Shiga toxin: Expression, distribution, and its role in the environment

Toxins

Volumn 3, Issue 6, 2011, Pages 608-625

  Mauro, Steven A ^a   Koudelka, Gerald B ^b  

^a MERCYHURST COLLEGE   (United States)

^b UNIVERSITY AT BUFFALO   (United States)

Author keywords

Bacteriophage; E. coli O157; Shiga toxin

Indexed keywords

SHIGA TOXIN; VEROTOXIN 1; VEROTOXIN 2;

AQUATIC ENVIRONMENT; BACTERIAL GENE; BACTERIOPHAGE; CYTOTOXICITY; FECES ANALYSIS; GENE CONSTRUCT; GENE CONTROL; GENE EXPRESSION REGULATION; GENE FUNCTION; GENE LOCATION; GENETIC ANALYSIS; MICROBIAL DIVERSITY; NONHUMAN; REGULATOR GENE; REGULATORY MECHANISM; REPRESSOR GENE; REVIEW; SHIGA TOXIN 1 GENE; SHIGA TOXIN 2 GENE; SHIGA TOXIN PRODUCING ESCHERICHIA COLI; SHIGELLA DYSENTERIAE; TOXIN ANALYSIS; TOXIN SYNTHESIS; WATER ANALYSIS;

BACTERIOPHAGES; ECOSYSTEM; ENVIRONMENTAL MICROBIOLOGY; ESCHERICHIA COLI INFECTIONS; ESCHERICHIA COLI O157; HOST-PATHOGEN INTERACTIONS; SHIGA TOXIN 1; SHIGA TOXIN 2; VIRULENCE FACTORS;

EID: 79959677993     PISSN: None     EISSN: 20726651     Source Type: Journal    
DOI: 10.3390/toxins3060608     Document Type: Review

References (110)

1. Riley, L.W.; Remis, R.S.; Helgerson, S.D.; McGee, H.B.; Wells, J.G.; Davis, B.R.; Hebert, R.J.; Olcott, E.S.; Johnson, L.M.; Hargrett, N.T. et al. Hemorrhagic colitis associated with a rare Escherichia-coli serotype. New Engl. J. Med. 1983, 308, 681-685.
2. O'Brien, A.D.; LaVeck, G.D.; Thompson, M.R.; Formal, S.B. Production of Shigella dysenteriae type 1-like cytotoxin by Escherichia coli. J. Infect. Dis. 1982, 146, 763-769.
3. Scotland, S.M.; Smith, H.R.; Rowe, B. Two distinct toxins active on VERO cells from Escherichia coli O157. Lancet 1985, 2, 885-886.
4. Strockbine, N.A.; Marques, L.R.; Newland, J.W.; Smith, H.W.; Holmes, R.K.; O'Brien, A.D. Two toxin-converting phages from Escherichia coli O157:H7 strain 933 encode antigenically distinct toxins with similar biologic activities. Infect. Immunol. 1986, 53, 135-140.
5. Strockbine, N.A.; Jackson, M.P.; Sung, L.M.; Holmes, R.K.; O'Brien, A.D. Cloning and sequencing of the genes for Shiga toxin from Shigella dysenteriae type 1. J. Bacteriol. 1988, 170, 1116-1122.
6. Bridgwater, F.A.; Morgan, R.S.; Rowson, K.E.; Wright, G.P. The neurotoxin of Shigella shigae: Morphological and functional lesions produced in the central nervous system of rabbits. Br. J. Exp. Pathol. 1955, 36, 447-453.
7. Moore, M.A.; Blaser, M.J.; Perez-Perez, G.I.; O'Brien, A.D. Production of a Shiga-like cytotoxin by Campylobacter. Microb. Pathog. 1988, 4, 455-462.
8. Schmidt, H.; Montag, M.; Bockemuhl, J.; Heesemann, J.; Karch, H. Shiga-like toxin II-related cytotoxins in Citrobacter freundii Strains from humans and beef samples. Infec. Immunol. 1993, 61, 534-543.
9. Haque, Q.M.; Sugiyama, A.; Iwade, Y.; Midorikawa, Y.; Yamauchi, T. Diarrheal and environmental isolates of Aeromonas spp. produce a toxin similar to Shiga-like toxin 1. Curr. Microbiol. 1996, 32, 239-245.
