Campylobacter jejuni multilocus sequence types in humans, northwest England, 2003-2004.Detailed understanding of the epidemiology of Campylobacter Campylobacter Genus of gram-negative spiral-shaped bacteria infecting mammals. Many species, especially C. fetus, cause miscarriage in sheep and cattle. C. jejuni is a common cause of food poisoning. Sources include meats (particularly chicken) and unpasteurized milk. is increasingly facilitated through use of universal and reproducible techniques for accurate strain differentiation and subtyping. 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) ) enables discriminatory subtyping and grouping of isolate types into genetically related clonal complexes; it also has the advantage of ease of application and repeatability. Recent studies suggest that a measure of host association may be distinguishable with this system. We describe the first continuous population-based survey to investigate the potential of MLST to resolve questions of campylobacteriosis epidemiology. We demonstrate the ability of MLST to identify variations in the epidemiology of campylobacteriosis between distinct populations and describe the distribution of key subtypes of interest. ********** Campylobacter has been the most commonly reported bacterial enteric enteric /en·ter·ic/ (en-ter´ik) within or pertaining to the small intestine. en·ter·ic adj. 1. Of, relating to, or within the intestine. 2. pathogen Pathogen Any agent capable of causing disease. The term pathogen is usually restricted to living agents, which include viruses, rickettsia, bacteria, fungi, yeasts, protozoa, helminths, and certain insect larval stages. causing gastrointestinal illness in England and Wales England and Wales are both constituent countries of the United Kingdom, that together share a single legal system: English law. Legislatively, England and Wales are treated as a single unit (see State (law)) for the conflict of laws. for at least the last 15 years (1). The main species infecting humans are Campylobacter jejuni Campylobacter jejuni Vibrio jejuni, Campylobacter fetus ssp jejuni A curved or spiral gram-negative bacillus with a single polar flagellum Epidemiology Linked to contact with domestic and farm animals, unpasteurized milk, primates, day care and C. coli; these species also colonize col·o·nize v. col·o·nized, col·o·niz·ing, col·o·niz·es v.tr. 1. To form or establish a colony or colonies in. 2. To migrate to and settle in; occupy as a colony. 3. many different animals, especially birds (2). In Europe, campylobacteriosis shows a marked seasonality with a peak during the summer months (3,4), although this pattern is more marked in some countries than others (5). Especially sharp and annually consistent rises in incidence are reported in the United Kingdom, Greece, the Netherlands, and Denmark (5). A study in northwest (NW) England also indicated a consistent peak of human infection in March (6). Some studies have shown a coincident co·in·ci·dent adj. 1. Occupying the same area in space or happening at the same time: a series of coincident events. See Synonyms at contemporary. 2. seasonality of infection in broiler broiler a young (about 8 weeks old) male or female chicken weighing 3 to 3.5 lb. chickens and humans in Scandinavia and contamination of retail raw chicken and infection in humans in the United Kingdom and suggest a common environmental trigger (2,7,8). In a recent study of the influence of rainfall, sunshine, and temperature on seasonality in different regions of England The region, also known as Government Office Region, is currently the highest tier of local government sub-national entity of England in the United Kingdom. History and Wales Wales, Welsh Cymru, western peninsula and political division (principality) of Great Britain (1991 pop. 2,798,200), 8,016 sq mi (20,761 sq km), west of England; politically united with England since 1536. The capital is Cardiff. , increasing incidence of campylobacteriosis was most strongly correlated with increases in air temperature (9). Studies in northern Europe have attempted to identify environmental reservoirs of infection in water sources and livestock that could explain the seasonality of human infection. These studies have demonstrated campylobacter carriage rates peaking in late spring and summer in broiler chicken flocks (10,11) and dairy cattle (12) but more constant infection in lambs and beef cattle (12,13). Campylobacter has been successfully isolated and cultured from surface water in Finland (14,15), Italy (16), and NW England (17), and sporadic campylobacteriosis (illness not associated with an outbreak) has been linked with exposure to untreated water in Scandinavia (18,19). C. jejuni has also been isolated from a wide range of animal and environmental samples in a rural area in NW England, which suggests a potential environmental risk for exposure (20). Molecular subtyping methods including pulsed-field gel electrophoresis gel electrophoresis n. Electrophoresis performed in a gel composed of agarose, polyacrylamide, or starch. (PFGE PFGE Pulsed-Field Gel Electrophoresis ) (21), fluorescent amplified fragment length polymorphisms Amplified fragment length polymorphism PCR, or "AFLP-PCR" (often AFLP), is a tool used in the study of genetics and in the practice of genetic engineering. Amplified Fragment Length Polymorphism (AFLP (22), and multilocus sequence typing (MLST) (23) have been applied to C. jejuni to overcome the problems associated with traditional phenotypic phe·no·type n. 1. a. The observable physical or biochemical characteristics of an organism, as determined by both genetic makeup and environmental influences. b. methods. Although PFGE has been accurately and reproducibly used for several years to investigate disease clusters (24), an advantage of MLST is its ability to sequence isolate types and thus group them into genetically related clonal complexes; this is especially useful for integrating newly identified sequence types. MLST has also provided conceptual advances in understanding the population biology Population biology is a study of biological populations of organisms, especially in terms of biodiversity, evolution, and environmental biology. Malthus can almost be considered an early population biologist, even though his training was in economics and the term population and epidemiology of C. jejuni (25). Previous studies that used phenotypic strain characterization methods failed to establish source/host associations for particular phenotypes (26), but recent studies with MLST, including a study in NW England, suggest that a measure of host association may be distinguishable when this system is used (27). No continuous population-based survey has been performed to investigate the potential of MLST to answer the unresolved questions of campylobacteriosis epidemiology, particularly the drivers of the seasonal peak in the United Kingdom. This article is the first report of a 3-year study that used MLST to