Enteroaggregative Escherichia coli related to uropathogenic clonal group A.Enteroaggregative Escherichia coli Escherichia coli (ĕsh'ərĭk`ēə kō`lī), common bacterium that normally inhabits the intestinal tracts of humans and animals, but can cause infection in other parts of the body, especially the urinary tract. (EAEC EAEC enteroadherent Escherichia coli. EAEC Enteroadherent Escherichia coli, see there ) are heterogeneous, diarrheagenic E. coli E. coli: see Escherichia coli. E. coli in full Escherichia coli Species of bacterium that inhabits the stomach and intestines. E. coli can be transmitted by water, milk, food, or flies and other insects. Of EAEC strains from Nigeria, 10 independent antimicrobial-resistant isolates belonged to the multilocus sequence type 69 clonal complex, to which uropathogenic E. coli clonal group A belongs. This finding suggests a recent common ancestor for these distinct groups of pathogenic E. coli. ********** Enteroaggregative Escherichia coli (EAEC) is an emerging category of diarrheagenic E. coli EAEC are heterogeneous, and the distribution of known virulence genes rarely correlates with phylogeny based on housekeeping loci loci [L.] plural of locus. loci Plural of locus, see there (1). We recently identified 2 loci, involved in iron acquisition, that are distributed among EAEC in a manner that correlates with multilocus enzyme electrophoresis typing based on 20 housekeeping enzymes (2). This finding supports the possibility that EAEC clonal groups with increased pathogenic potential exist. The Study To identify overrepresented o·ver·rep·re·sent·ed adj. Represented in excessive or disproportionately large numbers: "Some groups, and most notably some races, may be overrepresented and others may be underrepresented" subgroups of potential clonal origin, we examined 131 EAEC strains isolated from children in Nigeria (3), 73 from 187 children with diarrhea and 58 from 144 healthy control participants. All 131 strains had previously been categorized as EAEC by the standard HEp-2 cell adherence assay (3,4). To determine flagellin flagellin /fla·gel·lin/ (flah-jel´in) a protein of bacterial flagella; it is composed of subunits in several-stranded helical arrangement. types, we used an RsaI-based PCR-restriction fragment length polymorphism polymorphism, of minerals, property of crystallizing in two or more distinct forms. Calcium carbonate is dimorphous (two forms), crystallizing as calcite or aragonite. Titanium dioxide is trimorphous; its three forms are brookite, anatase (or octahedrite), and rutile. (RFLP RFLP abbr. restriction fragment length polymorphism RFLP restriction fragment length polymorphism. RFLP ) protocol. Using primers FFLIC FFLIC Families & Friends of Louisiana’s Incarcerated Children 1 (5'-ATGGCACAAGTCATTAATACCCAAC-3') and R-FLIC2 (5'-CTAACCCTGCAGCAGAGACA-3'), we obtained an internal fliC amplicon from 105 (80.1%) of the 131 isolates. RFLP analysis delineated [greater than or equal to] 31 flagellin genotypes among these amplicons (online Appendix Table, available at www.cdc.gov/EID/content/13/5/757-appT. htm). Using 16 control strains, which represented 12 antigenically distinct H-types verified at reference centers, we could associate a specific H antigen H antigen n. See flagellar antigen. H antigen see H antigen. H antigen Transfusion medicine The trisaccharide stem chain of the ABO blood group, located on RBC membranes; the enzyme, with 10 of these RFLP patterns. The most common pattern, which corresponded with the H18 antigen, accounted for 18 (17%) of the genotyped isolates (or 14% of all isolates). Two H7 control strains had different genotypes, which indicates that the products of 2 different alleles are recognized by anti-H7 antiserum antiserum /an·ti·se·rum/ (an´ti-se?rum) a serum containing antibody(ies), obtained from an animal immunized either by injection of antigen or by infection with microorganisms containing antigen. . Both genotypes were detected among the test EAEC strains (online Appendix Table) and together accounted for 10 (7.6%) of the isolates. Other predominant fliC variants were H2 (3%), H11 (3.8%), H21 (7.6%), and H45 (4.9%). Although somefliC variants were somewhat more common among strains from children with diarrhea than from control participants (online Appendix Table), differences were not statistically significant (p>0.05, Fisher exact test). Antimicrobial susceptibility profiles were determined by disk diffusion as specified by the Clinical and Laboratory Standards Institute (5). Disks containing ampicillin ampicillin (ăm'pĭsĭl`ĭn), a penicillin-type antibiotic that is effective against both gram-negative microorganisms and gram-positive microorganisms such as Escherichia coli. (10 lag), tetracycline tetracycline (tĕ'trəsī`klēn), any of a group of antibiotics produced by bacteria of the genus Streptomyces. They are effective against a wide range of Gram positive and Gram negative bacteria, interfering with protein (30 [micro]g), trimethoprim trimethoprim /tri·meth·o·prim/ (-meth´o-prim) an antibacterial closely related to pyrimethamine; almost always used in combination with a sulfonamide, primarily for the treatment of urinary tract infections. (5 [micro]g), nalidixic acid nalidixic acid /nal·i·dix·ic ac·id/ (nal-i-dik´sik) a synthetic antibacterial agent used in the treatment of genitourinary infections caused by gram-negative organisms. na·li·dix·ic acid n. (30 [micro]g), chloramphenicol chloramphenicol (klōr'ămfĕn`əkŏl'), antibiotic effective against a wide range of gram-negative and gram-positive bacteria (see Gram's stain). It was originally isolated from a species of Streptomyces bacteria. (30 [micro]g), sulfonamide sulfonamide /sul·fon·amide/ (sul-fon´ah-mid) a compound containing the sbondSO2NH2 group. The sulfonamides, or sulfa drugs, are derivatives of sulfanilamide, competitively inhibit folic acid synthesis in microorganisms, and formerly were (300 [micro]g), streptomycin streptomycin (strĕp'tōmī`sĭn), antibiotic produced by soil bacteria of the genus Streptomyces and active against both gram-positive and gram-negative bacteria (see Gram's stain), including species resistant to other (10 [micro]g), and ciprofloxacin ciprofloxacin /cip·ro·flox·a·cin/ (sip?ro-flok´sah-sin) a synthetic antibacterial effective against many gram-positive and gram-negative bacteria; used as the hydrochloride salt. cip·ro·flox·a·cin n. (5 [micro]g) were used for testing on Mueller-Hinton agar (Oxoid, Lenexa, Kansas Lenexa is a city in the central part of Johnson County, located in Northeast Kansas, in the Central United States. The population was estimated to be 43,434 in the year 2005.[] It is the fourth most populous city in the county. , USA). Of the 18 Nigerian H18 strains, 13 had the same resistance pattern: ampicillin, chloramphenicol, streptomycin, sulfonamide, tetracycline, trimethoprim (Table 1). PCR PCR polymerase chain reaction. PCR abbr. polymerase chain reaction Polymerase chain reaction (PCR) identified 3 EAEC-specific and 2 other virulence genes (2, 6). Markers of well-characterized aggregative adherence plasmids, present in typical EAEC strains, are aggregative adherence regulator gene regulator gene n. A gene that causes the production of a protein that represses the activity of another gene in an operon. aggR, anti-aggregative protein or dispersin gene aap, and empiric plasmid probe (CVD CVD Cardiovascular disease, see there 432) that represents part of the aat secretion system operon (6). Of the 131 isolates, <30% harbor these loci (7). However, 17 (89%) of 18 H18-positive EAEC isolates harbored [greater than or equal to] 1 of these aggregative adherence plasmid loci, and 15 (83%) of 17 harbored all 3 loci (Table 1). Moreover, 15 (83%) of the 18 isolates contained iucA (aerobactin synthesis), and 17 (94%) contained chuA (heme transport outer-membrane receptor), significantly more than the other 113 EAEC isolates (and 39.8% and 24.8% for iucA and chuA, respectively; p<0.001 for each). Multidrug-resistant, chuA-positive E. coli H18 strains are also frequently recovered from patients with urinary tract infection urinary tract infection (UTI), n infection in one or more of the structures that make up the urinary system. Occurs more often in women and is most commonly caused by bacteria. . Some of these strains derive from the successful and globally disseminated multidrug-resistant clonal group A (CGA (Color/Graphics Adapter) The first video display standard for the IBM PC. This low-resolution system was superseded by EGA and then VGA. CGA required a digital RGB Color Display monitor. See PC display modes. CGA - Color Graphics Adapter ) (8). Uropathogenic E. coli (UPEC UPEC Uropathogenic Escherichia Coli UPEC Upper Peninsula Environmental Coalition UPEC University Professional Education Council ) clonal group A strains occur in the United States United States, officially United States of America, republic (2005 est. pop. 295,734,000), 3,539,227 sq mi (9,166,598 sq km), North America. The United States is the world's third largest country in population and the fourth largest country in area. and Europe; typically exhibit serotypes O11:H18, O17:H18, O73:H18, or O77:H18; and share a common resistance and repetitive element. PCR profile (8). Recently, at certain US centers, [greater than or equal to] 33% of trimethoprim-resistant E. coli isolates from uncomplicated cases of pyelonephritis pyelonephritis: see nephritis. pyelonephritis Infection (usually bacterial) and inflammation of kidney tissue and the renal pelvis. Acute pyelonephritis is usually localized and may have no apparent cause. and cystitis cystitis (sĭstī`tĭs), common acute or chronic inflammation of the urinary bladder. The disease occurs primarily in young women and frequently results from bacterial invasion of the urethra from the adjacent rectum, most commonly with have represented CGA (9). CGA strains also can infect nonurinary, extraintestinal sites (10). Furthermore, CGA-like strains have been recovered from human and animal feces, which implies a commensal commensal /com·men·sal/ (kom-men´sil) 1. living on or within another organism, and deriving benefit without harming or benefiting the host. 2. a parasite that causes no harm to the host. reservoir (11,12). Accordingly, we assessed our H18 EAEC isolates for membership in CGA. A CGA-specific PCR protocol, which yields a 175-bp PCR product in strains that have 3 single-nucleotide polymorphisms Within fumC (13), was applied to the Nigerian EAEC H18 isolates. Positive controls were 2 reference CGA cystitis isolates with UMN UMN upper motor neuron. 026 and SEQ SEQ Sequence SEQ Sequential SEQ South East Queensland (Australia) SEQ Smart Equities Conference SEQ Sequens/Sequentes SEQ Senior Enlisted Quarters SEQ Short Essay Question SEQ Stigmatisation and Eczema Questionnaire SEQ Scientific Equipment 102 (ATCC ATCC American Type Culture Collection, see there BAA-457) (13); negative controls were non-CGA cystitis isolate 2P9 (O15:K52:H1), UPEC isolates CFT CFT complement fixation test; see under fixation. CFT complement fixation test. 073 (O6: K2:H1) and 536 (O6:K15:H31), and an H18 enterotoxigenic en·ter·o·tox·i·gen·ic adj. Of or being an organism containing or producing an enterotoxin. Enterotoxigenic E. coli (ETEC ETEC enterotoxigenic