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R.L. Whitman

Publications and source records attributed to R.L. Whitman.

At least 19 recordsLinked to original sources

Multi-scale temporal and spatial variation in genotypic composition of Cladophora-borne Escherichia coli populations in Lake Michigan

High concentrations of Escherichia coli in mats of Cladophora in the Great Lakes have raised concern over the continued use of this bacterium as an indicator of microbial water quality. Determining the impacts of these environmentally abundant E. coli, however, necessitates a better understanding of their ecology. In this study, the population structure of 4285 Cladophora-borne E. coli isolates, obtained over multiple three day periods from Lake Michigan Cladophora mats in 2007-2009, was examined by using DNA fingerprint analyses. In contrast to previous studies that have been done using isolates from attached Cladophora obtained over large time scales and distances, the extensive sampling done here on free-floating mats over successive days at multiple sites provided a large dataset that allowed for a detailed examination of changes in population structure over a wide range of spatial and temporal scales. While Cladophora-borne E. coli populations were highly diverse and consisted of many unique isolates, multiple clonal groups were also present and accounted for approximately 33% of all isolates examined. Patterns in population structure were also evident. At the broadest scales, E. coli populations showed some temporal clustering when examined by year, but did not show good spatial distinction among sites. E. coli population structure also showed significant patterns at much finer temporal scales. Populations were distinct on an individual mat basis at a given site, and on individual days within a single mat. Results of these studies indicate that Cladophora-borne E. coli populations consist of a mixture of stable, and possibly naturalized, strains that persist during the life of the mat, and more unique, transient strains that can change over rapid time scales. It is clear that further study of microbial processes at fine spatial and temporal scales is needed, and that caution must be taken when interpolating short term microbial dynamics from results obtained from weekly or monthly samples.

Water Research↗

Coastal loading and transport of Escherichia coli at an embayed beach in Lake Michigan

A Chicago beach in southwest Lake Michigan was revisited to determine the influence of nearshore hydrodynamic effects on the variability of Escherichia coli (E. coli) concentration in both knee-deep and offshore waters. Explanatory variables that could be used for identifying potential bacteria loading mechanisms, such as bed shear stress due to a combined wave-current boundary layer and wave runup on the beach surface, were derived from an existing wave and current database. The derived hydrodynamic variables, along with the actual observed E. coli concentrations in the submerged and foreshore sands, were expected to reveal bacteria loading through nearshore sediment resuspension and swash on the beach surface, respectively. Based on the observation that onshore waves tend to result in a more active hydrodynamic system at this embayed beach, multiple linear regression analysis of onshore-wave cases further indicated the significance of sediment resuspension and the interaction of swash with gull-droppings in explaining the variability of E. coli concentration in the knee-deep water. For cases with longshore currents, numerical simulations using the Princeton Ocean Model revealed current circulation patterns inside the embayment, which can effectively entrain bacteria from the swash zone into the central area of the embayed beach water and eventually release them out of the embayment. The embayed circulation patterns are consistent with the statistical results that identified that 1) the submerged sediment was an additional net source of E. coli to the offshore water and 2) variability of E. coli concentration in the knee-deep water contributed adversely to that in the offshore water for longshore-current cases. The embayed beach setting and the statistical and numerical methods used in the present study have wide applicability for analyzing recreational water quality at similar marine and freshwater sites. ?? 2010 American Chemical Society.

Environmental Science & Technology↗

Budget analysis of Escherichia coli at a southern Lake Michigan Beach

Escherichia coli (EC) concentrations at two beaches impacted by river plume dynamics in southern Lake Michigan were analyzed using three-dimensional hydrodynamic and transport models. The relative importance of various physical and biological processes influencing the fate and transport of EC were examined via budget analysis and a first-order sensitivity analysis of model parameters. The along-shore advective fluxofEC(CFU/m 2 ·s)was found to be higher compared to its crossshore counterpart; however, the sum of diffusive and advective components was of a comparable magnitude in both directions showing the importance of cross-shore exchange in EC transport. Examination of individual terms in the EC mass balance equation showed that vertical turbulent mixing in the water column dominated the overall EC transport for the summer conditions simulated. Dilution due to advection and diffusion accounted for a large portion of the total EC budget in the nearshore, and the net EC loss rate within the water column (CFU/m 3 ·s) was an order of magnitude smaller compared to the horizontal and vertical transport rates. This result has important implications for modeling EC at recreational beaches; however, the assessment of the magnitude of EC loss rate is complicated due to the strong coupling between vertical exchange and depth-dependent EC loss processes such as sunlight inactivation and settling. Sensitivity analysis indicated that solar inactivation has the greatest impact on EC loss rates. Although these results are site-specific, they clearly bring out the relative importance of various processes involved.