10. Paton, A.W.; Paton, J.C. Enterobacter cloacae producing a Shiga-like toxin II-Related Cytotoxin Associated with a Case of Hemolytic-Uremic Syndrome. J. Clin. Microbiol. 1996, 34, 463-465.
11. Grotiuz, G.; Sirok, A.; Gadea, P.; Varela, G.; Schelotto, F. Shiga toxin 2-producing Acinetobacter haemolyticus associated with a case of bloody diarrhea. J. Clin. Microbiol. 2006, 44, 3838-3841.
12. Degnan, P.H.; Moran, N.A. Diverse phage-encoded toxins in a protective insect endosymbiont. Appl. Environ. Microbiol. 2008, 74, 6782-6791.
13. Alperi, A.; Figueras, J. Human isolates of Aeromonas possess Shiga toxin genes (stx1 and stx2) highly similar to the most virulent gene variants of Escherichia coli. Clin. Microbiol. Infect. 2010, 16, 1563-1567.
14. O'Brien, A.D.; Tesh, V.L.; Donohue-Rolfe, A.; Jackson, M.P.; Olsnes, S.; Sandvig, K.; Lindberg, A.A.; Keusch, G.T. Shiga toxin: Biochemistry, genetics, mode of action, and role in pathogenesis. Curr. Top. Microbiol. Immunol. 1992, 180, 65-94.
15. Sato, T.; Shimizu, T.; Watarai, M.; Kobayashi, M.; Kano, S.; Hamabata, T.; Takeda, Y.; Yamasaki, S. Genome analysis of a novel Shiga toxin 1 (Stx1)-converting phage which is closely related to Stx2-converting phages but not to other Stx1-converting phages. J. Bacteriol. 2003, 185, 3966-3971.
16. Fattah, K.R.; Mizutani, S.; Fattah, F.J.; Matsushiro, A.; Sugino, Y. A comparative study of the immunity region of lambdoid phages including Shiga-toxin-converting phages: Molecular basis for cross immunity. Genes Genet. Syst. 2000, 75, 223-232.
17. Perna, N.T.; Plunkett, G., III; Burland, V.; Mau, B.; Glasner, J.D.; Rose, J.D.; Mayhew, G.F.; Evans, P.S.; Gregor, J.; Kirkpatrick, H.A. et al. Genome sequence of enterohaemorrhagic Escherichia coli O157:H7. Nature 2001, 409, 529-533.
18. Caprioli, A.; Morabito, S.; Brugere, H.; Oswald, E. Enterohaemorrhagic Escherichia coli: Emerging issues on virulence and modes of transmission. Vet. Res. 2005, 36, 289-311.
19. Hussein, H.S. Prevalence and pathogenicity of Shiga toxin-producing Escherichia coli in beef cattle their products. J. Anim. Sci. 2007, 85, E63-E72.
20. Fremaux, B.; Prignet-Combaret, C.; Vernozy-Rozand, C. Long-term survival of Shiga toxin-producing Escherichia coli in cattle effluents and environment: An updated review. Vet. Microbiol. 2008, 132, 1-18.
21. Ferens, W.; Hovde, C.J. Escherichia coli O157:H7: Animal reservoir and sources of human infection. Foodborne Pathog. Dis. 2011, 8, 465-487.
22. Wilson, J.B.; McEwen, S.A.; Clarke, R.C.; Leslie, K.E.; Waltner-Toews, D.; Gyles, C.L. A case-control study of selected pathogens including verocytotoxigenic Escherichia coli in calf diarrhea on an Ontario veal farm. Can. J. Vet. Res. 1992, 56, 184-188.
23. Imamovic, L.; Balleste, E.; Jofre, J.; Muniesa, M. Quantification of Shiga toxin-converting bacteriophages in wastewater and in fecal samples by real-time quantitative PCR. Appl. Environ. Microbiol. 2010, 76, 5693-5701.
24. Rogerie, F.; Marecat, A.; Gambade, S.; Duponda, F.; Beauboisb, P.; Lange, M. Characterization of Shiga toxin producing E. coli and O157 serotype E. coli isolated in France from healthy domestic cattle. Int. J. Food Microbiol. 2001, 63, 217-223.