Escherichia coli. ETEC Enterotoxic Escherichia coli, see there ) isolate from the Nigeria study (14). Of the 18 EAEC H18 isolates, 10 (including 8 from children with diarrhea) were positive, whereas 8 (and the H18 ETEC isolate) were negative. Lack of obvious familial or temporal clustering of patients from whom these strains were isolated suggests that the isolates are not likely to be directly linked through a single point source. Of 21 other EAEC strains from diverse non-African locales that were similarly screened, including 3 isolates bearing the H18 fliC allele allele (əlēl`): see genetics. allele Any one of two or more alternative forms of a gene that may occur alternatively at a given site on a chromosome. (1), only Thai isolate 144-1 (H18-positive) (1) exhibited the CGA-specific fumC single-nucleotide polymorphisms. The 2 other non-Nigerian H18 EAEC (Peruvian O44:H18 isolate 042 and Thai O36:H18 isolate 44-1) (1) were negative. To unequivocally assess clonal relationships, we subjected all H18 Nigerian EAEC isolates and the Thai isolate 144-1 to multilocus sequence typing Multilocus sequence typing (MLST) is a technique in molecular biology for the typing of multiple loci. The procedure characterizes isolates of bacterial species using the DNA sequences of internal fragments of multiple (usually seven) housekeeping genes. (MLST MLST Multi Locus Sequence Typing MLST Medical Logistics Support Team MLST Mini Losi Super Truck (1/18th scale radio control vehicle) ) (15). Briefly, we sequenced designated internal regions of the adk, fumC, gyrB, icd, mdh, purA, and recA genes. Allele comparisons and sequence type (ST) assignments were done by using the open-source E. coli MLST database (http://web.mpiibberlin.mpg.de/mlst/dbs/Ecoli). All 11 putative EAEC CGA isolates carried fumC allele 35, which has all 3 targets of the CGA single-nucleotide polymorphism screen (G270A, C271T, and C288T) (13). Of these isolates, 10 (9 Nigerian, 1 Thai) belonged to ST394, which shares 5 of 7 alleles with ST69 (the predominant ST of UPEC CGA; [12]) and, according to according to prep. 1. As stated or indicated by; on the authority of: according to historians. 2. In keeping with: according to instructions. 3. the e-BURST algorithm, is placed with ST69 in the same larger ST69 complex, indicative of a recent common ancestor (15). One Nigerian H18 EAEC isolate, strain E23, was assigned a new ST, ST432, because of its novel purA allele; however, ST432 shares alleles with ST394 at all 6 other loci and so also belongs with the ST69 complex. Of the 8 H18 isolates that were negative in the fumC35 single-nucleotide polymorphism assay, 7 belonged to STs not previously described. Only 1 of these, strain E62 (ST471), shared 6 alleles with ST394 and 4 with ST69. Another strain, C16 (ST512), shared 5 alleles with ST69 and only 4 with ST394. Of the 8fumC35-negative H18 isolates, 6 did not share 5 alleles with ST69 or ST394 and are therefore considered to be of a different clonal complex. Six fumC35-negative H18 isolates had 6 alleles in common and belonged to 1 of 3 STs: 31,449, or 474. ST31 includes other EAEC in the MLST database (www.mlst.net). One EAEC H18 isolate shared no allele with any other EAEC isolate from this study (Table 2). Although iron-utilization genes chuA and iucA are present in all EAEC and UPEC ST69 complex strains, the EAEC virulence plasmid markers aggR, aap, and CVD432 (aat) were found in all 11 EAEC ST394/432 isolates but not in reference UPEC CGA (ST69) isolates (Table 1). EAEC ST394/432 strains and UPEC ST69 (CGA) strains appear to represent diverging lineages of common ancestry, which are adapting to separate niches. Escobar-Paramo (14) proposed that certain E. coli backgrounds appear to be more likely to acquire virulence genes. Our findings suggest that the ST69 complex progenitor pro·gen·i·tor n. 1. A direct ancestor. 