Homologous recombination between bacterial strains is theoretically capable of preventing the separation of daughter clusters, and producing cohesive clouds of genotypes in sequence space. However, numerous barriers to recombination are known. Barriers may be essential such as adaptive incompatibility, or ecological, which is associated with the opportunities for recombination in the natural habitat. Campylobacter jejuni is a gut colonizer of numerous animal species and a major human enteric pathogen. We demonstrate that the two major generalist lineages of C. jejuni do not show evidence of recombination with each other in nature, despite having a high degree of host niche overlap and recombining extensively with specialist lineages. However, transformation experiments show that the generalist lineages readily recombine with one another in vitro. This suggests ecological rather than essential barriers to recombination, caused by a cryptic niche structure within the hosts.
Campylobacter jejuni and C. coli are the most frequent causes of bacterial gastroenteritis in Finland (www.ktl.fi/ttr). The number of laboratory-confirmed cases has nearly doubled over a 10-year period (2,197 cases in 1995 to 4,002 cases in 2005). Campylobacter species are frequently isolated from a wide variety of sources, including poultry, cattle, pigs, sheep, cats, dogs, wild birds, and water. Most Campylobacter infections are sporadic, and the relative contributions of different sources of infection remain unknown. Risk factors for Campylobacter infection, as identified by case-control studies, include eating or preparing raw or undercooked meat, especially chicken meat, drinking unpasteurized milk or untreated water, contact with domestic animals and pets, and foreign travel (14,18,26). A recent case-control study (19) revealed swimming in natural bodies of water to be a novel risk factor. In addition, eating undercooked meat and drinking from dug wells were independent risk factors for domestically acquired Campylobacter infections in Finland (19).Discriminatory typing methods for use in the study of the molecular epidemiology and population genetics of Campylobacter isolates are crucial to better understand the epidemiology and ecology of the organism. Once such information is obtained, it could be used to develop intervention strategies to limit the numbers of human infections. Dingle et al. (6) developed a multilocus sequence typing (MLST) scheme for C. jejuni, which has been shown to be a valuable tool for studying the diversity and population genetics of Campylobacter isolates.
A total of 139 surface water samples from seven lakes and 15 rivers in southwestern Finland were analyzed during five consecutive seasons from autumn 2000 to autumn 2001 for the presence of various enteropathogens (Campylobacter spp., Giardia spp., Cryptosporidium spp., and noroviruses) and fecal indicators (thermotolerant coliforms, Escherichia coli, Clostridium perfringens, and F-RNA bacteriophages) and for physicochemical parameters (turbidity and temperature); this was the first such systematic study. Altogether, 41.0% (57 of 139) of the samples were positive for at least one of the pathogens; 17.3% were positive for Campylobacter spp. (45.8% of the positive samples contained Campylobacter jejuni, 25.0% contained Campylobacter lari, 4.2% contained Campylobacter coli, and 25.0% contained Campylobacter isolates that were not identified), 13.7% were positive for Giardia spp., 10.1% were positive for Cryptosporidium spp., and 9.4% were positive for noroviruses (23.0% of the positive samples contained genogroup I and 77.0% contained genogroup II). The samples were positive for enteropathogens significantly (P < 0.05) less frequently during the winter season than during the other sampling seasons. No significant differences in the prevalence of enteropathogens were found when rivers and lakes were compared. The presence of thermotolerant coliforms, E. coli, and C. perfringens had significant bivariate nonparametric Spearman's rank order correlation coefficients (P < 0.001) with samples that were positive for one or more of the pathogens analyzed. The absence of these indicators in a logistic regression model was found to have significant predictive value (odds ratios, 1.15 ؋ 10 8 , 7.57, and 2.74, respectively; P < 0.05) for a sample that was negative for the pathogens analyzed. There were no significant correlations between counts or count levels for thermotolerant coliforms or E. coli or the presence of F-RNA phages and pathogens in the samples analyzed.
Waterborne outbreaks associated with contamination of drinking water by Campylobacter jejuni are rather common in the Nordic countries Sweden, Norway, and Finland, where in sparsely populated districts groundwater is commonly used without disinfection. Campylobacters, Escherichia coli, or other coliforms have rarely been detected in potential sources. We studied three waterborne outbreaks in Finland caused by C. jejuni and used sample volumes of 4,000 to 20,000 ml for analysis of campylobacters and sample volumes of 1 to 5,000 ml for analysis of coliforms and E. coli, depending on the sampling site. Multiple samples obtained from possible sources (water distribution systems and environmental water sources) and the use of large sample volumes (several liters) increased the chance of detecting the pathogen C. jejuni in water. Filtration of a large volume (1,000 to 2,000 ml) also increased the rate of detection of coliforms and E. coli. To confirm the association between drinking water contamination and illness, a combination of Penner serotyping and pulsed-field gel electrophoresis (digestion with SmaI and KpnI) was found to be useful. This combination reliably verified similarity or dissimilarity of C. jejuni isolates from patient samples, from drinking water, and from other environmental sources, thus confirming the likely reservoir of an outbreak.
A matched case-control study was conducted to study risk factors for domestically acquired sporadic
Campylobacter
infections in Finland. Swimming in natural sources of water was a novel risk factor. Eating undercooked meat and drinking dug-well water were also independent risk factors for
Campylobacter
infection.
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