Environmental Science & Technology↗

Hand-mouth transfer and potential for exposure to E. coli and F + coliphage in beach sand, Chicago, Illinois

Beach sand contains fecal indicator bacteria, often in densities greatly exceeding the adjacent swimming waters. We examined the transferability of Escherichia coli and F + coliphage (MS2) from beach sand to hands in order to estimate the potential subsequent health risk. Sand with high initial E. coli concentrations was collected from a Chicago beach. Individuals manipulated the sand for 60 seconds, and rinse water was analysed for E. coli and coliphage. E. coli densities transferred were correlated with density in sand rather than surface area of an individual's hand, and the amount of coliphage transferred from seeded sand was different among individuals. In sequential rinsing, percentage reduction was 92% for E. coli and 98% for coliphage. Using dose-response estimates developed for swimming water, it was determined that the number of individuals per thousand that would develop gastrointestinal symptoms would be 11 if all E. coli on the fingertip were ingested or 33 if all E. coli on the hand were ingested. These results suggest that beach sand may be an important medium for microbial exposure; bacteria transfer is related to initial concentration in the sand; and rinsing may be effective in limiting oral exposure to sand-borne microbes of human concern.

Journal of Water and Health↗

Seasonal stability of Cladophora-associated Salmonella in Lake Michigan watersheds

The bacterial pathogens Shigella, Salmonella, Campylobacter, and shiga toxin-producing E. coli (STEC) were recently found to be associated with Cladophora growing in southern Lake Michigan. Preliminary results indicated that the Salmonella strains associated with Cladophora were genetically identical to each other. However, because of the small sample size (n = 37 isolates) and a lack of information on spatial-temporal relationships, the nature of the association between Cladophora and Salmonella remained speculative. In this study, we investigated the population structure and genetic relatedness of a large number of Cladophora-borne Salmonella isolates from Lake Michigan (n = 133), as well as those isolated from stream and lake water (n = 31), aquatic plants (n = 8), and beach sands and sediments (n = 8) from adjacent watersheds. Salmonella isolates were collected during 2005-2007 between May and August from Lake Michigan beachsheds in Wisconsin, Illinois, and Indiana. The genetic relatedness of Salmonella isolates was examined by using the horizontal, fluorophore-enhanced rep-PCR (HFERP) DNA fingerprinting technique. While the Salmonella isolates associated with Cladophora exhibited a high degree of genetic relatedness (???92% similarity), the isolates were not all genetically identical. Spatial and temporal relationships were evident in the populations examined, with tight clustering of the isolates both by year and location. These findings suggest that the relationship between Salmonella and Cladophora is likely casual and is related to input sources (e.g. wastewater, runoff, birds) and the predominant Salmonella genotype surviving in the environment during a given season. Our studies indicate that Cladophora is likely an important reservoir for Salmonella and other enteric bacterial pathogens in Lake Michigan beachsheds, which in turn may influence nearshore water quality. ?? 2008 Elsevier Ltd.

Water Research↗

Pilot- and bench-scale testing of faecal indicator bacteria survival in marine beach sand near point sources

Aim: Factors affecting faecal indicator bacteria (FIB) and pathogen survival/persistence in sand remain largely unstudied. This work elucidates how biological and physical factors affect die-off in beach sand following sewage spills. Methods and Results: Solar disinfection with mechanical mixing was pilot-tested as a disinfection procedure after a large sewage spill in Los Angeles. Effects of solar exposure, mechanical mixing, predation and/or competition, season, and moisture were tested at bench scale. First-order decay constants for Escherichia coli ranged between -0??23 and -1??02 per day, and for enterococci between -0??5 and -1??0 per day. Desiccation was a dominant factor for E. coli but not enterococci inactivation. Effects of season were investigated through a comparison of experimental results from winter, spring, and fall. Conclusions: Moisture was the dominant factor controlling E. coli inactivation kinetics. Initial microbial community and sand temperature were also important factors. Mechanical mixing, common in beach grooming, did not consistently reduce bacterial levels. Significance and Impact of the Study: Inactivation rates are mainly dependent on moisture and high sand temperature. Chlorination was an effective disinfection treatment in sand microcosms inoculated with raw influent. ?? 2009 The Society for Applied Microbiology.