investigate campylobacteriosis in NW England in 2 defined human populations, 1 small town-based and rural and 1 metropolitan and suburban. Methods Study Population The study population was defined as all persons with confirmed Campylobacter from April 2003 to March 2004 reported by residents in 4 local authorities (government administrative boundaries) in NW England (Figure 1). Wyre (population 106,826) and Fylde (74,032) local authorities adjoin geographically in the county of Lancashire and largely consist of rural and small town-based populations. For the purposes of this study, this area is referred to as rural. Salford (216,178) and Trafford (209,760) local authorities also adjoin geographically within the conurbation of Greater Manchester Greater Manchester, former metropolitan county, 497 sq mi (1,288 sq km), W central England. It comprised ten administrative districts: Bolton, Bury, Manchester, Oldham, Rochdale, Salford, Stockport, Tameside, Trafford, and Wigan. and are predominantly a mix of metropolitan and suburban districts. This area is referred to as suburban in this study. The 2 areas are [approximately equal to] 50 km apart and share some of the same drinking water drinking water supply of water available to animals for drinking supplied via nipples, in troughs, dams, ponds and larger natural water sources; an insufficient supply leads to dehydration; it can be the source of infection, e.g. leptospirosis, salmonellosis, or of poisoning, e.g. sources but are environmentally located in different water catchment areas catchment area or drainage basin, area drained by a stream or other body of water. The limits of a given catchment area are the heights of land—often called drainage divides, or watersheds—separating it from neighboring drainage . [FIGURE 1 OMITTED] Data Collection Confirmed cases of campylobacteriosis (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 UK National Standard Method for diagnosis [28]) are routinely reported to the NW Health Protection Agency (HPA (1) (High Performance Addressing) Refers to a variety of earlier addressing techniques that improved the quality of a passive matrix (LCD) screen. (2) (High Power A ) surveillance system by local National Health Service laboratories. Case-patients were identified as residents in the study area through available geographic information or by patient names, when geographic information was not available. Reports included basic demographic information such as age, sex, and date of disease onset. Where date of onset was not available, date of report was used as a proxy. Travel overseas was not well recorded in these data. Positive isolates of Campylobacter from patients resident in the study area were sent by the main diagnostic laboratories to the NW HPA Laboratory in Manchester for sequence typing. Case-patients were determined to be residents in the study area by using the methods described above. Campylobacter Speciation speciation Formation of new and distinct species, whereby a single evolutionary line splits into two or more genetically independent ones. One of the fundamental processes of evolution, speciation may occur in many ways. C. jejuni isolates were spread onto Colombia blood agar blood agar n. A nutrient culture medium that is enriched with whole blood and used for the growth of certain strains of bacteria. (Oxoid CM331, Unipath, UK) containing 5% defibrinated horse blood and incubated at 37[degrees]C in anaerobic anaerobic /an·aer·o·bic/ (an?ah-ro´bik) 1. lacking molecular oxygen. 2. growing, living, or occurring in the absence of molecular oxygen; pertaining to an anaerobe. jars (Don Whitley Scientific, Shipley, UK) under microaerobic conditions (5% C[O.sub.2], 5% [O.sub.2], 3% [H.sub.2], 87% [N.sub.2]). DNA DNA: see nucleic acid. DNA or deoxyribonucleic acid One of two types of nucleic acid (the other is RNA); a complex organic compound found in all living cells and many viruses. It is the chemical substance of genes. was extracted (10% dilutions) and tested for C. jejuni or C. coli by using a previously described Taqman assay (29) with primers and probes for the genes ceuE (for C. coli)and mapA (for C. jejuni). Sequence Typing of C. jejuni Isolates MLST was performed as described (23). The amplification reactions were performed in a 50-BL volume containing [approximately equal to] 1 [micro]L C. jejuni chromosomal chromosomal, adj relating to chromosome, or a configuration within the cell's nucleus that contains a linear thread of DNA that conveys genetic data. chromosomal emanating from or pertaining to chromosome. DNA (10 ng/[micro]L), 5 [micro]L of each primer (10 pmol/[micro]L), 10 [micro]L of 1 mmol/L deoxynucleoside triphosphates (Roche, Welwyn Garden City Welwyn Garden City (wĕl`ĭn), city (1991 pop. 40,665), Hertfordshire, E central England. It is a garden city, founded by Ebenezer Howard in 1920. Its industries produce a variety of products, including radio and television sets. , UK), 5 [micro]L 10x PCR PCR polymerase chain reaction. PCR abbr. polymerase chain reaction Polymerase chain reaction (PCR) buffer (Qiagen, Crawley, UK), and 0.25 U Taq DNA polymerase DNA polymerase /DNA po·lym·er·ase/ (pah-lim´er-as) any of various enzymes catalyzing the template-directed incorporation of deoxyribonucleotides into a DNA chain, particularly one using a DNA template. (Qiagen). The thermal cycling conditions were as follows: initial denaturation denaturation, term used to describe the loss of native, higher-order structure of protein molecules in solution. Most globular proteins exhibit complicated three-dimensional folding described as secondary, tertiary, and quarternary structures. at 94[degrees]C for 5 min, 40 cycles of denaturation at 94[degrees]C for 2 min; primer annealing annealing (ənēl`ĭng), process in which glass, metals, and other materials are treated to render them less brittle and more workable. at 50[degrees]C for 1 min; and extension at 72[degrees]C for 1 min with a final elongation elongation, in astronomy, the angular distance between two points in the sky as measured from a third point. The elongation of a planet is usually measured as the angular distance from the sun to the planet as measured from the earth. step at 72[degrees]C for 10 min. Thermal cycling was conducted with an MJ PTC (PTC, Needham, MA, www.ptc.com) Long a world leader in mechanical computer-aided design, manufacturing and engineering software, PTC, through acquisitions and reorganization, has transformed itself into a leading provider of Internet-based B2B solutions for discrete manufacturers. 