25. Jenkins, C.; Pearce, M.C.; Chart, H.; Cheasty, T.; Willshaw, G.A.; Gunn, G.J.; Dougan, G.; Smith, H.R.; Synge, B.A.; Frankel, G. An eight-month study of a population of verocytotoxigenic Escherichia coli (VTEC) in a Scottish cattle herd. J. Appl. Microbiol. 2002, 93, 944-953.
26. Gilbreath, J.J.; Shields, M.S.; Smith, R.L.; Farrell, L.D.; Sheridan, P.P.; Spiegel, K.M. Shiga toxins, and the genes encoding them, in fecal samples from native idaho ungulates. Appl. Environ. Microbiol. 2009, 75, 862-865.
27. Menrath, A.; Wieler, L.H.; Heidemanns, K.; Semmler, T.; Fruth, A.; Kemper, N. Shiga toxin producing Escherichia coli: Identification of non-O157:H7-Super-Shedding cows and related risk factors. Gut Pathog. 2010, 2, 7.
28. Cerqueira, A.M.F.; Guth, B.E.C.; Joaquim, R.M.; Andrade, J.R.C. High occurrence of Shiga toxin-producing Escherichia coli (STEC) in healthy cattle in Rio de Janeiro State, Brazil. Vet. Microbiol. 1999, 70, 111-121.
29. Fernandez, D.; Rodriguez, E.M.; Arroyo, G.H.; Padola, N.L.; Parma, A.E. Seasonal variation of Shiga toxin-encoding genes (stx) and detection of E. coli O157 in dairy cattle from Argentina. J. Appl. Microbiol. 2009, 106, 1260-1267.
30. Urdahl, A.M.; Beutin, L.; Skjerve, E.; Zimmermann, S.; Wasteson, Y. Animal host associated differences in Shiga toxin-producing Escherichia coli isolated from sheep and cattle on the same farm. J. Appl. Microbiol. 2003, 95, 92-101.
31. Cookson, A.L.; Taylor, S.C.; Attwood, G.T. The prevalence of Shiga toxin-producing Escherichia coli in cattle and sheep in the lower North Island, New Zealand. N. Z. Vet. J. 2006, 54, 28-33.
32. Heuvelink, A.E.; Van Den Biggelaar, F.L.A.M.; De Boer, E.; Herbes, R.G.; Melchers, W.J.G.; Huis In'T Veld, J.H.J.; Monnens, L.A.H. Isolation and Characterization of Verocytotoxin-producing Escherichia coli o157 strains from Dutch cattle and sheep. J. Clin. Microbiol. 1998, 36, 878-882.
33. Zschock, M.; Hamann, H.P.; Kloppert, B.; Wolter, W. Shiga-toxin-producing Escherichia coli in faeces of healthy dairy cows, sheep and goats: Prevalence and virulence properties. Lett. Appl. Microbiol. 2000, 31, 203-208.
34. Corte's, C.; de la Fuente, R.; Blanco, J.; Blanco, M.; Blanco, J.E.; Dhabi, G.; Mora, A.; Justel, P.; Conteras, A.; Sanchez, A. et al. Serotypes, virulence genes and intimin types of verotoxin-producing Escherichia coli and enteropathogenic E. coli isolated from healthy dairy goats in Spain. Vet. Microbiol. 2005, 110, 67-76.
35. Wani, S.A.; Samantha, I.; Munshi, Z.H.; Bhat, M.A.; Nishikawa, Y. Shiga toxin-producing Escherichia coli and enteropathogenic Escherichia coli in healthy goats in India: Occurrence and virulence properties. J. Appl. Microbiol. 2006, 100, 108-113.
36. Vu-Khac, H.; Cornick, N.A. Prevalence and genetic profiles of Shiga toxin-producing Escherichia coli strains isolated from buffaloes, cattle, and goats in central Vietnam. Vet. Microbiol. 2008, 126, 356-363.
37. Galiero, G.; Conedera, G.; Alfano, D.; Caprioli, A. Isolation of verocytotoxin-producing Escherichia coli O157 from water buffaloes (Bubalus bubalis) in southern Italy. Vet Rec. 2005, 156, 382-383.
