Seasonal temperature compensation in the horse mussel, Modiolus modiolus: metabolic enzymes, oxidative stress and heat shock proteins

Seasonal collections of the subtidal horse mussel, Modiolus modiolus, from a depth of 10 m were made at the Isles of Shoals, New Hampshire to assess changes in overall energetic demand, measured as respiration, the maximal activities of rate-limiting enzymes of intermediate metabolism, level of oxidative stress, and the expression of heat shock proteins (HSP). Weighted respiration rates of mussels from winter collections were significantly lower than summer rates but decreased by less than 20%. Specific activities of several rate-limiting enzymes were measured in mussels from the summer and winter collections at the temperature of collection and the reciprocal seasonal temperature (15 and 5 degrees C). Comparisons of these enzyme activities and the protein concentrations of hexokinase and citrate synthase show that a quantitative strategy is used to acclimatize to winter temperatures by these rate-limiting enzymes of intermediate metabolism. The activities and protein concentrations of the antioxidant enzyme, Cu/Zn superoxide dismutase (SOD) is seasonally indistinguishable while the concentration of HSP 70 was greater in winter than in summer samples. These results show that mussels seasonally compensate for decreases in temperature by increasing the concentration of rate-limiting metabolic enzymes while maintaining the same level of antioxidant protection in summer and winter consistent with high aerobic metabolism in both winter and summer. Lastly, the significantly greater concentrations of HSP70 in winter samples suggests that protein chaperone functions must be maintained while other seasonal adjustments to cold temperatures are occurring.     

In situ enclosure experiments on the influence of cultured mussels (Perna canaliculus) on phytoplankton at times of high and low ambient nitrogen

The influence of the cultured mussel Perna canaliculus (Gmelin 1791) on the abundance of phytoplankton was investigated in Pelorus Sound, New Zealand. Four in situ enclosure experiments were undertaken, two in summer when ambient nitrogen was low, and two in winter when it was high. Each experiment had four manipulation types: added mussels; added nitrogen; both mussels and nitrogen added; and control (no additions). In summer, there was a significant increase of chlorophyll a in response to added nitrogen, indicating that the phytoplankton were nitrogen-limited. At this time, mussels caused an increase (11-17%) in phytoplankton biomass, possibly by converting particulate nitrogen to ammonium, making the nitrogen available for phytoplankton utilisation. The highest ambient chlorophyll a concentrations coincided with high ambient nitrogen in the winter. At this time, mussel grazing caused a significant decrease (5-14%) in phytoplankton concentration, indicating that within-farm depletion of phytoplankton is most likely to occur in winter. On an annual time scale, the mussels had a stabilising influence on phytoplankton biomass, reducing high ambient levels in winter and slightly increasing low levels in summer.     

Seasonal utilization of different seston carbon sources by the ribbed mussel, Geukensia demissa (Dillwyn) in a mid-Atlantic salt marsh

Seston in salt marshes contains a temporally and spatially complex mixture of natural microparticulate organic material, including phytoplankton, vascular plant detritus, bacteria, heterotrophic nanoflagellates and benthic diatoms. Quantitative information is available concerning how suspension-feeding consumers, such as the ribbed mussel, Geukensia demissa (Dillwyn), utilize some of these components to satisfy their carbon demands. Despite this information there is still a limited understanding of how the relative nutritive contribution of these different dietary items may shift during the year associated with variations in both seston composition and the mussel's physiological condition. To investigate if the mussel's ability to use specific constituents of natural seston varies seasonally, we ran a series of pulse-chase 14C feeding experiments under ambient conditions in March, May, August and November 1996. Phytoplankton, cellulosic detritus, bacteria, heterotrophic nanoflagellates and benthic diatoms were radiolabeled and supplemented in small amounts to natural marsh water for feeding to mussels. The fate of 14C in mussel tissues, feces, respiration and excretion was quantified and contrasted among the different diet types and seasons. Microcapsules containing radiolabeled carbohydrate and protein were used as standards to differentiate possible between-experiment variations in seston composition from seasonal changes in the mussel's feeding and digestive physiology. Mussel clearance rates for all diets were highest in summer and autumn and lowest in winter and spring. In contrast, seasonal shifts in digestive physiology were only found for certain diets. The seasonal range of assimilation efficiencies for microcapsule standards (18-29%) and field-collected microheterotrophs (bacteria 76-93% and heterotrophic nanoflagellates 87-94%) did not differ significantly during the year, whereas summer and autumn assimilation efficiencies for cellulosic detritus (22-24%), phytoplankton (71-79%) and benthic diatoms (89-93%) were up to twofold greater than those in winter and spring (13%, 40-59% and 45-81%, respectively). We conclude that the digestive physiology (e.g., digestive enzyme production) of mussels responds to shifts in dietary components during the year.     

