Impact of suspended mussels (Mytilus edulis L.) on plankton communities in a Magdalen Islands lagoon (Québec, Canada): A mesocosm approach

The Grande-Entrée Lagoon (Magdalen Islands, Canada) has supported mussel (Mytilus edulis) cultivation for the last 25 years. Algal biomass in this lagoon is relatively low while heterotrophic plankton biomass is high. Although often considered herbivorous, it is known that filter-feeding bivalves can consume various types of food, from bacteria to zooplankton. We hypothesize that along with phytoplankton, heterotrophs constitute an important food resource for the Grande-Entrée mussels. In situ mesocosm experiments were undertaken at different seasons using short socks filled with mussels from the same cohort taken from an aquaculture farm, in order to determine the impact of cultured mussels on local plankton communities and assess the role of heterotrophs. Filtration activity by the mussels and associated epibionts present in the socks was expressed as clearance rates (CR). The average CR over all taxa was lowest in June and highest in October. Diatoms, dinoflagellates and heterotrophic protists constituted the bulk of planktonic carbon removed by mussels. While smaller-sized taxa contributed little (< 5%) to mussel carbon intake, large-sized heterotrophs (namely ciliates) contributed 69 to 88%. Taxon-marker pigment analyses generally confirmed these observations for groups containing phototrophic pigments. The high heterotrophic biomass retained by mussels indicates they are a major food source for mussels in this environment and should be considered both in the evaluation of mussel feeding and in assessing the influence of cultured mussels on local plankton ecosystems.     

Animal effects on dissolved organic carbon bioavailability in an algal controlled ecosystem

1. Animals exert both direct and indirect controls over elemental cycles, linking primary producer-based (green) and decomposer-based (brown) food webs through top-down trophic interactions and bottom-up element regeneration. Where animals are aggregated at high biomass, they create hotspots of elemental cycling. The relative importance of animal control on elemental cycling depends on animal biomass, species functional traits (i.e. feeding mode and stoichiometry), and their overlap.
2. We evaluated how animal community complexity affects the mechanisms regulating energy flow to the brown food web. We conducted a mesocosm experiment where we varied the biomass and overlap of animals with different life history and stoichiometric traits (stream fish and mussels) and measured how this influenced the quantity and fraction of labile carbon available to microbes. We used linear models and structural equation modelling to evaluate direct (excretion) and indirect (herbivory, nutrient availability, and nutrient stoichiometry) effects of animals on bioavailable dissolved organic carbon (BDOC) concentration.
3. In experimental stream mesocosms, we found support for both direct (DOC excretion) and indirect (grazing) animal influences on BDOC concentration. Although we found that snail, fish, and mussel biomass increased nutrient concentrations, neither nutrient concentration nor stoichiometry had a significant effect on BDOC concentration. This has been due to the high background nutrient concentration context of our stream mesocosm water. Snails, probably due to their high biomass and small body size, exerted a significant positive direct control on BDOC concentration. Fish and mussels exerted a significant negative indirect control on BDOC via their effects (grazing and bioturbation) on algal biomass.
4. Our results imply that primary consumers with different feeding strategies provide a key mechanism regulating the flow of DOC into the brown food web through direct (excretion) and indirect (grazing) controls on primary producers. This highlights that animals can provide important controls on the production of bioavailable organic energy supporting microbes in aquatic ecosystems, but the importance of these controls depends on the nutrient context and the distribution of primary producer and animal biomasses.