Journal of Applied Microbiology↗

Clostridium botulinum type E occurs and grows in the alga Cladophora glomerata

In recent years, massive avian die-offs from Clostridium botulinum type E infection have occurred in the Sleeping Bear Dunes National Lakeshore (SLBE) area of Lake Michigan. These outbreaks have been coincidental with massive blooms of the green algae Cladophora , mostly Cladophora glomerata . We tested the hypothesis that Clostridium botulinum type E can grow under suitable conditions in these algal mats. In a lab mesocosm study, Cladophora from four outbreak-impacted beaches from SLBE were compared with four unimpacted beaches in the Milwaukee–Racine area for bontE gene of Clostridium botulinum . Frequency of the bontE gene was higher after incubation (25 °C for up to 6 weeks) of Cladophora from impacted vs. the unimpacted area. Since no type E gene was detected initially in Cladophora from any of the eight locations, we infer that the increased occurrence of type E gene arose from spore germination or vegetative Clostridium growth within the existing algal mats of SLBE. Moreover, we found that the congener Clostridium perfringens readily grows in mesocosms containing Cladophora .

Lake Michigan;Sleeping Bear Dunes National Lakesho↗

Geographic relatedness and predictability of Escherichia coli along a peninsular beach complex of Lake Michigan

To determine more accurately the real-time concentration of fecal indicator bacteria (FIB) in beach water, predictive modeling has been applied in several locations around the Great Lakes to individual or small groups of similar beaches. Using 24 beaches in Door County, Wisconsin, we attempted to expand predictive models to multiple beaches of complex geography. We examined the importance of geographic location and independent variables and the consequential limitations for potential beach or beach group models. An analysis of Escherichia coli populations over 4 yr revealed a geographic gradient to the beaches, with mean E. coli concentrations decreasing with increasing distance from the city of Sturgeon Bay. Beaches grouped strongly by water type (lake, bay, Sturgeon Bay) and proximity to one another, followed by presence of a storm or creek outfall or amount of shoreline enclosure. Predictive models developed for beach groups commonly included wave height and cumulative 48-h rainfall but generally explained little E. coli variation (adj. R 2 = 0.19-0.36). Generally low concentrations of E. coli at the beaches influenced the effectiveness of model results presumably because of low signal-to-noise ratios and the rarity of elevated concentrations. Our results highlight the importance of the sensitivity of regressors and the need for careful methods evaluation. Despite the attractiveness of predictive models as an alternative beach monitoring approach, it is likely that FIB fluctuations at some beaches defy simple prediction approaches. Regional, multi-beach, and individual beach predictive models should be explored alongside other techniques for improving monitoring reliability at Great Lakes beaches. Copyright ?? 2009 by the American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America. All rights reserved.

Journal of Environmental Quality↗

Summer E. coli patterns and responses along 23 Chicago beaches

Concentrations of E. coli in recreational beach water are highly variable both locally and temporally, but a broader understanding of these fluctuations may be explained through coastal observations. Currently, beach contamination study approaches tend to be site-specific underthe belief that politically delineated beaches are unique and management of beaches cannot be regionally oriented. E. coli data collected over five years from 23 Chicago beaches clearly identified ambient linked patterns at the regional scale. Temporal fluctuations were similar, with all beaches having simultaneous peaks and troughs of E. coli concentrations. Spatially, E. coli concentrations for beaches more closely situated were more closely correlated, indicating spatial autocorrelation. Julian day, wave height, and barometric pressure explained up to 40% of the variation, a value comparable to individual, less parsimonious site-specific models. Day of sampling could explain the majority of the variation in E. coli concentrations, more so than beach, depth, or time of day. Comparing beaches along a targeted coastline allows a better understanding of inherent background regional fluctuations and, ultimately, better predictions of E. coli concentrations in coastal recreational water.