Annual variation in condition,respiration and remineralisation ofMytilus edulis L. in the Sound,Denmark

By means of monthly in situ incubations, variations in oxygen uptake, nutrient release and C/N-ratio were monitored during a period of 14 months of a mussel population (Mytilus edulis L.) located on an exposed beach. A condition index calculated as weight/length³ showed that the condition of the mussels was highest in the spring. Specific oxygen uptake and nutrient release had separate maxima, with high oxygen uptake in the spring coinciding with a period of growth, and high nutrient release during summer when the temperature was highest. Oxygen uptake was significantly correlated with both the condition of the mussels and the temperature, while ammonium release was significantly correlated only with the temperature. Except in spring, the oxygen uptake, condition index and O/N-ratio were low, indicating a poor condition of the mussels. The mussels suffered from suboptimal conditions caused by inadequate food supply and failed to accumulate glycogen reserves essential for the development of mature gametes.     

The role of the blue mussel Mytilus edulis in the cycling of nutrients in the Oosterschelde estuary (The Netherlands)

The fluxes of particulate and dissolved material between bivalve beds and the water column in the Oosterschelde estuary have been measured in situ with a Benthic Ecosystem Tunnel. On mussel beds uptake of POC, PON and POP was observed. POC and PON fluxes showed a significant positive correlation, and the average C:N ratio of the fluxes was 9.4. There was a high release of phosphate, nitrate, ammonium and silicate from the mussel bed into the water column. The effluxes of dissolved inorganic nitrogen and phosphate showed a significant correlation, with an average N:P ratio of 16.5. A comparison of the in situ measurements with individual nutrient excretion rates showed that excretion by the mussels contributed 31–85% to the total phosphate flux from the mussel bed. Ammonium excretion by the mussels accounted for 17–94% of the ammonium flux from the mussel bed. The mussels did not excrete silicate or nitrate. Mineralization of biodeposition on the mussel bed was probably the main source of the regenerated nutrients.From the in situ observations net budgets of N, P and Si for the mussel bed were calculated. A comparison between the uptake of particulate organic N and the release of dissolved inorganic N (ammonium + nitrate) showed that little N is retained by the mussel bed, and suggested that denitrification is a minor process in the mussel bed sediment. On average, only 2/3 of the particulate organic P, taken up by the mussel bed, was recycled as phosphate. A net Si uptake was observed during phytoplankton blooms, and a net release dominated during autumn. It is concluded that mussel beds increase the mineralization rate of phytoplankton and affect nutrient ratios in the water column. A comparison of N regeneration by mussels in the central part of the Oosterschelde estuary with model estimates of total N remineralization showed that mussels play a major role in the recycling of nitrogen.     

Seasonal effects of zebra mussels on littoral nitrogen transformation rates in Gull Lake, Michigan, U.S.A.