38. Oliveira, M.G.; Brito, J.R.; Carvalho, R.R.; Guth, B.E.; Gomes, T.A.; Vieira, M.A.; Kato, M.A.; Ramos, I.I.; Vaz, T.M.; Irino, K. Water buffaloes (Bubalus bubalis) identified as an important reservoir of Shiga toxin-producing Escherichia coli in Brazil. Appl. Environ. Microbiol. 2007, 73, 5945-5948.
39. Fratamico, P.M.; Bhagwat, A.A.; Injaian, L.; Fedorka-Cray, P.J. Characterization of Shiga toxin-producing Escherichia coli strains isolated from swine feces. Foodborne Pathog. Dis. 2008, 5, 827-838.
40. Kim, Y.J.; Kim, J.H.; Hur, J.; Lee, J.H. Isolation of Escherichia coli from piglets in South Korea with diarrhea and characteristics of the virulence genes. Can. J. Vet. Res. 2010, 74, 59-64.
41. Yan, Y.; Shi, Y.; Cao, D.; Meng, X.; Xia, L.; Sun, J. Prevalence of Stx phages in environments of a pig farm and lysogenic infection of the field E. coli O157 isolates with a recombinant converting Phage. Curr. Microbiol. 2011, 62, 458-464.
42. Heuvelink, A.E.; Zwartkruis-Nahuis, J.T.; van den Biggelaar, F.L.; van Leeuwen, W.J.; de Boer, E. Isolation and characterization of Vero cytotoxin-producing Escherichia coli O157 from slaughter pigs and poultry. Int. J. Food. Microbiol. 1999, 52, 67-75.
43. García, A.; Fox, J.G. The rabbit as a new reservoir host of enter hemorrhagic Escherichia coli. Emerg. Infect. Dis. 2003, 9, 1592-1597.
44. Dipineto, L.; Santaniello, A.; Fontanella, M.; Lagos, K.; Fioretti, A.; Menna, L.F. Presence of Shiga toxin-producing Escherichia coli O157:H7 in living layer hens. Lett. Appl. Microbiol. 2006, 43, 293-295.
45. Farooq, S.; Hussain, I.; Mir, M.A.; Bhat, M.A.; Wani, S.A. Isolation of atypical enter pathogenic Escherichia coli and Shiga toxin 1 and 2f-producing Escherichia coli from avian species in India. Lett. Appl. Microbiol. 2009, 48, 692-697.
46. Asakura, H. Makino, S.; Shirahata, T.; Tsukamoto, T.; Kurazono, H.; Ikeda, T.; Takeshi, K. Detection and genetical characterization of Shiga toxin-producing Escherichia coli from wild deer. Microbiol. Immunol. 1998, 42, 815-822.
47. Sánchez, S.; García-Sánchez, A.; Martínez, R.; Blanco, J.; Blanco, J.E.; Blanco, M.; Dahbi, G.; Mora, A.; Hermoso de Mendoza, J.; Alonso, J.M. et al. Detection and characterization of Shiga toxin-producing Escherichia coli other than Escherichia coli O157:H7 in wild ruminants. Vet. J. 2009, 180, 384-388.
48. Kistler, W.; Mulugeta, S.; Mauro, S.A. Shiga toxin 2 genes are abundant in deer feces from Western Pennsylvania. Toxins 2011, submitted for publication.
49. Leotta, G.A.; Deza, N.; Origlia, J.; Toma, C.; Chinen, I.; Miliwebsky, E.; Iyoda, S.; Sosa-Estani, S.; Rivas, M. Detection and characterization of Shiga toxin-producing Escherichia coli in captive non-domestic mammals. Vet. Microbiol. 2006, 118, 151-157.
50. Bentancor, A.; Rumi, M.V.; Gentilini, M.V.; Sardoy, C.; Irino, K.; Agostini, A.; Cataldi, A. Shiga toxin-producing and attaching and effacing Escherichia coli in cats and dogs in a high hemolytic uremic syndrome incidence region in Argentina. FEMS Microbiol. Lett. 2007, 267, 251-256.