Environmental Science & Technology↗

Environmental occurrence of the enterococcal surface protein ( esp ) gene is an unreliable indicator of human fecal contamination

The enterococcal surface protein ( esp ) gene found in Enterococcus faecalis and E. faecium has recently been explored as a marker of sewage pollution in recreational waters but its occurrence and distribution in environmental enterococci has not been well-documented. If the esp gene is found in environmental samples, there are potential implications for microbial source tracking applications. In the current study, a total of 452 samples (lake water, 100; stream water, 129; nearshore sand, 96; and backshore sand, 71; Cladophora sp. (Chlorophyta), 41; and periphyton (mostly Bacillariophyceae ), 15) collected from the coastal watersheds of southern Lake Michigan were selectively cultured for enterococci and then analyzed for the esp gene by PCR, targeting E. faecalis/ E. faecium ( esp fs/fm ) and E. faecium ( esp fm ). Overall relative frequencies for esp fs/fm and esp fm were 27.4 and 5.1%. Respective percent frequency for the esp fs/fm and esp fm was 36 and 14% in lake water; 38.8 and 2.3% in stream water; 24 and 6.3% in nearshore sand; 0% in backshore sand; 24.4 and 0% in Cladophora sp.; and 33.3 and 0% in periphyton. The overall occurrence of both esp fs/fm and esp fm was significantly related ( χ 2 = 49, P < 0.0001). Post-rain incidence of esp fs/fm increased in lake and stream water and nearshore sand. Further, E. coli and enterococci cell densities were significant predictors for esp fs/fm occurrence in post-rain lake water, but esp fm was not. F + coliphage densities were not significant predictors for esp fm or esp fs/fm gene incidence. In summary, the differential occurrence of the esp gene in the environment suggests that it is not limited to human fecal sources and thus may weaken its use as a reliable tool in discriminating contaminant sources (i.e., human vs nonhuman).

Environmental Science & Technology↗

Coastal strategies to predict Escherichia coli concentrations for beaches along a 35 km stretch of southern Lake Michigan

To understand the fate and movement of Escherichia coli in beach water, numerous modeling studies have been undertaken including mechanistic predictions of currents and plumes and empirical modeling based on hydrometeorological variables. Most approaches are limited in scope by nearshore currents or physical obstacles and data limitations; few examine the issue from a larger spatial scale. Given the similarities between variables typically included in these models, we attempted to take a broader view of E. coli fluctuations by simultaneously examining twelve beaches along 35 km of Indiana's Lake Michigan coastline that includes five point-source outfalls. The beaches had similar E. coli fluctuations, and a best-fit empirical model included two variables: wave height and an interactive term comprised of wind direction and creek turbidity. Individual beach R2 was 0.32-0.50. Data training-set results were comparable to validation results (R2 = 0.48). Amount of variation explained by the model was similar to previous reports for individual beaches. By extending the modeling approach to include more coastline distance, broader-scale spatial and temporal changes in bacteria concentrations and the influencing factors can be characterized. ?? 2008 American Chemical Society.

Environmental Science & Technology↗

Sunlight, season, snowmelt, storm, and source affect E. coli populations in an artificially ponded stream

Reducing fecal indicator bacteria, such as Escherichia coli (E. coli), in streams is important for many downstream areas. E. coli concentrations within streams may be reduced by intervening ponds or wetlands through a number of physical and biological means. A section of Dunes Creek, a small coastal stream of southern Lake Michigan, was impounded and studied for 30??months from pre-through post-construction of the experimental pond. E. coli reduction became more predictable and effective with pond age. E. coli followed the hydrograph and increased several-fold during both rainfall and snowmelt events. Seasonally, the pond was more effective at reducing E. coli during summer than winter. Late summer, non-solar reduction or inactivation of E. coli in the pond was estimated at 72% and solar inactivation at 26%. E. coli DNA fingerprinting demonstrated that the winter population was genetically more homogeneous than the summer population. Detection of FRNA coliphages suggests that there was fecal contamination during heavy rain events. An understanding of how environmental factors interact with E. coli populations is important for assessing anticipated contaminant loading and the reduction of indicator bacteria in downstream reaches. ?? 2007.