2. An originator of a line of descent. progenitor ancestor, including parent. progenitor cell stem cells. , from which CGA UPEC and ST394 EAEC are derived, may have had a propensity to acquire virulence genes as well as antimicrobial resistance elements, thereby generating at least 2 clonal groups pathogenic for humans, with several nonoverlapping, horizontally acquired virulence factors. CGA-like strains recently isolated from animal feces and food samples have been proposed by Ramchandani et al. (11) as possible reservoirs for UPEC CGA strains. However, although they found similar resistance patterns and serotypes among animal CGA isolates as among humans, they did not find typical UPEC-associated virulence gene profiles (11). Tartof et al. (12) have subsequently shown that CGA-like strains from animal or environmental sources do not belong to ST69 (as do most human UPEC CGA isolates) but that they are part of the ST69 complex, particularly the ST394 type, which corresponds to the EAEC H18 clonal group we describe. Nonhuman ST394 isolates could possibly represent CGA-like EAEC and point to potential nonhuman reservoirs of EAEC, which remain to be identified. Conclusions ST69 and ST394 appear to represent successful, genetically related lineages; isolates belonging to both are commonly resistant to ampicillin, chloramphenicol, streptomycin, sulfonamides Sulfonamides Definition Sulfonamides are medicines that prevent the growth of bacteria in the body. Purpose Sulfonamides are used to treat many kinds of infections caused by bacteria and certain other microorganisms. , tetracycline, and trimethoprim. Widespread use of trimethoprim-sulfamethoxazole has been proposed as a reason for the emergence and spread of UPEC CGA (8). This combination, as well as other drugs to which ST394 strains are typically resistant, is commonly used (and misused) in Nigeria and other developing countries and could provide selective pressure for EAEC ST394/432. Although our numbers were too small to significantly associate ST394/432 strains with disease, 8 of 10 of these isolates were from children with diarrhea. Our study has unveiled what we believe to be a previously unrecognized EAEC clonal group. The fumC single-nucleotide polymorphism method, proposed for identifying UPEC CGA, could be useful for assessing its distribution. Acknowledgments We thank Team 56 at the Wellcome Trust The Wellcome Trust is a United Kingdom-based charity established in 1936 to administer the fortune of the American-born pharmaceutical magnate Sir Henry Wellcome. Its income was derived from what was originally called Burroughs Wellcome & Co, later renamed in the UK as the Sanger Institute The Wellcome Trust Sanger Institute (formerly the Sanger Centre) is a genome research centre in Cambridgeshire, England. It was set up in 1992 by the Wellcome Trust and the Medical Research Council, the purpose of which is stated on their website ([1] as "to further our , particularly Nick Matthews, for sequencing. Claire-Louise Crichton, Sheraz Said, and Amanda Muir provided other technical assistance. This work was funded by a Branco Weiss Fellowship from the Society in Science, Zurich, to I.N.O.F.W.