51. Sánchez, S.; Martínez, R.; García, A.; Vidal, D.; Blanco, J.; Blanco, M.; Blanco, J.E.; Mora, A.; Herrera-León, S.; Echeita, A. et al. Detection and characterisation of O157:H7 and non-O157 Shiga toxin-producing Escherichia coli in wild boars. Vet. Microbiol. 2010, 143, 420-423.
52. Hughes, L.A.; Bennett, M.; Coffey, P.; Elliott, J.; Jones, T.R.; Jones, R.C.; Lahuerta-Marin, A.; McNiffe, K.; Norman, D.; Williams, N.J. et al. Risk factors for the occurrence of Escherichia coli virulence genes eae, stx1 and stx2 in wild bird populations. Epidemiol. Infect. 2009, 137, 1574-1582.
53. Varma, J.K.; Greene, K.D.; Reller, M.E.; DeLong, S.M.; Trottier, J.; Nowicki, S.F.; DiOrio, M.; Koch, E.M.; Bannerman, T.L.; York, S.T. et al. An outbreak of Escherichia coli O157 infection following exposure to a contaminated building. J. Am. Med. Assoc. 2003, 290, 2709-2712.
54. Muniesa, M.; Jofre, J.; García-Aljaro, C.; Blanch, A.R. Occurrence of Escherichia coli O157:H7 and other enterohemorrhagic Escherichia coli in the environment. Environ. Sci. Technol. 2006, 40, 7141-7149.
55. Swerdlow, D.L.; Woodruff, B.A.; Brady, R.C.; Griffin, P.M.; Tippen, S.; Donnell, H.D., Jr.; Geldreich, E.; Payne, B.J.; Meyer, A., Jr.; Wells, J.G. et al. A waterborne outbreak in Missouri of Escherichia coli O157:H7 associated with bloody diarrhea and death. Ann. Intern. Med. 1992, 117, 812-819.
56. Licence, K.; Oates, K.R.; Synge, B.A.; Reid, T.M. An outbreak of E. coli O157 infection with evidence of spread from animals to man through contamination of a private water supply. Epidemiol. Infect. 2001, 126, 135-138.
57. Yatsuyanagi, J.; Saito, S.; Ito, I. A case of hemolytic-uremic syndrome associated with Shiga toxin 2-producing Escherichia coli O121 infection caused by drinking water contaminated with bovine feces. Jpn. J. Infect. Dis. 2002, 55, 174-176.
58. Bopp, D.J.; Sauders, B.D.; Waring, A.L.; Ackelsberg, J.; Dumas, N.; Braun-Howland, E.; Dziewulski, D.; Wallace, B.J.; Kelly, M.; Halse, T. et al. Detection, isolation, and molecular subtyping of Escherichia coli O157:H7 and Campylobacter jejuni associated with a large waterborne outbreak. J. Clin. Microbiol. 2003, 41, 174-180.
59. McCall, B.J.; Slinko, V.G.; Smith, H.V.; Heel, K.; Culleton, T.H.; Kelk, V.R.; Stafford, R.J. An outbreak of Shiga toxin-producing Escherichia coli infection associated with a school camp. Commun. Dis. Intell. 2010, 34, 54-56.
60. Lienemann, T.; Pitkänen, T.; Antikainen, J.; Mölsä, E.; Miettinen, I.; Haukka, K.; Vaara, M.; Siitonen, A. Shiga toxin-producing Escherichia coli O100:H(-): stx (2e) in drinking water contaminated by waste water in Finland. Curr. Microbiol. 2011, 62, 1239-1244.
61. Ram, S.; Vajpayee, P.; Shanker, R. Contamination of potable water distribution systems by multiantimicrobial-resistant enter hemorrhagic Escherichia coli. Environ. Health Perspect. 2008, 116, 448-452.
62. Halabi, M.; Orth, D.; Grif, K.; Wiesholzer-Pittl, M.; Kainz, M.; Schöberl, J.; Dierich, M.P.; Allerberger, F.; Würzner, R. Prevalence of Shiga toxin-, intimin- and haemolysin genes in Escherichia coli isolates from drinking water supplies in a rural area of Austria. Int. J. Hyg. Environ. Health 2008, 211, 454-457.