200 thermal cycler The Thermal cycler (also known as a thermocycler, PCR machine or DNA amplifier) is a laboratory apparatus used for PCR. The device has a thermal block with holes where tubes with the PCR reaction mixtures can be inserted. . Amplicons were detected on a 1.5% ethidium bromide Ethidium bromide (sometimes abbreviated as EtBr) is an intercalating agent commonly used as a nucleic acid stain in molecular biology laboratories for techniques such as agarose gel electrophoresis. agarose agarose more highly purified form of agar with similar uses to agar and widely used in the separation of nucleic acid fragments. gel and purified by using a UniFilter Multiscreen PCR cleanup plate (Whatman, Brentford, UK) according to the manufacturer's instructions. Sequencing reactions were conducted in 10 [micro]L of 1/4 reaction volumes containing 2 [micro]L purified DNA, 0.5 [micro]L primer (10 pmol/[micro]L), 1 [micro]L sequencing buffer (Genetix, New Milton New Milton is a market town in south west Hampshire, England. The town has a small high street, which has various shops to suit all residents, and also holds a very successful market every Wednesday. , UK), 2 [micro]L DTCS DTCS Design Technology Consulting Services (Dallas, Texas) DTCS Senior Chief Dental Technician (Naval Rating) DTCS DSCS Tactical Control Subsystem DTCS Distributed Table Construction Scheme Quick Start Master Mix (Beckman Coulter This article needs sources or references that appear in reliable, third-party publications. Alone, primary sources and sources affiliated with the subject of this article are not sufficient for an accurate encyclopedia article. , Fullerton, CA, USA), and 4.5 [micro]L molecular grade water. Thermal cycling conditions for sequencing reactions and ethanol cleanup of sequenced products were set up according to the manufacturer's instructions (Beckman Coulter). The products were analyzed on a Beckman Coulter CEQ CEQ Council On Environmental Quality CEQ Course Experience Questionnaire (higher education) CEQ Centrale de l'Enseignement du Québec CEQ Cinema Equalizer 8000 automated DNA sequencer A DNA sequencer is an instrument used to automate the DNA sequencing process. DNA sequencers have become more important due to large genomics projects and the need to increase productivity. (Beckman Coulter). All sequence assemblage assemblage: see collage. assemblage Three-dimensional construction made from household materials such as rope and newspapers or from any found materials. and editing were performed with Sequencher 4.0 software (GeneCodes Corporation, Ann Arbor Ann Arbor, city (1990 pop. 109,592), seat of Washtenaw co., S Mich., on the Huron River; inc. 1851. It is a research and educational center, with a large number of government and industrial research and development firms, many in high-technology fields such as , MI, USA). MLST 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. and ST Assignment MLST alleles, sequence types (STs), and clonal complexes were assigned by using the Campylobacter PubMLST database (30). Sequences were submitted for allele designation as appropriate. Statistical Analysis Incidence rates quoted were calculated per 100,000 population by using the relevant 2003 annual population estimate for each local authority area, purchased from the UK Office for National Statistics. Incidence ratios, associated 95% confidence intervals confidence interval, n a statistical device used to determine the range within which an acceptable datum would fall. Confidence intervals are usually expressed in percentages, typically 95% or 99%. (CIs), and p values were calculated by using StatsDirect statistical software (http://www.statsdirect.com/), which used Fisher exact test to analyze the difference between 2 crude rates. A mid-p approach to Fisher exact test was also used to test the significance of observed differences in proportions (by using StatsDirect). The level of statistical significance chosen for all analyses was p<0.05. Results Seasonal Incidence of Reported Disease During the first 12 months of the study period (April 2003-March 2004), 493 cases of laboratory-confirmed Campylobacter sp. were reported through the NW surveillance system (Table 1) from residents of Fylde, Wyre, Salford, and Trafford local authorities (Figure 1). This corresponded to approximate annual incidences of 100/100,000 for the area encompassing Fylde and Wyre (rural area) and 73.3/100,000 for the area encompassing Salford and Trafford (suburban area); this difference was significant (p<0.001). The incidence for the whole region of NW England in this period was 69.8/100,000 (data not shown). A similar seasonal pattern of cases was seen in each of the 2 areas in the study, with a large increase in reported cases between weeks 17 to 20 and weeks 21 to 24 (May/June), an elevated incidence through the summer months (weeks 21-36) that declines between weeks 37 and 48 (September to November) to the baseline level of incidence seen in weeks 17-20 (Figure 2). Incidence appeared in increase during the Christmas holiday period (weeks 49-52); this increase was sustained in the rural area until week 8 (February). With the exception of one 4-week period (weeks 25-28), the monthly incidence of campylobacteriosis reported was higher in the rural area, but this finding was only statistically significant during weeks 29-32 in 2003 and weeks 1-8 in 2004. [FIGURE 2 OMITTED] Sequence Typing of Isolates Of 493 cases reported, 388 (79%) laboratory specimens A laboratory specimen is a sample of a species which is preserved and made available to Zoology students in educational institutions. The purpose is to educate the student about the structure, general appearance, various organs, and details related to the specimen's body. were obtained for typing. A proportion of the other cases may not have been identified at the point of diagnosis as study isolates because incomplete demographic information was supplied with the sample. Of the specimens submitted for typing, 30 were C. coli, and 32 did not yield a culture, which left 326 isolates (66% of reported cases) of C. jejuni typed. From these isolates, 93 distinct MLST sequence types of C. jejuni were identified and assigned to 21 clonal complexes, with 20 remaining unassigned until further identification of types enables the designation of new complexes. The most common clonal complex isolated was ST-21 (102 cases, 28.7% of all typed cases, including C. jejuni and C. coli), and this complex was almost 3 times more common than the next complex ST-45 (35 cases, 9.8% of all typed cases) (Table 1). Almost 10% of typed cases were unassigned to clonal complexes at time of writing. Geographic Distribution of Sequence Types The geographic distribution of clonal complexes across the study area varied. Of the 10 most commonly reported clonal complexes (including ones not yet assigned), several were reported with higher incidence in the rural area, although greater numbers are required to give sufficient power to test the significance of these differences in many groups (Table 2). The greatest variation was seen among cases with clonal complexes ST-45 (incidence ratio [IR] 3.53, 95% CI 1.71-7.51) and ST-206 (IR = 1.88, 95%CI 0.65-5.30), although ST-45 was the only complex with a significantly (p<0.001) higher incidence in the rural area. Although not sequence typed, the incidence of C. coli (IR 2.69, 95%CI 1.23-5.95) was also