-G. was supported by a Bryn Mawr College Bryn Mawr College, at Bryn Mawr, Pa; undergraduate for women, graduate coeducational; opened 1885 by the Society of Friends, with a bequest from Joseph W. Taylor of Burlington, N.J. Modeled on a group curriculum plan at Johns Hopkins Univ. Mellon Mays Undergraduate fellowship and a Haverford College Haverford College Private liberal arts college in Haverford, Pa., near Philadelphia. Founded by Quakers in 1833 as a men's college, it became coeducational in 1980. It is consistently ranked as one of the top U.S. colleges. Undergraduate Howard Hughes Medical Institute Howard Hughes Medical Institute, (HHMI), nonprofit medical research organization founded in 1953 by Howard Hughes and largly funded from proceeds of the 1984–85 sale of Hughes Aircraft. Headquartered in Chevy Chase, Md. Multicultural scholarship. J.W. was funded by the Wellcome Trust of Great Britain Great Britain, officially United Kingdom of Great Britain and Northern Ireland, constitutional monarchy (2005 est. pop. 60,441,000), 94,226 sq mi (244,044 sq km), on the British Isles, off W Europe. The country is often referred to simply as Britain. and J.R.J. by the Office of Research and Development, Medical Research Service, Department of Veteran Affairs. J.R.J. has received research grants and consultancies from Ortho-McNeil, Bayer, Wyeth-Ayerst, Merck, Procter & Gamble, and Rochester Medical. References (1.) Czeczulin JR, Whittam TS, Henderson IR, Navarro-Garcia F, Nataro JP. Phylogenetic phy·lo·ge·net·ic adj. 1. Of or relating to phylogeny or phylogenetics. 2. Relating to or based on evolutionary development or history. analysis of virulence genes in enteroaggregative and diffusely adherent adherent /ad·her·ent/ (-ent) sticking or holding fast, or having such qualities. Escherichia coli. Infect Immun. 1999;67: 2692-9. (2.) Okeke IN, Scaletsky IC, Soars EH, Macfarlane MacFarlane or Macfarlane is a surname shared by:
(3.) Okeke IN, Lamikanra A, Czeczulin J, Dubovsky F, Kaper JB, Nataro JP. Heterogeneous virulence of enteroaggregative Eschericihia coli strains isolated from children in Southwest Nigeria. J Infect Dis. 2000;181:252 60. (4.) Fields PI, Blom K, Hughes HJ, Helsel LO, Feng P, Swaminathan B. Molecular characterization of the gene encoding H antigen in Escherichia coli and development of a PCR-restriction fragment length polymorphism test for identification of E. coli O157:H7 and O157: NM. J Clin Microbiol. 1997;35:1066-70. (5.) National Committee for Clinical Laboratory Standards. Performance standards for antimicrobial disk susceptibility tests, 8th edition; Approved standard. Villanova (PA): The Committee; 2003. p. 130. (6.) Cerna JF, Nataro JP, Estrada-Garcia T. Multiplex PCR for detection of three plasmid-borne genes of enteroaggregative Escherichia coli strains. J Clin Microbiol. 2003;41:2138-40. (7.) Okeke IN, Lamikanra A, Steinruck H, Kaper JB. Characterization of Escherichia coli strains from cases of childhood diarrhea in provincial southwestern Nigeria. J Clin Microbiol. 2000;38:7-12. (8.) Johnson JR, Manges AR, O'Bryan TT, Riley LW. A disseminated multidrug-resistant clonal group of uropathogenic Eseherichia coli in pyelonephritis. Lancet. 2002;359:2249-51. (9.) Manges AR, Johnson JR, Foxman B, O'Bryan TT, Fullerton KE, Riley LW. Widespread distribution of urinary tract infections caused by a multidrug-resistant Escherichia coli clonal group. N Engl J Med. 2001;345:1007 13. (10.) Manges AR, Perdreau-Remington F, Solberg O, Riley LW. Multidrug-resistant Escherichia coli clonal groups causing community-acquired bloodstream infections. J Infect. 2005;53:25-9. (11.) Ramchandani M, Manges AR, DebRoy C, Smith SP, Johnson JR, Riley LW. Possible animal origin of human-associated, multidrug-resistant, uropathogenic Escherichia coli. Clin Infect Dis. 2005;40:251-7. (12.) Tartof SY, Solberg OD, Manges AR, Riley LW. Analysis of a uropathogenic Escherichia coli clonal group by multilocus sequence typing. J Clin Microbiol. 2005;43:5860-4. (13.) Johnson JR, Owens K, Manges AR, Riley LW. Rapid and specific detection of Escheriehia coli clonal group A by gene-specific PCR. J Clin Microbiol. 