63. McCarthy, T.A.; Barrett, N.L.; Hadler, J.L.; Salsbury, B.; Howard, R.T.; Dingman, D.W.; Brinkman, C.D.; Bibb, W.F.; Cartter, M.L. Hemolytic-uremic syndrome and Escherichia coli O121 at a Lake in Connecticut, 1999. Pediatrics 2001, 108, E59.
64. Paunio, M.; Pebody, R.; Keskimäki, M.; Kokki, M.; Ruutu, P.; Oinonen, S.; Vuotari, V.; Siitonen, A.; Lahti, E.; Leinikki, P. Swimming-associated outbreak of Escherichia coli O157:H7. Epidemiol. Infect. 1999, 122, 1-5.
65. Feldman, K.A.; Mohle-Boetani, J.C.; Ward, J.; Furst, K.; Abbott, S.L.; Ferrero, D.V.; Olsen, A.; Werner, S.B. A cluster of Escherichia coli O157: Nonmotile infections associated with recreational exposure to lake water. Public Health Rep. 2002, 117, 380-385.
66. Smith, C.J.; Olszewski, A.M.; Mauro, S.A. Correlation of Shiga toxin gene frequency with commonly used microbial indicators of recreational water quality. Appl. Environ. Microbiol. 2009, 75, 316-21.
67. Walters, S.P.; Thebo, A.L.; Boehm, A.B. Impact of urbanization and agriculture on the occurrence of bacterial pathogens and stx genes in coastal waterbodies of central California. Water Res. 2011, 45, 1752-1762.
68. Casas, V.; Miyake, J.; Balsley, H.; Roark, J.; Telles, S.; Leeds, S.; Zurita, I.; Breitbart, M.; Bartlett, D.; Azam, F. et al. Widespread occurrence of phage-encoded exotoxin genes in terrestrial and aquatic environments in Southern California. FEMS Microbiol. Lett. 2006, 261, 141-149.
69. Ram, S.; Vajpayee, P.; Shanker, R. Prevalence of multi-antimicrobial-agent resistant, Shiga toxin and enterotoxin producing Escherichia coli in surface waters of river Ganga. Environ. Sci. Technol. 2007, 41, 7383-7388.
70. Duris, J.W.; Haack, S.K.; Fogarty, L.R. Gene and antigen markers of Shiga-toxin producing E. coli from Michigan and Indiana river water: Occurrence and relation to recreational water quality criteria. J. Environ. Qual. 2009, 38, 1878-1886.
71. Kurokawa, K.; Tani, K.; Ogawa, M.; Nasu, M. Abundance and distribution of bacteria carrying sltII gene in natural river water. Lett. Appl. Microbiol. 1999, 28, 405-410.
72. Blanch, A.R.; García-Aljaro, C.; Muniesa, M.; Jofre, J. Detection, enumeration and isolation of strains carrying the stx2 gene from urban sewage. Water Sci. Technol. 2003, 47, 109-116.
73. García-Aljaro, C.; Muniesa, M.; Jofre, J.; Blanch, A.R. Prevalence of the stx2 gene in coliform populations from aquatic environments. Appl. Environ. Microbiol. 2004, 70, 3535-3540.
74. Loukiadis, E.; Kérourédan, M.; Beutin, L.; Oswald, E.; Brugère, H. Characterization of Shiga toxin gene (stx)-positive and intimin gene (eae)-positive Escherichia coli isolates from wastewater of slaughterhouses in France. Appl. Environ. Microbiol. 2006, 72, 3245-3251.
75. Chern, E.C.; Tsai, Y.L.; Olson, B.H. Occurrence of genes associated with enterotoxigenic and enterohemorrhagic Escherichia coli in agricultural waste lagoons. Appl. Environ. Microbiol. 2004, 70, 356-362.
76. Muniesa, M.; Jofre, J. Occurrence of phages infecting Escherichia coli O157:H7 carrying the Stx2 gene in sewage from different countries. FEMS Microbiol. Lett. 2000, 183, 197-200.
77. Muniesa, M.; Jofre, J. Abundance in sewage of bacteriophages that infect Escherichia coli O157:H7 and that carry the Shiga toxin 2 gene. Appl. Environ. Microbiol. 1998, 64, 2443-2448.
78. Dumke, R.; Schröter-Bobsin, U.; Jacobs, E.; Röske, I. Detection of phages carrying the Shiga toxin 1 and 2 genes in waste water and river water samples. Lett. Appl. Microbiol. 2006, 42, 48-53.