significantly higher in the rural area (p<0.01). Temporal Distribution of Sequence Types Temporal distribution of different clonal complexes in each of the study areas also varied through the first year of the study. Cases of clonal complex ST-45 were most often reported in the rural area during weeks 25-28, although cases were reported continuously throughout the summer months (weeks 21-40, May to September, Figure 3). The proportion of cases reported in weeks 25 to 28 that were typed ST-45 was significantly higher that reported in the annual rural dataset (proportion difference 0.188, exact mid-p = 0.038) (data not shown). In the suburban area, clonal complex ST-45 was most often reported during weeks 21-28 (May to July), and the temporal distribution appeared reduced. The only period in which the difference in ST-45 incidence between the areas approached significance was weeks 25-28 (IR 3.3, 95% CI 0.9-13.17, p = 0.052) (data not shown). Clonal complex ST-45 was not reported before week 21 in either area. [FIGURE 3 OMITTED] Cases of clonal complex ST-21 were reported more or less throughout the 12-month period in both areas, but the incidence fluctuated far less in the suburban area. Almost 50% of the cases reported in the suburban area in weeks 49 to 52 were clonal complex ST-21 compared with 30% in the annual suburban dataset, but this difference was not significant (proportion difference 0.176, exact mid-p = 0.113) (data not shown). Clonal complex ST-206 was most often reported in the rural area in weeks 25 to 28, and more remarkably in weeks 41 to 44, when the proportion of cases typed as ST-206 was significantly higher than in the annual dataset (proportion difference 0.367, exact mid-p = 0.006) (data not shown). Cases of clonal complex ST-206 were consistently reported from weeks 25 through 44 (June to October) in the suburban area, but the proportion of ST-206 cases in weeks 37 to 40 was significantly higher than in the annual dataset (proportion difference 0.101, exact mid-p = 0.035). Age Distribution of Sequence Types Analysis of age-specific incidence by clonal complex was hampered by low numbers of cases; only data for the 6 most commonly reported complexes are shown (Table 3). Overall, the higher incidence of campylobacteriosis described in the rural area was also evident in each of the age bands analyzed, although the difference in 0- to 14-year-old patients was only marginally significant (IR 1.72, 95% CI 0.91-3.18). Among cases in the most commonly reported clonal complex, ST-21, incidence among younger case-patients (0- to 14-year-olds) was higher in the suburban area, but for all other age groups, the incidence was higher in the rural area. None of these differences were statistically significant. The only significant difference was a higher incidence of ST-45 among those [greater than or equal to] 55 years of age in the rural area compared with those [greater than or equal to] 55 years in the suburban area. Discussion We described some characteristics of human infection with Campylobacter in 2 environmentally distinct areas of NW England and some preliminary results of a study that used MLST to better define the epidemiology of campylobacteriosis within a distinct population. MLST identified possible variations in the epidemiology of campylobacteriosis between populations. However, our sample sizes are small, and the role of chance in the associations described cannot be excluded without further analysis of a larger dataset. The seasonality of incidence in the 2 study areas is broadly similar to that previously described (3,6) with a sharp rise in cases around weeks 21 to 24 (May and June) that is sustained through the summer months. Throughout the first year of the study, incidence was higher in the rural area of Fylde and Wyre than in the suburban area of Salford and Trafford; this difference was most significant in the first 8 weeks of 2004. This difference in incidence was seen for all analyzed age groups, although it was not statistically significant among 0- to 14-year-olds. These observations may indicate increased exposure of the more rural population to sources of Campylobacter and true differences in distribution season between the 2 areas, perhaps indicating distinct transmission routes. However, a variety of other factors likely influenced these observations, including differences in healthcare-seeking behavior between the populations and differences in frequency of travel abroad. Although some adjustments have been made for the underlying population structure of each study area (the use of age-specific 4-weekly incidence), estimates of population do not take into account the seasonal movement of persons, such as students (Salford has a large student population) and age-related travel (31,32). For instance, the winter rise in incidence in the rural setting may be due, in part, to winter vacations Winter vacation has been proposed in modern times (the 20th and 21st centuries) as a more practical alternative to summer vacation in areas that have harsh winters and mild summers. taken by those with the flexibility and finances to travel abroad out of season (generally the older generation, who are also more represented in the rural area of this study). The exclusion of travel-related cases will be key in further exploring some of these trends, and matching more detailed epidemiologic information from case questionnaires will facilitate this as the study progresses. Subsequent years of data analysis will also clarify any true differences in incidence between the populations. Sequence typing isolates collected throughout the first 12 months of the study showed a wide range of identified types, with many represented only by single cases and relatively few identified in significant numbers. Several new types were described in this study and await assignment to clonal complexes. Almost 30% of typed isolates from the study area align with the largest clonal complex so far defined, complex ST-21 (23,30), and previously reported isolates from this clonal complex originate from a wide variety of sources other than human cases, including cattle, chicken, milk, sand, and water (23). Most cases arising from a large waterborne outbreak of campylobacteriosis in Walkerton, Ontario Walkerton is a town and is the county seat of Bruce County, Ontario. It is located on the Saugeen River and is 75 km southwest of Owen Sound. The town was incorporated in 1871 and was named after Joseph Walker, who settled in this area in 1850. , that originated from infected in·fect tr.v. in·fect·ed, in·fect·ing, in·fects 1. To contaminate with a pathogenic microorganism or agent. 