• 1 Zebra mussels (Dreissena polymorpha) are successful colonisers of lake littoral habitats and they interact strongly with littoral benthos. Previous research suggests that localised areas colonised by zebra mussels may be hotspots of nitrogen (N) cycling.
• 2 The effects of zebra mussels on nitrification and denitrification rates were examined approximately every other month for 1 year in Gull Lake, Michigan, U.S.A. Littoral sediment was collected from an area free of zebra mussels and distributed into shallow trays; rocks colonised with zebra mussels were placed in half of the trays, while uncolonised rocks were placed in the remaining trays. After an incubation period of 6–8 weeks in the lake, sediment and zebra mussels were collected from the trays, replaced with new sediment and zebra mussels, and placed in the lake for the next interval. In the laboratory, sediment nitrification and denitrification rates were measured for each tray.
• 3 Sediment nitrification rates did not increase in the presence of zebra mussels; instead nitrification rates were sensitive to changes in water temperature and increased with increasing exchangeable sediment ammonium. In contrast, denitrification rates increased in sediment trays with zebra mussels in the winter when nitrate (NO₃⁻) availability was high and when Chara did not grow in the trays.
• 4 Sediment denitrification was NO₃⁻‐limited in all seasons, regardless of zebra mussel treatment. However, sediment in the presence of zebra mussels responded less to NO₃⁻ addition, suggesting that NO₃⁻ limitation of denitrification can be reduced by zebra mussel activity. Zebra mussels have a seasonally variable impact on sediment denitrification rates, and this may translate into altered seasonal patterns of N cycling in localised areas of lakes where they are particularly abundant.

     

Consumer Aggregations Drive Nutrient Dynamics and Ecosystem Metabolism in Nutrient-Limited Systems

Differences in animal distributions and metabolic demands can influence energy and nutrient flow in an ecosystem. Through taxa-specific nutrient consumption, storage, and remineralization, animals may influence energy and nutrient pathways in an ecosystem. Here we show these taxa-specific traits can drive biogeochemical cycles of nutrients and alter ecosystem primary production and metabolism, using riverine systems that support heterogeneous freshwater mussel aggregations. Freshwater unionid mussels occur as distinct, spatially heterogeneous, dense aggregations in rivers. They may influence rates of production and respiration because their activities are spatially concentrated within given stream reaches. Previous work indicates that mussels influence nutrient limitation patterns, algal species composition, and producer and primary consumer biomass. Here, we integrate measures of organismal rates, stoichiometry, community-scaled rates, and ecosystem rates, to determine the relative source–sink nutrient dynamics of mussel aggregations and their influence on net ecosystem processes. We studied areas with and without mussel aggregations in three nitrogen-limited rivers in southeastern Oklahoma, USA. We measured respiration and excretion rates of mussels and collected a subset of samples for tissue chemistry and for thin sectioning of the shell to determine growth rates at each site. This allowed us to assess nutrient remineralization and nutrient sequestration by mussels. These rates were scaled to the community. We also measured stream metabolism at three sites with and without mussels. We demonstrated that mussel species have distinct stoichiometric traits, vary in their respiration rates, and that mussel aggregations influence nutrient cycling and productivity. Across all mussel aggregations, we found that mussels excreted more nitrogen than they sequestered into tissue and excreted more phosphorus than they sequestered except at one site. Furthermore, gross primary productivity was significantly greater at reaches with mussels. Collectively, our results indicate that mussels have ecosystem-level impacts on nutrient availability and production in nutrient-limited rivers. Within these streams, mussels are affecting the movement of nutrients and altering nutrient spiralling.     

Species traits and environmental conditions govern the relationship between biodiversity effects across trophic levels

Changing environments can have divergent effects on biodiversity–ecosystem function relationships at alternating trophic levels. Freshwater mussels fertilize stream foodwebs through nutrient excretion, and mussel species-specific excretion rates depend on environmental conditions. We asked how differences in mussel diversity in varying environments influence the dynamics between primary producers and consumers. We conducted field experiments manipulating mussel richness under summer (low flow, high temperature) and fall (moderate flow and temperature) conditions, measured nutrient limitation, algal biomass and grazing chironomid abundance, and analyzed the data with non-transgressive overyielding and tripartite biodiversity partitioning analyses. Algal biomass and chironomid abundance were best explained by trait-independent complementarity among mussel species, but the relationship between biodiversity effects across trophic levels (algae and grazers) depended on seasonal differences in mussel species’ trait expression (nutrient excretion and activity level). Both species identity and overall diversity effects were related to the magnitude of nutrient limitation. Our results demonstrate that biodiversity of a resource-provisioning (nutrients and habitat) group of species influences foodweb dynamics and that understanding species traits and environmental context are important for interpreting biodiversity experiments.     