79. Muniesa, M.; Lucena, F.; Jofre, J. Comparative survival of free Shiga toxin 2-encoding phages and Escherichia coli strains outside the gut. Appl. Environ. Microbiol. 1999, 65, 5615-5618.
80. Haack, S.K.; Duris, J.W.; Fogarty, L.R.; Kolpin, D.W.; Focazio, M.J.; Furlong, E.T.; Meyer, M.T. Comparing wastewater chemicals, indicator bacteria concentrations, and bacterial pathogen genes as fecal pollution indicators. J. Environ. Qual. 2009, 38, 248-258.
81. Ishii, S.; Yan, T.; Shively, D.A.; Byappanahalli, M.N.; Whitman, R.L.; Sadowsky, M.J. Cladophora (Chlorophyta) spp. harbor human bacterial pathogens in near shore water of Lake Michigan. Appl. Environ. Microbiol. 2006, 72, 4545-4553.
82. Vanden Heuvel, A.; McDermott, C.; Pillsbury, R.; Sandrin, T.; Kinzelman, J.; Ferguson, J.; Sadowsky, M.; Byappanahalli, M.; Whitman, R.; Kleinheinz, G.T. The green alga, Cladophora, promotes Escherichia coli growth and contamination of recreational waters in Lake Michigan. J. Environ. Qual. 2009, 39, 333-344.
83. Gourmelon, M.; Montet, M.P.; Lozach, S.; Le Mennec, C.; Pommepuy, M.; Beutin, L.; Vernozy-Rozand, C. First isolation of Shiga toxin 1d producing Escherichia coli variant strains in shellfish from coastal areas in France. J. Appl. Microbiol. 2006, 100, 85-97.
84. Neely, M.N.; Friedman, D.I. Arrangement and functional identification of genes in the regulatory region of lambdoid phage H-19B, a carrier of a Shiga-like toxin. Gene 1998, 223, 105-113.
85. Wagner, P.L.; Neely, M.N.; Zhang, X.; Acheson, D.W.; Waldor, M.K.; Friedman, D.I. Role for a phage promoter in Shiga toxin 2 expression from a pathogenic Escherichia coli strain. J. Bacteriol. 2001, 183, 2081-2085.
86. Wagner, P.L.; Livny, J.; Neely, M.N.; Acheson, D.W.; Friedman, D.I.; Waldor, M.K. Bacteriophage control of Shiga toxin 1 production and release by Escherichia coli. Mol. Microbiol. 2002, 44, 957-970.
87. Neely, M.N.; Friedman, D.I. Functional and genetic analysis of regulatory regions of coliphage H-19B: Location of Shiga-like toxin and lysis genes suggest a role for phage functions in toxin release. Mol. Microbiol. 1998, 28, 1255-1267.
88. Neely, M.N.; Friedman, D.I. N-mediated transcription antitermination in lambdoid phage H-19B is characterized by alternative NUT RNA structures and a reduced requirement for host factors. Mol. Microbiol. 2000, 38, 1074-1085.
89. Calderwood, S.B.; Mekalanos, J.J. Iron regulation of Shiga-like toxin expression in Escherichia coli is mediated by the fur locus. J. Bacteriol. 1987, 169, 4759-4764.
90. Sung, L.M.; Jackson, M.P.; O'Brien, A.D.; Holmes, R.K. Transcription of the Shiga-like toxin type II and Shiga-like toxin type II variant operons of Escherichia coli. J. Bacteriol. 1990, 172, 6386-6395.
91. Roberts, J.W.; Roberts, C.W.; Craig, N.L.; Phizicky, E.M. Activity of the Escherichia coli recA-gene product. Cold Spring Harbor Symp. Quant. Biol. 1979, 43 (Pt 2), 917-920.
92. Craig, N.L.; Roberts, J.W. E. coli recA protein-directed cleavage of phage lambda repressor requires polynucleotide. Nature 1980, 283, 26-30.
93. Phizicky, E.M.; Roberts, J.W. Kinetics of RecA protein-directed inactivation of repressors of phage lambda and phage P22. J. Mol. Biol. 1980, 139, 319-328.