     

Tracing Consumer-Derived Nitrogen in Riverine Food Webs

The flux of consumer-derived nutrients is recognized as an important ecosystem process, yet few studies have quantified the impact of these fluxes on freshwater ecosystems. The high abundance of bivalves in both marine and freshwater suggests that bivalves can exert large effects on aquatic food webs. The objective of our study was to determine the importance of unionid mussel-derived nitrogen (MDN) to the food web. We used a stable isotope tracer approach in conjunction with nutrient uptake and excretion experiments. We fed mussels (Lampsilis siliquiodea, n = 249) a ¹⁵N-enriched algal diet and placed them into a N-limited stream for 63 days. Mussel hemolymph was non-lethally sampled over the course of the experiment to measure tissue turnover of δ¹⁵N and excretion experiments were done to model the amount of N mussels provided in comparison to stream N uptake demand. Multiple food web pools were sampled twice prior and five times following the mussel addition to trace the ¹⁵N through the food web. Our mussel excretion rates in comparison to areal uptake demand suggested that mussel excretion can account for 40% of the total N demand in this stream. Our enrichment showed that MDN was entering the food web and supplied up to 19% of the N in specific compartments of the food web near the mussel bed. When scaled to a natural mussel aggregation, our results suggest up to 74% of N in the food web may be mussel-derived. Our results show that N supplied by mussels can be an important nutrient subsidy that provides food web support.     

Effects of zebra mussels on cladoceran communities under eutrophic conditions

The purpose of this study was to determine how zebra mussels affected cladoceran community structure under eutrophic conditions. We conducted a mesocosm study where we manipulated the presence of zebra mussels and the presence of large-bodied Daphnia (Daphnia magna and Daphnia pulicaria). We also conducted a complimentary life-table experiment to determine how water from the zebra mussel treatment affected the life history characteristics of the cladoceran species. We anticipated that small- and large-bodied cladoceran species would respond differently to changes in algal quality and quantity under the effects of zebra mussels. Large-bodied Daphnia successfully established in the zebra mussel treatment but failed to grow in the control. We did not observe positive relationships between food concentrations and cladoceran abundances. However, the phosphorus content in the seston indicated that food quality was below the threshold level for large-bodied cladocerans at the beginning of the experiment. We believe that zebra mussels quickly enhanced the phosphorus content in the seston due to the excretion of inorganic phosphorus, thus facilitating the development of large-bodied Daphnia. In conclusion, our results suggest that zebra mussels can alter the phosphorus content of seston in lakes and this can affect the dynamics of crustacean zooplankton.     

Effects of invasive zebra mussels on phytoplankton,turbidity, and dissolved nutrients in reservoirs

Few experiments have quantified the effects of invasive zebra mussels (Dreissena polymorpha) on man-made reservoirs relative to other aquatic habitats. Reservoirs, however, are the dominate water body type in many of the states that are at the current front of the zebra mussel invasion into the western United States. The objective of this research, therefore, was to determine how zebra mussels affected phytoplankton, turbidity, and dissolved nutrients in water that was collected from three Kansas reservoirs that varied in trophic state (mesotrophic to hypereutrophic), but all experienced frequent cyanobacterial blooms. Laboratory mesocosm experiments were conducted to document the effects of zebra mussels on cyanobacteria and general water quality characteristics in the reservoir water. Zebra mussels significantly reduced algal biomass, and the total biovolume of cyanobacteria (communities were dominated by Anabaena) in each reservoir experiment. The effects of zebra mussels on other major algal groups (diatoms, flagellates, and green algae) and algal diversity were less consistent and varied between the three reservoir experiments. Similarly, the effects of zebra mussels on nutrient concentrations varied between experiments. Zebra mussels increased dissolved phosphorus concentrations in two of the reservoir experiments, but there was no effect of zebra mussels on dissolved phosphorus in the mesotrophic reservoir experiment. Combined, our results strongly suggest that zebra mussels have the potential to significantly impact reservoirs as they continue to expand throughout the western United States. Moreover, the magnitude of these effects may be context dependent and vary depending on the trophic state and/or resident phytoplankton communities of individual reservoirs as has similarly been reported for natural lakes.     