2004;42:2618-22. (14.) Escobar-Paramo P, Clermont O, Blanc-Potard AB, Bui H, Le Bouguenec C, Denamur E. A specific genetic background is required for acquisition and expression of virulence factors in Escherichia coli. Mol Biol Evol. 2004;21:1085 94. (15.) Wirth T, Falush D, Lan R, Colles F, Mensa MENSA. This comprehends all goods and necessaries for livelihood. Obsolete. P, Wieler LH, et al. Sex and virulence in Escherichia coli: an evolutionary perspective. Mol Microbiol. 2006;60:1136 51. Ms Wallace-Gadsden is a graduate student in the department of microbiology and molecular biology molecular biology, scientific study of the molecular basis of life processes, including cellular respiration, excretion, and reproduction. The term molecular biology was coined in 1938 by Warren Weaver, then director of the natural sciences program at the Rockefeller at Tufts University Tufts University, main campus at Medford, Mass.; coeducational; chartered 1852 by Universalists as a college for men. It became a university in 1955. Jackson College, formerly a coordinate undergraduate college for women, merged with the College of Liberal Arts in . Her main interests are bacterial colonization and pathogenesis. Address for correspondence: Iruka N. Okeke, Department of Biology, Haverford College, 370 Lancaster Ave, Haverford, PA 19041, USA; email: iokeke@haverford.edu Faith Wallace-Gadsden, * (1) James R. Johnson, ([dagger]) ([double dagger double dagger n. A reference mark (  ) used in printing and writing. Also called diesis.Noun 1. ]) John Wain Noun 1. John Wain - English writer (1925-1994) John Barrington Wain, Wain , ([section]) and Iruka N. Okeke * * Haverford College, Haverford, Pennsylvania Haverford is an unincorporated community located mostly in Haverford Township in Delaware County, but also partially in Lower Merion Township in Montgomery County, about 10 miles west of Philadelphia. It is on the Main Line which is known for its large amount of money. , USA; ([dagger]) Minneapolis Veterans Administration Medical Center, Minneapolis, Minnesota “Minneapolis” redirects here. For other uses, see Minneapolis (disambiguation). Minneapolis (pronounced IPA: /ˌmɪniˈæpəlɪs/) is the largest city in the U.S. , USA; ([double dagger]) University of Minnesota (body, education) University of Minnesota - The home of Gopher. http://umn.edu/. Address: Minneapolis, Minnesota, USA. , Minneapolis, Minnesota, USA; and ([section]) Wellcome Trust Sanger Institute, Cambridgeshire, United Kingdom (1) Current affiliation: Tufts University School of Medicine The Tufts University School of Medicine is one of the eight schools that comprise Tufts University. Located on the university's health sciences campus in the Chinatown district of Boston, Massachusetts, the medical school has clinical affiliations with thousands of doctors and , Boston, Massachusetts, USA
Table 1. Properties of H18 EAEC isolates and selected reference
strains *
Country of Clinical
Strain isolation condition Serotype
C08 Nigeria Diarrhea 086:H18
C14 Nigeria Diarrhea N/K
E23 Nigeria Diarrhea N/K
E30 Nigeria Diarrhea N/K
G10 Nigeria Diarrhea N/K
G17a Nigeria Diarrhea N/K
G59 Nigeria Diarrhea N/K
G67b Nigeria Diarrhea N/K
C16 Nigeria Diarrhea N/K
G55 Nigeria Diarrhea N/K
E64 Nigeria Healthy N/K
G108 Nigeria Healthy N/K
E56 Nigeria Healthy N/K
E62 Nigeria Healthy N/K
E68 Nigeria Healthy N/K
G103 Nigeria Healthy N/K
G121a Nigeria Healthy N/K
G149 Nigeria Healthy N/K
042 Peru Diarrhea 044:H18
44-1 Thailand Diarrhea 036:H18
144-1 Thailand Diarrhea 077:NM
E02 Nigeria Diarrhea Ont:H18
DH5[alpha]([dagger]) N/A N/A N/A
2P9 ([double dagger]) USA UTI 015:K52:H1
SEQ102 ([double dagger]) USA UTI 011:NT
UMN026 ([double dagger]) USA UTI 017:K52:H18
CFT073 ([double dagger]) USA UTI 06:K2:H1
CGA
(fUMC
Strain Motility SNP)
C08 + +
C14 + +
E23 + +
E30 + +
G10 + +
G17a - +
G59 - +
G67b + +
C16 + -