94. Phizicky, E.M.; Roberts, J.W. Induction of SOS functions: regulation of proteolytic activity of E. coli RecA protein by interaction with DNA and nucleoside triphosphate. Cell 1981, 25, 259-267.
95. Little, J.W. Autodigestion of lexA and phage lambda repressors. Proc. Natl. Acad. Sci. USA 1984, 81, 1375-1379.
96. Wagner, P.L.; Acheson, D.W.K.; Waldor, M.K. Human neutrophils and their products induce Shiga toxin production by Enterohemorrhagic Escherichia coli. Infect. Immunol. 2001, 69, 1934-1937.
97. Lainhart, W.; Stolfa, G.; Koudelka, G.B. Shiga toxin as a bacterial defense against a eukaryotic predator, Tetrahymena thermophila. J. Bacteriol. 2009, 191, 5116-5122.
98. Kimmitt, P.T.; Harwood, C.R.; Barer, M.R. Toxin gene expression by Shiga toxin-producing Escherichia coli: The role of antibiotics and the bacterial SOS response. Emerg. Infect. Dis. 2001, 6, 458-465.
99. Neely, M.N.; Friedman, D.I. Functional and genetic analysis of regulatory regions of coliphage H- 19B: Location of Shiga-like toxin and lysis genes suggest a role for phage functions in toxin release. Mol. Microbiol. 1998, 28, 1255-1267.
100. Wagner, P.L.; Livny, J.; Neely, M.N.; Acheson, D.W.; Friedman, D.I.; Waldor, M.K. Bacteriophage control of Shiga toxin 1 production and release by Escherichia coli. Mol. Microbiol. 2002, 44, 957-970.
101. Livny, J.; Friedman, D.I. Characterizing spontaneous induction of Stx encoding phages using a selectable reporter system. Mol. Microbiol. 2004, 51, 1691-1704.
102. Bullwinkle, T.J.; Koudelka, G.B. The Lysis-Lysogeny Decision of Bacteriophage 933W: A 933W repressor-mediated long distance loop has no role in regulating 933W PRM activity. J. Bacteriol. 2011, in press.
103. Casas, V.; Miyake, J.; Balsley, H.; Roark, J.; Telles, S.; Leeds, S.; Zurita, I.; Breitbart, M.; Bartlett, D.; Azam, F. et al. Widespread occurrence of phage-encoded exotoxin genes in terrestrial and aquatic environments in Southern California. FEMS Microbiol. Lett. 2006, 261, 141-149.
104. Matz, C.; Kjelleberg, S. Off the hook-how bacteria survive protozoan grazing. Trends Microbiol. 2005, 13, 302-307.
105. Gamage, S.D.; Strasser, J.E.; Chalk, C.L.; Weiss, A.A. Nonpathogenic Escherichia coli can contribute to the production of Shiga toxin. Infect. Immunol. 2003, 71, 3107-3115.
106. Wagner, P.L.; Neely, M.N.; Zhang, X.; Acheson, D.W.; Waldor, M.K.; Friedman, D.I. Role for a phage promoter in Shiga toxin 2 expression from a pathogenic Escherichia coli strain. J. Bacteriol. 2001, 183, 2081-2085.
107. Wildschutte, H.; Wolfe, D.M.; Tamewitz, A.; Lawrence, J.G. Protozoan predation, diversifying selection, and the evolution of antigenic diversity in Salmonella. Proc. Natl. Acad. Sci. USA 2004, 101, 10644-10649.
108. Jurgens, K.; Matz, C. Predation as a shaping force for the phenotypic and genotypic composition of planktonic bacteria. Antonie van Leeuwenhoek 2002, 81, 413-434.
109. Simek, K.; Vrba, J.; Pernthaler, J.; Posch, T.; Hartman, P.; Nedoma, J.; Psenner, R. Morphological and compositional shifts in an experimental bacterial community influenced by protists with contrasting feeding modes. Appl. Environ. Microbiol. 1997, 63, 587-595.
110. Jurgens, K.; Pernthaler, J.; Schalla, S.; Amann, R. Morphological and compositional changes in a planktonic bacterial community in response to enhanced protozoan grazing. Appl. Environ. Microbiol. 1999, 65, 1241-1250.