2. To communicate a pathogen or disease to. 3. To invade and produce infection in. cattle (33) were later identified as clonal complex ST-21 (34), which suggests that this clonal complex can be associated with environmental and foodborue transmission. Study of a farm ecosystem in NW England demonstrated that most clonal complex ST-21 isolates came from livestock, especially cattle (27). Isolates of clonal complex ST-21 have also been described in cloacal cloacal emanating from or pertaining to cloaca. cloacal kiss the contact which occurs during insemination in birds when the vent of the female is everted exposing the cloacal mucosa against which the phallus of the male is pressed. and excreta excreta /ex·cre·ta/ (eks-kret´ah) excretion (2). ex·cre·ta pl.n. Waste matter, such as sweat or feces, discharged from the body. samples from broiler chickens (35) and from a variety of farmyard isolates, including sheep and wild birds (36). Isolates from the next most represented clonal complex in the study area, complex ST-45, have previously been reported largely from humans and chicken (23,37) and were almost exclusively reported in broiler chicken and turkey chicks in a study sampling various livestock and wild birds in NW England (36). However, complex ST-45 was more frequently identified in wildlife than in livestock in the farm ecosystem study above (27) (although the ecosystem studied did not include poultry). Initial data from this current study suggest that ST-45 complex is more frequently identified in the rural component of the study population than in the metropolitan component, and particularly in those >55 years of age. This finding raises the possibility that this complex may be associated with an environmental transmission pathway, a specific set of behavioral traits, or both. This hypothesis is supported by the identification of this clonal complex in wildlife isolates of Campylobacter (27), which suggests a widespread distribution maintained in the environment. Cases of C. coli in this study have a similar distribution, and in previous studies this species has been the most dominant one isolated from surface waters (20,38). The seasonal clustering of ST-45 complex human isolates (compared with ST-21 complex) in summer months in the study populations may also indicate an environmental source. However, seasonal carriage rates in broiler flocks in other north European settings (10,11) correlate with the seasonality of ST-45 complex in this population, in which infection of beef cattle (13) correlates more closely to the seasonality of ST-21 complex. Given the apparent host preferences (not exclusively) of these 2 complexes, the seasonality of human complexes in this study may only reflect Campylobacter distribution in food animals rather than a common environmental source, as suggested by previous studies (2,7,8). No single sequence type was associated with the late spring seasonal rise in human campylobacteriosis, a finding consistent with that of a previous study in England that used serotyping (26). However, we have shown complex ST-45 to be significantly more prevalent during summer months in rural than in suburban areas. In addition, some evidence exists that a Christmas rise in infection in the suburban population may in part be mediated me·di·ate v. me·di·at·ed, me·di·at·ing, me·di·ates v.tr. 1. To resolve or settle (differences) by working with all the conflicting parties: by cases of ST-21. Eating out at a restaurant is a recognized risk factor for campylobacteriosis (32,39), and the Christmas season involves a variety of public as well as private parties in the United Kingdom. Identifying specific sequence types associated with such seasonal activity may help clarify the role of subpopulations of Campylobacter in human epidemiology. Some evidence also suggests that MLST is able to distinguish temporal clusters of campylobacteriosis such as for ST-206, which is significantly 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" in weeks 41 to 44 in the rural area. As this 3-year study progresses, we will match MLST complexes with common epidemiologic exposures through the use of case questionnaire data. The ease of use of the technique and its repeatability in a variety of laboratories are distinct advantages, and the increased use of MLST will enable valuable interlaboratory comparisons of types from similar population-based studies. We have demonstrated the ability to improve linking of apparently sporadic cases encountered in routine surveillance by assigning isolates to sequence type complexes. We believe that MLST will be a valuable tool in testing the significance of suspected epidemiologic exposures in human campylobacteriosis and thus support improved surveillance and development of effective interventions. Acknowledgments We acknowledge the work of the local National Health Service laboratories in managing and making available the study area isolates from routine diagnostic work. We used the Campylobacter jejuni multilocus sequence typing website (http://pubmlst.org/campylobacter/) developed by Keith Jolley and Man-Suen Chan and sited at the University of Oxford. The Food Standards Agency The Food Standards Agency is a non-ministerial government department of the Government of the United Kingdom. It is responsible for protecting public health in relation to food throughout the United Kingdom and is led by an appointed board that is intended to act in the public , United Kingdom, funded this project. 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. 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Int J Biometeorol. 2005;49:207-14. (6.) Sopwith W, Ashton M, Frost JA, Tocque K, O'Brien S O'Bri·en , Edna Born 1932. Irish writer whose works, including The Lonely Girl (1962) and Johnny I Hardly Knew You (1977), explore the lives of women in modern-day Ireland. Noun 1. , Regan M, et al. Enhanced surveillance of campylobacter infection in the North West of England The West of England is a loose term given to the area surrounding the City and County of Bristol, England. It is increasingly used - e.g. by the West of England Partnership - as a synonym for the former Avon (county) area. 1997-1999. J Infect. 2003;46:35-45. (7.) Wilson IG. Salmonella salmonella Any of the rod-shaped, gram-negative, non-oxygen-requiring bacteria that make up the genus Salmonella. Their main habitat is the intestinal tract of humans and other animals. and Campylobacter contamination of raw retail chickens from different producers: a six year survey. Epidemiol Infect. 2002;129:635-45. (8.) Meldrum RJ, Griffiths JK, Smith RM, Evans MR. The seasonality of human campylobacter infection and Campylobacter isolates from fresh, retail chicken in Wales. Epidemiol Infect. 2005;133:49-52. (9.) Louis VR, Gillespie IA, O'Brien SJ, Russek-Cohen E, Pearson AD, Colwell RR. Temperature-driven Campylobacter seasonality in England and Wales. Appl Environ Microbiol. 2005;71:85-92. (10.) Jacobs-Reitsma WF, Bolder NM, Mulder RW. Cecal cecal /ce·cal/ (se´k'l) 1. ending in a blind passage. 