Consumption rates of two rotifer species by zebra mussels Dreissena polymorpha

We demonstrated that zebra mussels Dreissena polymorpha collected from the Hudson River could consume two rotifer species that were common before the zebra mussel invasion. The clearance rates (volume of water filtered per hour) of zebra mussels differed when feeding on the two rotifer species but both decreased with an increase in rotifer density. The ingestion rates (biomass of rotifers per hour) for the two rotifer species also differed, but both increased with increasing rotifer density. This is the first experiment to measure zooplankton consumption by bivalve molluscs at different zooplankton densities. The feeding rates of mussels in this study were compared with those of phytoplankton and other zooplankton from previous studies. The diameter of the mussel inhalent siphon was correlated linearly with the shell length and tissue weight, and was usually an order of magnitude wider than rotifer sizes. It is concluded that bivalve suspension feeders not only act as food competitors but also as predators on zooplankton in the aquatic ecosystem.     

Ecosystem Processes Performed by Unionid Mussels in Stream Mesocosms: Species Roles and Effects of Abundance

Unionid mussels are a guild of freshwater, sedentary filter-feeders experiencing a global decline in both species richness and abundance. To predict how these losses may impact stream ecosystems we need to quantify the effects of both overall mussel biomass and individual species on ecosystem processes. In this study we begin addressing these fundamental questions by comparing rates of ecosystem processes for two common mussel species, Amblema plicata and Actinonaias ligamentina, across a range of abundance levels and at two trophic states (low and high productivity) in stream mesocosms. At both low and high productivity, community respiration, water column ammonia, nitrate, and phosphorus concentrations, and algal clearance rates were all linearly related to overall mussel biomass. After removing the effects of biomass with ANCOVA, we found few differences between species. In a separate series of experiments, nutrient excretion (phosphorus, ammonia, and molar N:P) and biodeposition rates were only marginally different between species. For the species studied here, functional effects of unionids in streams were similar between species and linearly related to biomass, indicating the potential for strong effects when overall mussel biomass is high and hydrologic residence times are long.     

Broad thermal capacity facilitates the primarily pelagic existence of northern fur seals (Callorhinus ursinus)

Thermoregulatory capacity may constrain the distribution of marine mammals despite having anatomical and physiological adaptations to compensate for the thermal challenges of an aquatic lifestyle. We tested whether subadult female northern fur seals (Callorhinus ursinus) experience increased thermoregulatory costs in water temperatures potentially encountered during their annual migration in the Bering Sea and North Pacific Ocean. Metabolic rates were measured seasonally in 6 captive female northern fur seals (2.75–3.5 yr old) in ambient air and controlled water temperatures of 2°C, 10°C, and 18°C. Rates of oxygen consumption in ambient air (1°C–18°C) were not related to environmental temperature except below 2.5°C (winter only). However, metabolism was significantly higher during the fall seasonal trials (September–October) compared to other times of year, perhaps due to the costs of molting. The fur seals appeared thermally neutral in all seasons for all water temperatures tested (2°C–18°C) except during the summer when metabolic rates were higher in the 2°C water. Comparing this broad thermal neutral zone to the average sea surface temperatures potentially encountered during annual migrations indicates wild fur seals can likely exploit a large geographic area without added thermal metabolic costs.     