Science of the Total Environment↗

Incidence of the enterococcal surface protein ( esp ) gene in human and animal fecal sources

The occurrence of the enterococcal surface protein ( esp ) gene in the opportunistic pathogens Enterococcus faecalis and E. faecium is well-documented in clinical research. Recently, the esp gene has been proposed as a marker of human pollution in environmental waters; however, information on its relative incidence in various human and animal fecal sources is limited. We have determined the occurrence of the esp gene in enterococci from human ( n = 64) and animal ( n = 233) fecal samples by polymerase chain reaction using two primer sets: one presumably specific for E. faecium ( esp fm ) and the other for both E. faecalis and E. faecium ( esp fs/fm ). We believe that this research is the first to explore the use of esp fs/fm for the detection of human waste in natural environmental settings. The incidence in human sources was 93.1% esp fm and 100% esp fs/fm in raw sewage influent; 30% for both esp fm and esp fs/fm in septic waste; and 0% esp fm and 80% esp fs/fm in active pit toilets. The overall occurrence of the gene in animal feces was 7.7% ( esp fs/fm ) and 4.7% ( esp fm ); animal types with positive results included dogs (9/43, all esp fm ), gulls (10/34, esp fs/fm ; 2/34, esp fm ), mice (3/22, all esp fs/fm ), and songbirds (5/55, all esp fs/fm ). The esp gene was not detected in cat (0/34), deer (0/4), goose (0/18), or raccoon (0/23) feces. The inconsistent occurrence, especially in septic and pit toilet sewage, suggests a low statistical power of discrimination between animal and human sources, which means a large number of replicates should be collected. Both esp fm and esp fs/fm were common in raw sewage, but neither one efficiently differentiated between animal and other human sources.

Environmental Science & Technology↗

TiO 2 as a photocatalyst for control of the aquatic invasive alga, Cladophora, under natural and artificial light

Cladophora, a nuisance and invasive, filamentous algae (Chlorophyta), massively accumulates along the shores of the lower Great Lakes each summer causing great economic damage and compromising recreational opportunity and perhaps public health. In vitro experiments showed that Cladophora samples were physically and biologically degraded when subjected to TiO 2 -mediated photocatalysis. For the most successful photocatalytic process, TiO 2 was immobilized on a glass surface and used in combination with either sunlight or artificial UV light. The loss of vital algal pigments was monitored using UV&ndash;vis spectrophotometry, and cell structural changes were determined by microscopic observation. Cladophora, in the presence of TiO 2 -covered glass beads, experienced a loss of chloroplast pigments after 2 h of UV lamp light irradiation. In a separate experiment, sunlight exposure over 4 days (&sim;24 h) resulted in the complete oxidative degradation of the green chloroplast pigments, verified by the UV spectra of the algal extracts. These results suggest that TiO 2 , mobilized on sunlit silicates may be useful in controlling growth and survival of this alga in the Great Lakes, thus mitigating many of the economic, aesthetic ecological impacts of this invasive alga.

Journal of Photochemistry and Photobiology A: Chem↗

Population structure of Cladophora -borne Escherichia coli in nearshore water of Lake Michigan

We previously reported that the macrophytic green alga Cladophora harbors high densities (up to 10 6 colony-forming units/g dry weight) of the fecal indicator bacteria, Escherichia coli and enterococci, in shoreline waters of Lake Michigan. However, the population structure and genetic relatedness of Cladophora -borne indicator bacteria remain poorly understood. In this study, 835 E. coli isolates were collected from Cladophora tufts (mats) growing on rocks from a breakwater located within the Indiana Dunes National Lakeshore in northwest Indiana. The horizontal fluorophore enhanced rep-PCR (HFERP) DNA fingerprinting technique was used to determine the genetic relatedness of the isolates to each other and to those in a library of E. coli DNA fingerprints. While the E. coli isolates from Cladophora showed a high degree of genetic relatedness (⩾92% similarity), in most cases, however, the isolates were genetically distinct. The Shannon diversity index for the population was very high (5.39). Both spatial and temporal influences contributed to the genetic diversity. There was a strong association of isolate genotypes by location (79% and 80% for lake- and ditch-side samplings, respectively), and isolates collected from 2002 were distinctly different from those obtained in 2003. Cladophora -borne E. coli isolates represented a unique group, which was distinct from other E. coli isolates in the DNA fingerprint library tested. Taken together, these results indicate that E. coli strains associated with Cladophora may be a recurring source of indicator bacteria to the nearshore beach.