G55 + -
E64 + +
G108 + +
E56 + -
E62 - -
E68 + -
G103 + -
G121a + -
G149 + -
042 + -
44-1 + -
144-1 - +
E02 + -
DH5[alpha]([dagger]) + -
2P9 ([double dagger]) N/D -
SEQ102 ([double dagger]) N/D +
UMN026 ([double dagger]) N/D +
CFT073 ([double dagger]) N/D -
Strain Resistance pattern
C08 Ap St Su Tc Tp
C14 ApCmStSuTcTp
E23 ApCmStSuTcTp
E30 Ap Cm St Su Tc Tp
G10 Ap Cm St Su Tp
G17a Ap Cm St Su Tc Tp
G59 Ap Cm St Su Tc Tp
G67b Ap Cm St Su TcTp
C16 Ap Cm St Su Tc Tp
G55 Ap Cm St Su Tc Tp
E64 Ap Cm St Su Tc Tp
G108 Ap Cm St Su Tc Tp
E56 Ap Cm St Su Tc
E62 Tc
E68 Ap Cm St Su TcTp
G103 ApCmStSuTcTp
G121a Ap Cm St Su Tc Tp
G149 Ap St Su Tc Tp
042 Cm St Su Tc Tp
44-1 Ap Cm St Su Tc
144-1 Cm
E02 ApCmStSuTcTp
DH5[alpha]([dagger]) -
2P9 ([double dagger]) St
SEQ102 ([double dagger]) ApCmStSuTcTp
UMN026 ([double dagger]) Ap Cm St Tc Tp
CFT073 ([double dagger]) -
CVD432
Strain aggR aap (aat)
C08 + + +
C14 + + +
E23 + + +
E30 + + +
G10 + + +
G17a + + +
G59 + + +
G67b + + +
C16 + + +
G55 - - -
E64 + + +
G108 - - +
E56 + + +
E62 + + +
E68 - - +
G103 - - +
G121a + + +
G149 + + +
042 + + +
44-1 + + +
144-1 + + +
E02 - - -
DH5[alpha]([dagger]) - - -
2P9 ([double dagger]) - - -
SEQ102 ([double dagger]) - - -
UMN026 ([double dagger]) - - -
CFT073 ([double dagger]) - - -
Strain astA chuA iucA
C08 - + +
C14 + + +
E23 - + +
E30 - + +
G10 - + +
G17a - + +
G59 - + +
G67b - + +
C16 - + -
G55 - -
E64 + + +
G108 + + +
E56 - + +
E62 - + +
E68 + + +
G103 + + +
G121a - + +
G149 - + +
042 - + -
44-1 - + +
144-1 - + +
E02 - + -
DH5[alpha]([dagger]) - - -
2P9 ([double dagger]) - + +
SEQ102 ([double dagger]) -
UMN026 ([double dagger]) - + +
CFT073 ([double dagger]) - + +
* EAEC, enteroaggregative Escherichia coli; CGA, clonal group A;
SNP, single nucleotide polymorphism; Ap, ampicillin; St, streptomycin;
Tc, tetracycline; Tp, trimethoprim; N/K, not known; Cm,
chloramphenicol; N/A, not applicable; UTI, urinary tract infection;
N/D, not Su, sulfonamide; determined. All strains were susceptible to
nalidixic acid and ciprofloxacin. All pathotypes are EAEC (H18) unless
otherwise noted.
([dagger]) Pathotype K-12.
([double dagger]) Pathotype uropathogenic E. coli.
Table 2. Multilocus sequence types of CGA-associated fumd single-
nucleotide polymorphism-positive strains *
Clinical condition,
Strain country adk fUMC
C08 Diarrhea, Nigeria 21 35
C14 Diarrhea, Nigeria 21 35
E23 Diarrhea, Nigeria 21 35
E30 Diarrhea, Nigeria 21 35
G10 Diarrhea, Nigeria 21 35
G17a Diarrhea, Nigeria 21 35
G59 Diarrhea, Nigeria 21 35
G67b Diarrhea, Nigeria 21 35
E64 Healthy, Nigeria 21 35
G108 Healthy, Nigeria 21 35
144-1 Diarrhea, Thailand 21 35
SEQ102 ([dagger]) UTI (CGA), USA 21 35
C16 Healthy, Nigeria 21 22
G55 Healthy, Nigeria 6 4
E56 Healthy, Nigeria 18 22
E62 Healthy, Nigeria 21 125
E68 Healthy, Nigeria 18 22
G103 Healthy, Nigeria 18 22
G121a Healthy, Nigeria 18 22
G149 Healthy, Nigeria 18 22
Allele profile
Strain gyrB icd mdh purA
C08 61 52 5 5
C14 61 52 5 5
E23 61 52 5 72
E30 61 52 5 5
G10 61 52 5 5
G17a 61 52 5 5
G59 61 52 5 5
G67b 61 52 5 5
E64 61 52 5 5
G108 61 52 5 5
144-1 61 52 5 5
SEQ102t 27 6 5 5
C16 2 6 5 5
G55 33 1 20 12
E56 17 6 5 5
E62 61 52 5 5
E68 94 6 5 5
G103 94 6 5 5
G121a 1 6 5 5
G149 1 6 5 5
Alele profile
Strain recA ST ST complex
C08 4 394 69
C14 4 394 69
E23 4 432 69
E30 4 394 69
G10 4 394 69
G17a 4 394 69
G59 4 394 69
G67b 4 394 69
E64 4 394 69
G108 4 394 69
144-1 4 394 69
SEQ102t 4 69 69
C16 4 512 Unassigned
G55 7 423 Unassigned
E56 4 31 31
E62 4 471 Unassigned
E68 4 449 Unassigned
G103 4 449 Unassigned
G121a 4 474 Unassigned
G149 4 474 Unassigned
* GCA, cional group A; ST, sequence type; UTI, urinary tract infection.
([dagger]) Data from reference (14). Data for all other isolates are
from this study.
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