Dreissenid mussels enhance nutrient efflux, periphyton quantity and production in the shallow littoral zone of a large lake

Dreissenid mussels are notorious invasive organisms whose establishment is associated with large, ecosystem-scale changes to patterns of productivity in aquatic systems. We investigated how localized impacts of dreissenids affect the potential of littoral substrates to support primary and secondary production using in situ incubations and comparisons of natural mussel-colonized and mussel-free substrates in the littoral zone of a large, shallow lake. We compared dreissenid-colonized and dreissenid-free substrates in terms of nutrient balance, surface area, periphyton loads as well as benthic primary production and respiration rates. Dreissenid-colonized substrates acted as sources of dissolved nutrients to the water column, with mussel mass-specific rates of dissolved phosphorus and ammonia excretion averaging 7.2 ± 5.6 (mean ± SD), and 92.6 ± 64.7 μg/g mussel shell free dry mass/h, respectively. Mussel-colonized substrates also had higher surface area, and supported approximately double the amount of periphyton and organic matter loads compared to mussel-free substrates, as well as having higher rates of primary production and community respiration. We suggest that the localized effects of dreissenids can play an important role in changing whole-ecosystem production patterns, with the extent of dreissenid impacts strongly dependent on lake size and morphometry.     

Effects of season, temperature, and body mass on the standard metabolic rate of tegu lizards (Tupinambis merianae)

Abstract This study examined how the standard metabolic rate of tegu lizards, a species that undergoes large ontogenetic changes in body weight with associated changes in life-history traits, is affected by changes in body mass, body temperature, season, and life-history traits. We measured rates of oxygen consumption (Vo(2)) in 90 individuals ranging in body mass from 10.4 g to 3.75 kg at three experimental temperatures (17 degrees , 25 degrees , and 30 degrees C) over the four seasons. We found that standard metabolic rate scaled to the power of 0.84 of body mass at all experimental temperatures in all seasons and that thermal sensitivity of metabolism was relatively low (Q(10) approximately 2.0-2.5) over the range from 17 degrees to 30 degrees C regardless of body size or season. Metabolic rates did vary seasonally, being higher in spring and summer than in autumn and winter at the same temperatures, and this was true regardless of animal size. Finally, in this study, the changes in life-history traits that occurred ontogenetically were not accompanied by significant changes in metabolic rate.     

Seasonal Differences in Hypoxia Tolerance in Gulf Killifish, Fundulus Grandis (Fundulidae)

Many estuarine habitats are characterized by episodes of hypoxia, the frequency and severity of which may vary seasonally. Accordingly, resident fish species may show seasonal differences in their capacity to tolerate hypoxia. We have tested this hypothesis in the gulf killifish, Fundulus grandis, sampled from the Lake Pontchartrain estuary (Louisiana) at different times of the year. We measured 2 indicators of hypoxia tolerance, the frequency of aquatic surface respiration (ASR) during gradual reduction in dissolved oxygen (D.O.) and survival time during severe hypoxic stress, and found both to be significantly affected by season. Fish collected during the summer did not engage in ASR until the D.O. concentration dropped to values lower than that required to elicit ASR by fish collected during other seasons. Laboratory acclimation of fish to low oxygen did not change the relationship between ASR behavior and D.O., suggesting that the observed seasonal effect on ASR was not simply due to previous exposure of summer fish to environmental hypoxia. Furthermore, fish collected during the summer and winter had significantly longer survival times during exposure to severe hypoxia than fish collected during the fall. Survival analysis indicated that the condition of fish was positively associated with survival time, and seasonal variation in condition accounted for about half of the observed difference between survival times of fish collected during the summer and fall. Seasonal variation in ASR and survival, when taken together, demonstrate that hypoxia tolerance in F. grandis may be subject to acclimatization. An increase in hypoxia tolerance during the summer could increase survivorship of fish when exposed to elevated temperatures and low oxygen concentrations which prevail during the summer months.     