The functional role of native freshwater mussels in the fluvial benthic environment

1. Freshwater mussels are the dominant consumer biomass in many fluvial systems. As filter feeding grazers, mussels can remove large amounts of particulate matter from the water column and transfer these resources to the substrate as biodeposits (agglutinated mussel faeces and pseudofaeces). Mussel biodeposits are a nutrient rich and easily assimilated food source and therefore may have significant relevance to benthic community structure. This study examines the functional role of Margaritifera falcata in the South Fork Eel River, California. 2. We addressed two main questions: (i) Do mussels increase benthic resources in this system? (ii) If so, does this alter macroinvertebrate community structure? 3. Measurements and enclosure experiments in the South Fork Eel River show that mussels can play a significant role in local food webs by increasing available fine particulate matter (both organic and inorganic) on the substrate. We document increased benthic macroinvertebrate biomass for predators and collectors (Leptophlebidae) in the presence of mussels, but only in late summer.     

Body size and food thresholds for zero growth in Dreissena polymorpha: a mechanism underlying intraspecific competition

1. Dreissena polymorpha is an extraordinarily successful invasive species that shows high recruitment of small juvenile mussels on established mussel banks. Such juvenile settlement on, and overgrowth of, large adult mussels; however, leads to competition with adults, and often at high densities and low‐food concentrations. 2. The concept of food thresholds for zero growth has been a powerful approach to explaining size‐related exploitative competition in different zooplankton species. We applied it to investigate whether food threshold concentrations for zero growth (C₀) differ between juvenile and adult zebra mussels. 3. By determining body mass growth at various concentrations of a diet mixture (Nannochloropsis limnetica and Isochrysis aff. galbana) we demonstrate that the threshold food concentration for growth of juvenile mussels (C₀ = 0.08 mg C L⁻¹) is substantially lower than that for adults (C₀ = 0.36 mg C L⁻¹). 4. This indicates that, at low food availability, juvenile zebra mussels are competitively superior to their larger conspecifics. Within zebra mussel banks plankton food is substantially depleted and so the observed mechanism might ensure juvenile success and therefore the regeneration of mussel banks in nature.     

Water clarity and eelgrass responses to nitrogen reductions in the eutrophic Skive Fjord, Denmark

Eelgrass depth limits and water clarity in the Skive Fjord estuarine system have not improved despite nutrient input reductions of 30%. Long-term monitoring data (1989–2010) were used to investigate the underlying causes. Dissolved inorganic and organic nitrogen concentrations decreased significantly over time, whereas particulate organic nitrogen concentration, assumed to consist primarily of phytoplankton and phytoplankton detritus and calculated as a proportional factor to chlorophyll a, did not change. Total organic carbon, mostly of autochthonous origin, remained constant despite reduced nitrogen concentrations, resulting in an increasing C:N ratio of the organic material in the water column. Phytoplankton primary production also remained constant suggesting that phytoplankton growth was only limited by nitrogen to a minor degree. Alleviated grazing pressure caused by a reduction in the blue mussel standing stock and a pelagic food web dominated by jellyfish may have contributed to the constantly high phytoplankton levels. Particulate inorganic matter, likely reflecting sediment resuspension, increased over time, most probably in response to removal of blue mussels and declining eelgrass cover. The Skive Fjord estuarine system is affected by multiple pressures—nutrient enrichment, mussel dredging and climate change that must be addressed together for water clarity to improve and eelgrass to recover.     

Understanding how nutrient cycles and freshwater mussels (Unionoida) affect one another

Nutrient loads and nutrient cycling, especially of phosphorus and nitrogen, are among the most important controls on the character of freshwater ecosystems and have been greatly affected by human actions. Despite the widespread importance of nutrients in freshwater ecosystems, the varied linkages between nutrient cycling and freshwater mussel populations have not been thoroughly described. Here, I explore three of these linkages. First, I suggest that nutrient loads are related to the well-being of mussel populations through several mechanisms, probably producing a nonlinear and non-monotonic relationship between nutrient loads and mussel populations. Second, I discuss the ability of mussels to spatially focus nutrients from the overlying water onto the sediments, which has not been fully appreciated, perhaps because nutrient cycling has been viewed chiefly from the viewpoint of the well-mixed water column rather than the patchy sediments. Third, I discuss the ability of mussel populations to accumulate and release nutrients, introducing time lags into nutrient dynamics and stoichiometry (“nutrient capacitance”). Finally, I propose a speculative analysis of the role of freshwater mussels in the nutrient cycles of pristine river systems, which must have been much greater than in modern rivers, with their high nutrient loads and depleted mussel populations.     