Water Research↗

Interaction and influence of two creeks on Escherichia coli concentrations of nearby beaches: Exploration of predictability and mechanisms

The impact of river outfalls on beach water quality depends on numerous interacting factors. The delivery of contaminants by multiple creeks greatly complicates understanding of the source contributions, especially when pollution might originate up- or down-coast of beaches. We studied two beaches along Lake Michigan that are located between two creek outfalls to determine the hydrometeorologic factors influencing near-shore microbiologic water quality and the relative impact of the creeks. The creeks continuously delivered water with high concentrations of Escherichia coli to Lake Michigan, and the direction of transport of these bacteria was affected by current direction. Current direction reversals were associated with elevated E. coli concentrations at Central Avenue beach. Rainfall, barometric pressure, wave height, wave period, and creek specific conductance were significantly related to E. coli concentration at the beaches and were the parameters used in predictive models that best described E. coli variation at the two beaches. Multiple inputs to numerous beaches complicates the analysis and understanding of the relative relationship of sources but affords opportunities for showing how these complex creek inputs might interact to yield collective or individual effects on beach water quality.

Journal of Environmental Quality↗

Modeling the transport and inactivation of E. coli and enterococci in the near-shore region of Lake Michigan

To investigate the transport and fate of fecal pollution at Great Lakes beaches and the health risks associated with swimming, the near-shore waters of Lake Michigan and two tributaries discharging into it were examined for bacterial indicators of human fecal pollution. The enterococcus human fecal pollution marker, which targets a putative virulence factor the enterococcal surface protein (esp) in Enterococcus faecium , was detected in 2/28 samples (7%) in the tributaries draining into Lake Michigan and in 6/30 samples (20%) in Lake Michigan beaches. This was indicative of human fecal pollution being transported in the tributaries and occurrence at Lake Michigan beaches. To understand the relative importance of different processes influencing pollution transport and inactivation, a finite-element model of surf-zone hydrodynamics (coupled with models for temperature, E. coli and enterococci) was used. Enterococci appear to survive longer than E. coli , which was described using an overall first-order inactivation coefficient in the range 0.5&minus;2.0 per day. Our analysis suggests that the majority of fecal indicator bacteria variation can be explained based on loadings from the tributaries. Sunlight is a major contributor to inactivation in the surf-zone and the formulation based on sunlight, temperature and sedimentation is preferred over the first-order inactivation formulation.

Environmental Science & Technology↗

Seasonal persistence and population characteristics of Escherichia coli and entercocci in deep backshore sand of two freshwater beaches

We studied the shoreward and seasonal distribution of E. coli and enterococci in sand (at the water table) at two southern Lake Michigan beaches—Dunbar and West Beach (in Indiana). Deep, backshore sand (∼20 m inland) was regularly sampled for 15 months during 2002–2003. E. coli counts were not significantly different in samples taken at 5-m intervals from 0–40 m inland (P=0.25). Neither E. coli nor enterococci mean counts showed any correlation or differences between the two beaches studied. In laboratory experiments, E. coli readily grew in sand supplemented with lake plankton, suggesting that in situ E. coli growth may occur when temperature and natural organic sources are adequate. Of the 114 sand enterococci isolates tested, positive species identification was obtained for only 52 (46%), with E. faecium representing the most dominant species (92%). Genetic characterization by ribotyping revealed no distinct genotypic pattern (s) for E. coli , suggesting that the sand population was rather a mixture of numerous strains (genotypes). These findings indicate that E. coli and enterococci can occur and persist for extended periods in backshore sand at the groundwater table. Although this study was limited to two beaches of southern Lake Michigan, similar findings can be expected at other temperate freshwater beaches. The long-term persistence of these bacteria, perhaps independent of pollution events, complicates their use as indicator organisms. Further, backshore sand at the water table may act as a reservoir for these bacteria and potentially for human pathogens.

Michigan, Indiana↗