Empirical models for predicting the excretion of nutrients (N and P) by aquatic metazoans: taxonomic differences in rates and element ratios

1. We developed empirical models for predicting the release of nutrients [nitrogen (N) and phosphorus (P)] by aquatic metazoans (zooplankton, mussels, benthic macroinvertebrates and fish). 2. The number of species represented in each model ranged from 9 to 74 (n = 40 – 1122), organism dry mass from 1 × 10^?5 to 8 × 10⁴ mg and water temperature from ?1.8 to 32 °C for all models. Organisms were from marine and freshwater (both lotic and lentic) environments. 3. Rates and ratios of nutrient excretion were modelled and intra‐ and intertaxon differences in excretion were examined. Rates of N and P excretion were not significantly different between marine and freshwater species within the same taxon (e.g. zooplankton). However, rates of excretion (as a function of organism dry mass and water temperature) were significantly different among different orders of zooplankton, mussels and fish. However, excretion of N was similar among different orders of benthic macroinvertebrates. 4. Detritivorous fish excreted both N and P at rates greater than all other taxa; whereas mussels excreted N and P generally at rates less than other taxa. There were no significant differences in the rate of N and P excretion between zooplankton and fish (i.e. the allometry of N and P excretion was similar between zooplankton and fish). 5. Molar N : P ratios of nutrients excreted increased with increasing organism dry mass for each group of metazoans, except for zooplankton and detritivorous fish (where N : P ratios declined with increasing organism dry mass). Molar N : P ratios in the excretions of aquatic metazoans were generally below the Redfield ratio of 16:1. 6. We examined the influence of variable abundance of zooplankton, benthic macroinvertebrates and fish on assemblage excretion rates. Rates of N and P excretion were calculated by applying our models to metazoan biomass and abundance data over seven consecutive years in two oligotrophic lakes. Rates of N and P excretion (g ha^?1 day^?1) increased linearly with increasing assemblage biomass (kg ha^?1). However, rates of N and P excretion were significantly and negatively correlated with the relative abundance of fish and positively correlated with the relative abundance of zooplankton.     

Respiration and Dry Matter Accumulation Around the Time of Anthesis in Field Stands of Winter Wheat (Triticum aestivum)

Rates of CO₂ efflux from the aerial parts of eight winter wheat(Triticum aestivum L. em Thell) genotypes were determined duringspike emergence, anthesis and early grain filling over two seasons.Dry weight data were also recorded at flag leaf full expansionand anthesis Respiration rates on a ground area basis did not differ (P >0–05) among the times of day when measurements were made.In contrast, highly significant day-to-day effects (P > 0–001)were evident. Daily respiratory efflux was consistently highduring spike emergence, declining to lower values before andafter anthesis in each year. Changes in daily respiration wereanalysed in terms of a two-parameter respiration model. Modelpredictions suggested that changes in chemical composition ofthe dry matter formed during ontogeny could explain the day-to-dayeffect Although no genotypic differences in respiration (ground areabasis) and crop growth rate were found, significant negativecorrelations between crop d. wt and respiration rate per unitd. wt were evident. Genotypic differences in crop d. wt alsobecame more distinct from the first harvest to the second. Theseobservations are discussed in terms of genotypic variation inmaintenance respiration Wheat, respiration, genotype, ontogeny     

Context-dependent effects of freshwater mussels on stream benthic communities

1. We asked whether unionid mussels influence the distribution and abundance of co‐occurring benthic algae and invertebrates. In a yearlong field enclosure experiment in a south‐central U.S. river, we examined the effects of living mussels versus sham mussels (shells filled with sand) on periphyton and invertebrates in both the surrounding sediment and on mussel shells. We also examined differences between two common unionid species, Actinonaias ligamentina (Lamarck 1819) and Amblema plicata (Say 1817). 2. Organic matter concentrations and invertebrate densities in the sediment surrounding mussels were significantly higher in treatments with live mussels than treatments with sham mussels or sediment alone. Organic matter was significantly higher in the sediment surrounding Actinonaias than that surrounding Amblema. Actinonaias was more active than Amblema and may have increased benthic organic matter through bioturbation. 3. Living mussels increased the abundance of periphyton on shells and the abundance and richness of invertebrates on shells, whereas effects of sham mussels were similar to sediment alone. Differences in the amount of periphyton growing on the shells of the two mussel species reflected differences in mussel activity and shell morphology. 4. Differences between living and sham mussel treatments indicate that biological activities of mussels provide ecosystem services to the benthic community beyond the physical habitat provided by shells alone. In treatments containing live mussels we found significant correlations between organic matter and chlorophyll a concentrations in the sediment, organic matter concentrations and invertebrate abundance in the sediment and the amount of chlorophyll a on the sediment and invertebrate abundance. There were no significant correlations among these response variables in control treatments. Thus, in addition to providing biogenic structure as habitat, mussels likely facilitate benthic invertebrates by altering the availability of resources (algae and organic matter) through nutrient excretion and biodeposition. 5. Effects of mussels on sediment and shell periphyton concentrations, organic matter concentrations and invertebrate abundance, varied seasonally, and were strongest in late summer during periods of low water volume, low flow, and high water temperature. 6. Our study demonstrates that freshwater mussels can strongly influence the co‐occurring benthic community, but that effects of mussels are context‐dependent and may vary among species.     