2. pertaining to the cecum. ce·cal adj. Of, relating to, or having the characteristics of the cecum. carriage of Campylobacter and Salmonella in Dutch broiler flocks at slaughter: a one-year study. Poult poult a young turkey. Sci. 1994;73:1260-6. (11.) Kapperud G, Skjerve E, Vik L, Hauge K, Lysaker A, Aalmen I, et al. Epidemiological investigation of risk factors for campylobacter colonization colonization, extension of political and economic control over an area by a state whose nationals have occupied the area and usually possess organizational or technological superiority over the native population. in Norwegian broiler flocks. Epidemiol Infect. 1993;111:245-55. (12.) Stanley KN, Wallace JS, Currie cur·rie n. Variant of curry2. JE, Diggle PJ, Jones K. The seasonal variation of thermophilic ther·mo·phil·ic adj. Requiring high temperatures for normal development, as certain bacteria. campylobacters in beef cattle, dairy cattle and calves calves 1 n. Plural of calf1. calves Noun the plural of calf . J Appl Microbiol. 1998;85:472-80. (13.) Stanley KN, Wallace JS, Currie JE, Diggle PJ, Jones K. Seasonal variation of thermophilic campylobacters in lambs at slaughter. J Appl Microbiol. 1998;84:1111-6. (14.) Aho M, Kurki M, Rautelin H, Kosunen TU. Waterborne outbreak of Campylobacter enteritis campylobacter enteritis Infectious disease A water-borne gastroenteritis caused by C jejuni, a cause of travelers' diarrhea Epidemiology Linked to ingestion of contaminated eggs, poultry, water; 2-4 day incubation period Clinical Abdominal pain, ± after outdoors infantry drill in Utti, Finland. Epidemiol Infect. 1989;103:133-41. (15.) Hanninen ML, Niskanen M, Korhonen L. Water as a reservoir for Campylobacter jejuni infection in cows studied by serotyping and pulsed-field gel electrophoresis (PFGE). Zentralblatt Fuer Veterinaermedizin Reihe B. 1998;45:37-42. (16.) Baffone W, Bruscolini F, Pianetti A, Biffi MR, Brandi G, Salvaggio L, et al. Diffusion of thermophilic Campylobacter in the Pesaro-Urbino area (Italy) from 1985 to 1992. Eur J Epidemiol. 1995;11: 83-6. (17.) Jones K, Betaieb M, Telford DR. Thermophilic campylobacters in surface waters around Lancaster, UK: negative correlation Noun 1. negative correlation - a correlation in which large values of one variable are associated with small values of the other; the correlation coefficient is between 0 and -1 indirect correlation with Campylobacter infections in the community. J Appl Bacteriol. 1990;69:758-64. (18.) Schonberg-Norio D, Takkinen J, Hanninen ML, Katila ML, Kankoranta SS, Mattila L, et al. Swimming and Campylobacter infections. Emerg Infect Dis. 2004;10:1474-7. (19.) Carrique-Mas J, Andersson Y, Hjertqvist M, Svensson A, Torner A, Giesecke J. Risk factors for domestic sporadic campylobacteriosis among young children in Sweden. Scand J Infect Dis. 2005;37:101-10. (20.) Brown PE, Christensen OF, Clough n. 1. A cleft in a hill; a ravine; a narrow valley. 2. A sluice used in returning water to a channel after depositing its sediment on the flooded land. 1. (Com.) An allowance in weighing. See Cloff. HE, Diggle PJ, Hart CA, Hazel S, et al. Frequency and spatial distribution of environmental Campylobacter spp. Appl Environ Microbiol. 2004;70:6501-11. (21.) Olsen SJ, Hansen GR, Bartlett L, Fitzgerald C, Sonder A, Manjrekar R, et al. An outbreak of Campylobacter jejuni infections associated with food handler A software routine that performs a particular task. It often refers to a routine that "handles" an exception of some kind, such as an error, but it can refer to mainstream processes as well. The term is typically used in operating systems and other system software. contamination: the use of pulsed-field gel electrophoresis. J Infect Dis. 2001;183:164-7. (22.) Champion OL, Best EL, Frost JA. Comparison of pulsed-field gel electrophoresis and amplified fragment length polymorphism techniques for investigating outbreaks of enteritis enteritis (ĕn'tərī`tĭs), inflammation of the gastrointestinal tract. Acute enteritis is not usually serious except in infants and older people, in whom the accompanying diarrhea can cause dehydration through the loss of fluids. due to campylobacters. J Clin Microbiol. 2002;40:2263-5. (23.) Dingle KE, Colles FM, Wareing DR, Ure R, Fox AJ, Bolton FE, et al. Multilocus sequence typing system for Campylobacter jejuni. J Clin Microbiol. 2001;39:14-23. (24.) Gerner-Smidt P, Hise K, Kincaid J, Hunter S, Rolando S Rolando Jorge Pires da Fonseca (born 31 August, 1985 in São Vicente, Cape Verde) is a Portuguese football player. He currently plays for Belenenses. C.F. , Hyytia-Trees E, et al. PulseNet USA: a five-year update. Foodborne Pathog Dis. 2006;3:9-19. (25.) Dingle KE, Colles FM, Ure R, Wagenaar JA, Duim B, Bolton FJ, et al. Molecular characterization of Campylobacter jejuni clones: a basis for epidemiologic investigation. Emerg Infect Dis. 2002;8:949-55. (26.) Owen RJ, Slater E, Telford D, Donovan T, Barnham M. Subtypes of Campylobacter jejuni from sporadic cases of diarrhoeal disease at different locations in England are highly diverse. Eur J Epidemiol. 1997;13:837-40. (27.) French N, Barrigas M, Brown P, Ribiero P, Williams N, Leatherbarrow H, et al. Spatial epidemiology Spatial epidemiology is the study of the spatial distribution of disease. and natural population structure of Campylobacter jejuni colonizing a farmland ecosystem. Environ Microbiol. 2005;7:1116-26. (28.) Health Protection Agency Standards Unit. Identification of Campylobacter species, BSOP BSOP Biblical Seminary of the Philippines BSOP Brazilian Series of Poker BSOP Bonsai Society of Portland (Portland, Oregon) BSOP Big Suites of Polls (Gaia online) ID 23. [cited 2005 Nov]. Available from http://www.hpa-standardmethods.org.uk/documents/bsopid/ pdf/bsopid23.pdf (29.) Best EL, Powell EJ, Swift C, Grant KA, Frost JA. Applicability of a rapid duplex (communications) duplex - Used to describe a communications channel that can carry signals in both directions, in contrast to a simplex channel which only ever carries a signal in one direction. real-time PCR assay for speciation of Campylobacter jejuni and Campylobacter coli directly from culture plates. FEMS FEMS Federation of European Microbiological Societies FEMS Federation of European Materials Societies FEMS Fabrication Engineering Management System FEMS Facility Equipment Maintenance System (PMEL/TMDE) Microbiol Lett. 2003;229:237-41. (30.) Jolley K, Chan M-S M-S Master-Slave M-S Mid-Side (stereo recording technique) M-S Miznay-Shardin (mine plate charge) . Campylobacter jejuni and Campylobacter coli MLST Home Page [cited 2005 Nov]. Available from http:// pubmlst.org/campylobacter/ (31.) Campylobacter Sentinel Surveillance Scheme Collaborators. Foreign and domestic travel and the risk of Campylobacter infection: results from a population-based sentinel surveillance scheme. J Travel Med. 2003;10:136-8. (32.) Rodrigues LC, Cowden JM, Wheeler JG, Sethi D, Wall PG, Cumberland P, et al. The study of infectious intestinal disease in England: risk factors for cases of infectious intestinal disease with Campylobacter jejuni infection. Epidemiol Infect. 