the Afa/Dr/AAF family. Infect Immun. 2008, 76: 3281-3292. 10.1128/IAI.01646-07. Bernier C, Gounon P, Le Bouguénec C: Identification of an aggregative adhesion fimbria (AAF) type III-encoding operon in enteroaggregative Escherichia coli as a sensitive probe for detecting the AAF-encoding operon family. Infect Immun. 2002, 70: 4302-4311. 10.1128/IAI.70.8.4302-4311.2002. Cravioto A, Tello A, Navarro A, Ruiz J, Villafan H, Uribe F, Eslava C: Association of Escherichia coli HEp-2 adherence patterns with type and duration of diarrhoea. Lancet. 1991, 337: 262-264. 10.1016/0140-6736(91)90868-P. Servin AL: Pathogenesis of Afa/Dr diffusely adhering Escherichia coli. Clin Microbiol Rev. 2005, 18: 264-292. 10.1128/CMR.18.2.264-292.2005. Nataro JP: Enteroaggregative Escherichia coli pathogenesis. Curr Opin Gastroenterol. 2005, 21: 4-8. Farfan MJ, Inman KG, Nataro JP: The major pilin subunit of the AAF/II fimbriae from enteroaggregative Escherichia coli mediates binding to extracellular matrix proteins. Infect Immun. 2008, 76: 4378-4384. 10.1128/IAI.00439-08. Chaudhuri RR, Sebaihia M, Hobman JL, Webber MA, Leyton DL, Goldberg MD, Cunningham AF, Scott-Tucker A, Ferguson PR, Thomas CM, Frankel G, Tang CM, Dudley EG, Roberts IS, Rasko DA, Pallen MJ, Parkhill J, Nataro JP, Thomson NR, Henderson IR: Complete genome sequence and comparative metabolic profiling of the prototypical enteroaggregative Escherichia coli strain O42. PLoS One. 2010, 5: e8801-10.1371/journal.pone.0008801. Feng P, Lampel KA, Karch H, Whittam TS: Genotypic and phenotypic changes in the emergence of Escherichia coli O157:H7. J Infect Dis. 1998, 177: 1750-1753. 10.1086/517438. Feng PC, Monday SR, Lacher DW, Allison L, Siitonen A, Keys C, Eklund M, Nagano H, Karch H, Keen J, Whittam TS: Genetic diversity among clonal lineages within Escherichia coli O157:H7 stepwise evolutionary model. Emerg Infect Dis. 2007, 13: 1701-1706. 10.3201/eid1311.070381. Whittam TS, Wolfe ML, Wachsmuth IK, Orskov F, Orskov I, Wilson RA: Clonal relationships among Escherichia coli strains that cause hemorrhagic colitis and infantile diarrhea. Infect Immun. 1993, 61: 1619-1629. Brzuszkiewicz E, Thurmer A, Schuldes J, Leimbach A, Liesegang H, Meyer FD, Boelter J, Petersen H, Gottschalk G, Daniel R: Genome sequence analyses of two isolates from the recent Escherichia col outbreak in Germany reveal the emergence of a new pathotype: Entero-Aggregative-Haemorrhagic Escherichia coli (EAHEC). Arch Microbiol. 2011, 193: 883-891. 10.1007/s00203-011-0725-6.