Seston quality drives feeding,stoichiometry and excretion of zebra mussels

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Overland transport of recreational boats as a spreading vector of zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis>

In aquatic ecosystems invasive species are among the most important threats to biodiversity worldwide. Understanding the dispersal mechanisms of aquatic invaders is very important for protection and management of vulnerable water bodies. Here we ask how recreational boats that are transported overland could contribute to the dispersal of invasive zebra mussels among lakes in Switzerland. Using a questionnaire sent to registered boat owners, we surveyed properties of transported boats and collected information on self-reported mussel fouling and transport activities of boat owners. We also sampled boat hulls at launching ramps and harbors for biofouling invertebrates. Boats that were kept seasonally or year-round in water were found to have high vector potential with mussel fouling rates of more than 40 %. However, only about 6 % of boats belonging to these groups were transported overland to other water bodies. Considering that approximately 100,000 recreational boats are registered in Switzerland, we estimated that every year around 1400 boats fouled with mussels are transported overland. Such boats pose a high risk of distributing zebra mussels between water bodies. Our results suggest that there is a considerable risk that recreational boats may spread new fouling species to all navigable water bodies within the study area. We speculate that one such species could be the quagga mussel, which has not yet invaded lakes in Switzerland. On a more positive note, our study has identified the group of high-risk boats so that possible control measures would only affect a relatively small number of boat owners.     

Cost of selective feeding by the blue mussel (Mytilus trossulus) as measured by respiration and ammonia excretion rates

The metabolic cost as measured by respiration and ammonia excretion rates associated with a selective as compared to a non-selective feeding behaviour was determined for the blue mussel, Mytilus trossulus. Mussels were challenged with four environmentally relevant seston matrices of different quality and quantity, which were known to evoke either a sorting response (i.e., selective feeding) where organic-rich particles were selected over inorganic particles as compared to no sorting (i.e., non-selective) where either inorganic or organic particles were ingested by the bivalve. Seston matrices were prepared by mixing known quantities of silt and algae such that the following extent of feeding responses would occur; no pre-sorting of ingested material, (1) no algae+50 mg silt l⁻¹, (2) 150×10⁶ cells l⁻¹ of algae+no silt, and, where pre-sorting of ingested material occurred, (3) 20×10⁶ cells l⁻¹ of algae and 20 mg l⁻¹ of silt, and (4) 150×10⁶ cells l⁻¹ of algae and 50 mg l ¹ of silt. A control, which represented basal metabolism consisting of mussels exposed only to filtered seawater (0.45 μm), was included for a total of five treatments. Mussel respiration and ammonia excretion rates were independent of whether mussels were pre-sorting or not sorting ingested material. Of the four matrices, only rates determined for conditions of maximum seston quality and quantity where pre-sorting of the seston occurred were significantly different from control mussels (p<0.05, ANOVA). Estimates of net energy used for feeding, where net energy is total energy intake (food) minus energy expenditure (energy lost through respiration and excreta as measured by ammonia excretion rates), indicated that feeding, whether selective or non-selective required only 0.92% of net energy intake. Hence, mussels appear to be highly adapted to a dynamic food environment with negligible costs associated with the feeding process, even when significant pre-selection of organic-rich particles occurs.