2001;127:185-93. (33.) Clark CG, Price L, Ahmed R, Woodward DL, Melito PL, Rodgers FG, et al. Characterization of waterborne outbreak-associated Campylobacter jejuni, Walkerton, Ontario. Emerg Infect Dis. 2003;9:1232-41. (34.) Clark CG, Bryden L, Cuff cuff (kuf) a small, bandlike structure encircling a part or object. musculotendinous cuff one formed by intermingled muscle and tendon fibers. WR, Johnson PL, Jamieson F, Ciebin B, et al. Use of the oxford multilocus sequence typing protocol and sequencing of the flagellin flagellin /fla·gel·lin/ (flah-jel´in) a protein of bacterial flagella; it is composed of subunits in several-stranded helical arrangement. short variable region to characterize isolates from a large outbreak of waterborne Campylobacter sp. strains in Walkerton, Ontario, Canada. J Clin Microbiol. 2005;43:2080-91. (35.) Connerton PL, Loc CCM CCM Contemporary Christian Music CCM Critical Care Medicine CCM County College of Morris (New Jersey) CCM Chama Cha Mapinduzi (political party, Tanzania) CCM CORBA Component Model , Swift C, Dillon E, Scott A, Rees CED (Capacitance Electronic Disc) An earlier videodisc technology from RCA that was released in 1981 and abandoned five years later. Like phonograph records, the analog disc contained grooves that a stylus rode over. , et al. Longitudinal study longitudinal study a chronological study in epidemiology which attempts to establish a relationship between an antecedent cause and a subsequent effect. See also cohort study. of Campylobacter jejuni bacteriophages and their hosts from broiler chickens. Appl Environ Microbiol. 2004;70:3877-83. (36.) Colles FM, Jones K, Harding RM, Maiden MC. Genetic diversity of Campylobacter jejuni isolates from farm animals and the farm environment. Appl Environ Microbiol. 2003;69:7409-13. (37.) Manning G; Dowson CG, Bagnall MC, Ahmed IH, West M, Newell DG. Multilocus sequence typing for comparison of veterinary and human isolates of Campylobacter jejuni. Appl Environ Microbiol. 2003;69:6370-9. (38.) Rosef O, Rettedal G, Lageide L. Thermophilic campylobacters in surface water: a potential risk of campylobacteriosis. Int J Environ Health Res. 2001;11:321-7. (39.) Friedman CR, Hoekstra RM, Samuel M, Marcus R, Bender J, Shiferaw B, et al. Risk factors for sporadic Campylobacter infection 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. : a case-control study case-control study, n an investigation employing an epidemiologic approach in which previously existing incidents of a medical condition are used in lieu of gathering new information from a randomized population. in FoodNet sites. Clin Infect Dis. 2004;38(Suppl 3):S285-96. Address for correspondence: Will Sopwith, Health Protection Agency NW, 105 Boundary St, Liverpool, UK L5 9YJ; email: will.sopwith@ hpa.org.uk Use of trade names is for identification only and does not imply endorsement by the Public Health Service or by the U.S. Department of Health and Human Services Noun 1. Department of Health and Human Services - the United States federal department that administers all federal programs dealing with health and welfare; created in 1979 Health and Human Services, HHS . Will Sopwith, * Andrew Birtles, ([dagger]) Margaret Matthews Margaret Matthews (born 5 August, 1935) is an American athlete who mainly competed in the 100 meters. She competed for United States in the 1956 Summer Olympics held in Melbourne, Australia where she won the bronze medal in the 4 x 100 meters with her teammates Mae Faggs, , ([dagger]) Andrew Fox Andrew Fox is an American author from New Orleans. He has written two comic novels, Fat White Vampire Blues and Bride of the Fat White Vampire. Both novels feature Jules Duchon, a morbidly obese vampire who resides in New Orleans and works as a taxi driver. , ([dagger]) Steven Gee, ([double dagger double dagger n. A reference mark (  ) used in printing and writing. Also called diesis.Noun 1. ]) Michael Painter, * Martyn Regan, * Qutub Syed, * and Eric Bolton ([dagger]) * Health Protection Agency (North West), Liverpool, United Kingdom; ([dagger]) Regional Health Protection Agency Laboratory, Manchester, United Kingdom; and ([double dagger]) Cumbria and Lancashire Health Protection Unit, Preston, United Kingdom Dr Sopwith is involved in developing web-based surveillance, running training days and conferences, producing regularly published analytical reports, developing disease mapping, and participating in infectious disease Infectious disease A pathological condition spread among biological species. Infectious diseases, although varied in their effects, are always associated with viruses, bacteria, fungi, protozoa, multicellular parasites and aberrant proteins known as prions. research and development projects with the Health Protection Agency in Liverpool, United Kingdom. Current projects include the molecular epidemiology molecular epidemiology Molecular medicine An evolving field that combines the tools of standard epidemiology–case studies, questionnaires and monitoring of exposure to external factors with the tools of molecular biology–eg, restriction endonucleases, of campylobacteriosis in NW England, the epidemiology of cryptosporidiosis Cryptosporidiosis Definition Cryptosporidiosis refers to infection by the sporeforming protozoan known as Cryptosporidia. Protozoa are a group of parasites that infect the human intestine, and include the better known Giardia. , and the effect of rotavirus infection rotavirus infection Virology RI is usually mild, but may be severe in children ≤ 2 yrs due to intense vomiting Morbidity > 870,000 children < age 5 die of rotavirus infection in developing countries, in contrast to 75 to 150 in the US Epidemiology among children.
Table 1. Distribution of Campylobacter jejuni multilocus sequence
typing clonal complexes by study area, April 2003 to March 2004 *
Clonal
complex Fylde Wyre Salford Trafford Total
ST-21 15 21 22 44 102
ST-45 11 10 7 7 35
JA 6 6 6 16 34
ST-257 5 4 9 10 28
ST-443 1 5 6 9 21
ST-48 3 3 2 11 19
ST-206 6 2 3 7 18
ST-353 3 1 1 8 13
ST-22 2 0 2 3 7
ST-49 0 2 2 3 7
ST-42 1 0 2 3 6
ST-354 1 0 0 4 5
ST-61 2 1 1 1 5
ST-283 0 1 0 3 4
ST-52 1 0 2 1 4
ST-573 1 2 0 1 4
ST-658 1 0 2 1 4
ST-403 0 0 0 3 3
ST-508 1 0 0 2 3
ST-460 0 0 2 0 2
ST-177 1 0 0 0 1
ST-362 0 0 1 0 1
C. coli 5 11 4 10 30
No typing 22 24 39 52 137 ([dagger])
Total 88 93 113 199 493
Clonal % of all
complex typed
ST-21 28.7
ST-45 9.8
JA 9.6
ST-257 7.9
ST-443 5.9
ST-48 5.3
ST-206 5.1
ST-353 3.7
ST-22 2.0
ST-49 2.0
ST-42 1.7
ST-354 1.4
ST-61 1.4
ST-283 1.1
ST-52 1.1
ST-573 1.1
ST-658 1.1
ST-403 0.8
ST-508 0.8
ST-460 0.6
ST-177 0.3
ST-362 0.3
C. coli 8.4
No typing
Total
* Including UA (new sequence types as yet unassigned to a clonal
complex). Data show the number of human isolates per local authority
and the percentage of all typed isolates attributed to each complex.
Numbers of C. coli isolates are also shown and included in the
denominator of "all typed cases." Isolates with "no typing" represent
reports of campylobacteriosis to the surveillance system that do not
have a corresponding typed isolate.
([dagger]) Not included in the "all typed cases" denominator.
