Richard W. Merritt

Ecology and transmission of Buruli ulcer disease: a systematic review.

Buruli ulcer is a neglected emerging disease that has recently been reported in some countries as the second most frequent mycobacterial disease in humans after tuberculosis. Cases have been reported from at least 32 countries in Africa (mainly west), Australia, Southeast Asia, China, Central and South America, and the Western Pacific. Large lesions often result in scarring, contractual deformities, amputations, and disabilities, and in Africa, most cases of the disease occur in children between the ages of 4–15 years. This environmental mycobacterium, Mycobacterium ulcerans, is found in communities associated with rivers, swamps, wetlands, and human-linked changes in the aquatic environment, particularly those created as a result of environmental disturbance such as deforestation, dam construction, and agriculture. Buruli ulcer disease is often referred to as the “mysterious disease” because the mode of transmission remains unclear, although several hypotheses have been proposed. The above review reveals that various routes of transmission may occur, varying amongst epidemiological setting and geographic region, and that there may be some role for living agents as reservoirs and as vectors of M. ulcerans, in particular aquatic insects, adult mosquitoes or other biting arthropods. We discuss traditional and non-traditional methods for indicting the roles of living agents as biologically significant reservoirs and/or vectors of pathogens, and suggest an intellectual framework for establishing criteria for transmission. The application of these criteria to the transmission of M. ulcerans presents a significant challenge.

Nutrient dynamics, bacterial populations, and mosquito productivity in tree hole ecosystems and microcosms

Water-filled treeholes provide an experimental setting for examining pro- cesses within an ecosystem, and influences of external factors on those processes. Using a limnological, experimental approach involving both natural tree holes and laboratory mi- crocosms of the tree hole ecosystem, we identified and studied interacting, biotic processes, including dynamics of bacterial populations and variation in concentration of inorganic nutrients in tree hole water, and density-dependent competition for food among larvae of the mosquito Aedes triseriatus. We characterized the influence of external factors (inputs of leaf detritus and stemflow) on those processes. Analyses of water samples over time showed that tree hole water was rich and dynamic in nutrients (nitrite, nitrate, ammonium, phosphate, and sulfate); ammonium was the dominant form ofinorganic nitrogen. Variation in nutrient concentrations in microcosms depended upon exogenous inputs (leaf detritus and stemflow water), dilution of nutrients by stemflow, nutrient cycling processes (nitri- fication, denitrification, and sulfate reduction), and ammonium excretion by mosquito larvae. The densities of bacteria in tree hole water, obtained using direct counts of DAPI- fluorochrome stained samples and epifluorescence microscopy, ranged from 2.0 x 106 to 6.0 x 107 cells/mL, and in microcosms from 4.6 x 105 to 2.6 x 108 cells/mL. Experi- mentation involving microcosms revealed that bacterial abundance was reduced by mos- quito feeding and stemflow flushing. Further experiments showed that stemflow flushing increased mosquito productivity from microcosms several-fold and released mosquitoes from density-dependent competition. This effect was likely related to nutrient input and the simultaneous removal of toxic metabolites owing to inputs of stemflow water. We conclude that disturbance by a physical factor, stemflow, has a major influence on the interactions of nutrient dynamics, bacterial populations, and mosquito productivity in temperate tree-hole ecosystems.

The use of invertebrate functional groups to characterize ecosystem attributes in selected streams and rivers in south Brazil

An analysis conducted at nine stream/river sites in the Atlantic Forest region in the State of Paraná, Brazil used macroinvertebrate functional feeding group (FFG) assessments to evaluate ecological condition of the sites. The FFG approach categorizes qualitative macroinvertebrate collections according to their morphological-behavioral adaptations for food acquisition (e.g. scrapers that harvest non-filamentous, attached algae from stable surfaces in flowing water). FFG ratios were employed as surrogates for stream/river ecosystem attributes: balance between autotrophy and heterotrophy; linkage between riparian inputs of coarse particulate organic matter and in-stream food webs; relative dominance of fine particulate organic matter in transport (suspended load) compared to that deposited in the sediments; and geomorphic stability of the channel. The analyses indicated that all nine sites were heterotrophic, six of the nine carried expected levels of suspended organic load and showed below expected linkage with riparian inputs, and in only two were stable substrates limiting. The implications of the findings and recommendations for further analysis and modifications of the protocol are discussed.

Buruli Ulcer (M. ulcerans Infection): New Insights, New Hope for Disease Control

Buruli ulcer is a disease of skin and soft tissue caused by Mycobacterium ulcerans. It can leave affected people scarred and disabled. What are the prospects for disease control?

Distribution of Mycobacterium ulcerans in buruli ulcer endemic and non-endemic aquatic sites in Ghana.

Mycobacterium ulcerans, the causative agent of Buruli ulcer, is an emerging environmental bacterium in Australia and West Africa. The primary risk factor associated with Buruli ulcer is proximity to slow moving water. Environmental constraints for disease are shown by the absence of infection in arid regions of infected countries. A particularly mysterious aspect of Buruli ulcer is the fact that endemic and non-endemic villages may be only a few kilometers apart within the same watershed. Recent studies suggest that aquatic invertebrate species may serve as reservoirs for M. ulcerans, although transmission pathways remain unknown. Systematic studies of the distribution of M. ulcerans in the environment using standard ecological methods have not been reported. Here we present results from the first study based on random sampling of endemic and non-endemic sites. In this study PCR-based methods, along with biofilm collections, have been used to map the presence of M. ulcerans within 26 aquatic sites in Ghana. Results suggest that M. ulcerans is present in both endemic and non-endemic sites and that variable number tandem repeat (VNTR) profiling can be used to follow chains of transmission from the environment to humans. Our results suggesting that the distribution of M. ulcerans is far broader than the distribution of human disease is characteristic of environmental pathogens. These findings imply that focal demography, along with patterns of human water contact, may play a major role in transmission of Buruli ulcer.

Aquatic Snails, Passive Hosts of Mycobacterium ulcerans

ABSTRACT Accumulative indirect evidence of the epidemiology of Mycobacterium ulcerans infections causing chronic skin ulcers (i.e., Buruli ulcer disease) suggests that the development of this pathogen and its transmission to humans are related predominantly to aquatic environments. We report that snails could transitorily harbor M. ulcerans without offering favorable conditions for its growth and replication. A novel intermediate link in the transmission chain of M. ulcerans becomes likely with predator aquatic insects in addition to phytophage insects. Water bugs, such as Naucoris cimicoides, a potential vector of M. ulcerans, were shown to be infected specifically by this bacterium after feeding on snails experimentally exposed to M. ulcerans.

Field and laboratory investigations on the effects of road salt (NaCl) on stream macroinvertebrate communities

Field and laboratory experiments were conducted to examine the effects of road salt (NaCI) on stream macroinvertebrates. Field studies investigated leaf litter processing rates and functional feeding group composition at locations upstream and downstream from point source salt inputs in two Michigan, USA streams. Laboratory studies determined the effects of increasing NaCl concentrations on aquatic invertebrate drift, behavior, and survival. Field studies revealed that leaves were processed faster at upstream reference sites than at locations downstream from road salt point source inputs. However, it was sediment loading that resulted in partial or complete burial of leaf packs, that affected invertebrate activity and confounded normal leaf pack colonization. There were no significant differences that could be attributed to road salt between upstream and downstream locations in the diversity and composition of invertebrate functional feeding groups. Laboratory drift and acute exposure studies demonstrated that drift of Gammarus (Amphipoda) may be affected by NaCl at concentrations greater than 5000 mg/l for a 24-h period. This amphipod and two species of limnephilid caddisflies exhibited a dose response to salt treatments with 96-h LC50 values of 7700 and 3526 mg NaCl/l, respectively. Most other invertebrate species and individuals were unaffected by NaCl concentrations up to 10,000 mg/l for 24 and 96 h, respectively.

Development and application of a macroinvertebrate functional-group approach in the bioassessment of remnant river oxbows in southwest Florida

Invertebrate richness, density, mass, and functional-group analyses were used as surrogates for ecosystem attributes to evaluate conditions of 10 remnant oxbows along the channelized lower Caloosahatchee River in southwest Florida. Replicate 30-sec dipnet samples were taken in Hydrocotyle and Nuphar plant beds that accounted for >80% of cover in the oxbows. Dissolved oxygen (DO) was <65% saturation at the bottom in most oxbows, and all beds, except for 1 Hydrocotyle bed, were rated as heterotrophic. Invertebrate analyses indicated that most oxbows had sufficient coarse particulate organic matter to support normal summer shredder populations and an abundant supply of fine particulate organic matter to support large populations of filtering collectors. High ratios of predators to other functional groups were coupled with many rapid-turnover (i.e., short-life-cycle) prey taxa. Availability of invertebrate food for drift- and benthic-feeding fish was evaluated as generally poor. Overall rankings of ecological condition of oxbows, accomplished using all invertebrate categories and DO levels, formed the basis for recommendations about oxbow restoration along the river.

The effect of an industrial effluent on an urban stream benthic community: Water quality vs. habitat quality

We studied the effect of an industrial effluent on the water quality, habitat quality, and benthic macroinvertebrates of an urban stream in southwestern Michigan (USA). The effluent affected water quality by raising in-stream temperatures 13-18 degree C during colder months and carrying high amounts of iron (> 20 x higher than ambient) that covered the streambed. The effluent also affected habitat conditions by increasing total stream discharge by 50-150%, causing a significant change in substrate and flow conditions. We used three methods to collect benthic macroinvertebrates in depositional and erosional habitats and to understand the relative importance of habitat quality and water quality alterations. Macroinvertebrate response variables included taxonomic richness, abundance, and proportional abundance of sensitive taxonomic groups. Results indicated that the effluent had a positive effect on macroinvertebrate communities by increasing the quantity of riffle habitat, but a negative effect on macroinvertebrate communities by reducing water quality. Results illustrated the need for careful consideration of habitat quality and water quality in restoration or remediation programs.

Aquatic invertebrates as unlikely vectors of Buruli ulcer disease.

Biting water bugs were not correlated with pathogen occurrence.