Table 2. Comparative distributions of the most common MLST clonal
complexes by study area, April 2003 to March 2004 *
Isolates Incidence/100,000
Clonal complex Rural Suburban Rural Suburban
ST-21 36 66 19.91 15.50
ST-45# 21# 14# 11.61# 3.29#
UA 12 22 6.64 5.17
ST-257 9 19 4.98 4.46
ST-443 6 15 3.32 3.52
ST-48 6 13 3.32 3.05
ST-206 8 10 4.42 2.35
ST-353 4 9 2.21 2.11
ST-22 2 5 1.11 1.17
ST-49 2 5 1.11 1.17
C. coli# 16# 14# 8.85# 3.29#
All cases ([dagger])# 181# 312# 100.08# 73.25#
95% confidence
Clonal complex Incidence ratio intervals p value
ST-21 1.28 0.83-1.96 --
ST-45# 3.53# 1.71-7.51# <0.001#
UA 1.28 0.58-2.71 --
ST-257 1.12 0.44-2.59 --
ST-443 1.01 0.32-2.80 --
ST-48 1.09 0.34-3.07 --
ST-206 1.88 0.65-5.30 --
ST-353 1.05 0.24-3.75 --
ST-22 0.94 0.09-5.75 --
ST-49 0.94 0.09-5.75 --
C. coli# 2.69# 1.23-5.95# <0.01#
All cases ([dagger])# 1.37# 1.13-1.65# <0.001#
Note: Those data with a statistically significant difference between
the study areas are indicated with # (level of significance p<0.05).
* Including UA (new sequence types as yet unassigned to a clonal
complex), Campylobactercoli isolates, and all cases of
campylobacteriosis reported. Annual estimated incidence was calculated
for each clonal complex and incidence ratios were calculated for rural
incidence/suburban incidence, including confidence intervals. Those
data with a statistically significant difference between the study
areas are shown in boldface type (level of significance p<0.05). MLST,
multilocus sequence typing.
([dagger]) Refers to the entire dataset for reference, i.e., those
shown in this table plus all others.
Table 3. Comparative age distributions of the most common MLST clonal
complexes by study area (first year of the study) *
Isolates
Clonal Age
complex group Rural Suburban
ST-21 0-14 2 14
15-34 9 24
35-54 12 18
[greater than or equal to] 55 13 13
ST-45 0-14 3 1
15-34 1 4
35-54 7 6
[greater than or equal to] 55# 10# 3#
ST-257 0-14 2 1
15-34 1 8
35-54 3 8
[greater than or equal to] 55# 4 4
ST-443 0-14 1 0
15-34 3 5
35-54 0 6
[greater than or equal to] 55# 1 6
ST-48 0-14 1 2
15-34 5 7
35-54 0 4
[greater than or equal to] 55# 1 2
ST-206 0-14 0 0
15-34 1 6
35-54 5 5
[greater than or equal to] 55# 2 1
All cases 0-14 19 29
15-34# 48# 87#
35-54# 54# 87#
[greater than or equal to] 55# 57# 66#
Incidence/100,000
Clonal Age
complex group Rural Suburban
ST-21 0-14 6.65 17.68
15-34 24.27 21.02
35-54 24.38 15.14
[greater than or equal to] 55 20.17 11.43
ST-45 0-14 9.97 1.26
15-34 2.70 3.50
35-54 14.22 5.05
[greater than or equal to] 55# 15.51# 2.64#
ST-257 0-14 6.65 1.26
15-34 2.70 7.01
35-54 6.09 6.73
[greater than or equal to] 55# 6.21 3.52
ST-443 0-14 3.32 0.00
15-34 8.09 4.38
35-54 0.00 5.05
[greater than or equal to] 55# 1.55 5.28
ST-48 0-14 3.32 2.53
15-34 13.48 6.13
35-54 0.00 3.36
[greater than or equal to] 55# 1.55 1.76
ST-206 0-14 0.00 0.00
15-34 2.70 5.26
35-54 10.16 4.21
[greater than or equal to] 55# 3.10 0.88
All cases 0-14 63.13 36.62
15-34# 129.45# 76.20#
35-54# 109.71# 73.18#
[greater than or equal to] 55# 88.43# 58.05#
95% confidence
Clonal Age Incidence intervals
complex group ratio
ST-21 0-14 0.38 0.04-1.64
15-34 1.15 0.47-2.57
35-54 1.61 0.71-3.53
[greater than or equal to] 55 1.76 0.75-4.13
ST-45 0-14 7.89 0.63-414.40
15-34 0.77 0.02-7.78
35-54 2.82 0.81-10.15
[greater than or equal to] 55# 5.88# 1.51-33.24#
ST-257 0-14 5.26 0.27-310.48
15-34 0.38 0.01-2.87
35-54 0.91 0.15-3.77
[greater than or equal to] 55# 1.76 0.33-9.47
ST-443 0-14 -- --
15-34 1.85 0.29-9.50
35-54 -- --
[greater than or equal to] 55# 0.29 0.01-2.42
ST-48 0-14 1.32 0.02-25.27
15-34 2.20 0.55-8.05
35-54 -- --
[greater than or equal to] 55# 0.88 0.01-16.94
ST-206 0-14 -- --
15-34 0.51 0.01-4.23
35-54 2.42 0.56-10.49
[greater than or equal to] 55# 3.53 0.18-208.11
All cases 0-14 1.72 0.91-3.18
15-34# 1.70# 1.17-2.44#
35-54# 1.50# 1.05-2.13#
[greater than or equal to] 55# 1.52# 1.05-2.20#
Clonal Age p value
complex group
ST-21 0-14 --
15-34 --
35-54 --
[greater than or equal to] 55 --
ST-45 0-14 --
15-34 --
35-54 --
[greater than or equal to] 55# <0.01#
ST-257 0-14 --
15-34 --
35-54 --
[greater than or equal to] 55# --
ST-443 0-14 --
15-34 --
35-54 --
[greater than or equal to] 55# --
ST-48 0-14 --
15-34 --
35-54 --
[greater than or equal to] 55# --
ST-206 0-14 --
15-34 --
35-54 --
[greater than or equal to] 55# --
All cases 0-14 --
15-34# <0.01#
35-54# 0.02#
[greater than or equal to] 55# 0.02#
Note: Those data with a statistically significant difference between
the study areas are indicated with # (level of significance p<0.05).
* Including all cases of campylobacteriosis reported for which age was
available. Annual estimated incidences in 4 age groups were calculated
for each clonal complex using estimates of the age-specific annual
population for each local authority. Incidence ratios were calculated
for rural incidence/suburban incidence and 95% confidence intervals
calculated. Those data with a statistically significant difference
between the study areas are shown in boldface type (level of
significance p<0.05). MLST, multilocus sequence typing.
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