Group living and the effects of spatial position in aggregations of Mytilus edulis

Summary The mussel Mytilus edulis typically occurs in aggregations and several consequences of living in groups were studied. Isolated individuals and individuals associated in relatively small groups (6–9 mussles/group) grew more and therefore had greater reproductive output than mussels associated with relatively large groups of 21–28 individuals. Mussels located in the centers of groups exhibited reduced growth and thus lower reproduction relative to mussels located on the edges of groups whose growth and reproduction was similar to that of isolated individuals. Sampling from natural populations indicated that most mussels grow within the matrix of very large groups and hence will experience reduced growth and reproduction. Patterns of growth exhibited by mussels in association with living and model mussels showed that the adverse effects on growth exhibited by mussels in relatively large groups are not a function of the mere physical relief of a mussel clump, but are caused by some property of living neighbors.Laboratory experiments on mussel predation by the crab Pachygrapsus crassipes indicated that crabs prey disproportionately on mussels growing on the edges of groups.The consequences of group living in mobile and nonmobile organisms are considered, and it is suggested that a greater number of negative effects will arise in groups as mobility decreases. In addition, the noted ecological similarity between groups of sessile organisms and spreading clones and its evolutionary implications are discussed.This is contribution no 171 from the Smithsonian Marine Station at Link Port     

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.     

Effects of a small dam on freshwater mussel growth in an Alabama (U.S.A.) stream

1. Dams, ubiquitous features in many lotic ecosystems, are believed to have many broad‐ranging and predominantly negative effects on stream biota. Whereas the impacts of larger dams are well studied, few studies have quantified effects of small dams on streams. 2. Recent surveys found numerous locations where mussels were abundant and larger in reaches immediately downstream from small dams. We examined mussel shell growth and resource conditions in Sandy Creek, a small (third‐order) tributary of the Tallapoosa River in east‐central Alabama (U.S.A.), to determine whether larger populations and individuals result from more rapid growth or longer lifespans of mussels downstream from the dam. 3. Growth rates for populations occurring immediately downstream from the dam (mill reach), c. 5 km downstream from the dam (downstream reach) and upstream from the impoundment (upstream reach) were compared with environmental conditions (seasonal measures of nutrient concentrations and water chemistry) and food availability [total suspended solids (TSS)]. Water temperature was continuously monitored using data loggers. 4. Analysis of length‐at‐age data using multiple growth models found that mill reach mussels grew faster than both up‐ and downstream populations. This dam appears to substantially increase water temperatures and may extend the shell growth period in the mill reach. TSS quantity varied seasonally between sites but was generally highest in the impoundment and mill reach during spring and autumn. TSS quality was highest in the upstream reach from spring through autumn but was highest in the impoundment and mill reach during winter. 5. Our data suggest that some small impoundments enhance conditions for freshwater mussel growth in downstream reaches. However, we do not know how far downstream this subsidy extends or how different species respond to mill dam augmentation. Regardless, mounting evidence suggests that this phenomenon is geographically and taxonomically widespread in eastern North America. Heretofore, undocumented positive effects of small dams suggest that some older dams may warrant protection or restoration if downstream reaches support imperilled mussel populations. Further, some small dams may prove useful conservation tools for natural resource managers attempting to identify sites for mussel culture facilities or translocation refugia.     

Physical forces constrain the depth distribution of the abundant native mussel Elliptio complanata in lakes

1. Unionid mussels often account for a large portion of benthic biomass and contribute to nutrient cycling and sediment processes, but are thought to be limited to shallow areas (<2–3 m). 2. The depth distribution and body size of Elliptio complanata were compared in seven Canadian Shield lake basins of different sizes to test what factors determine the upper and lower limit of their depth range. Specifically, I tested whether (i) the upper range of their distribution is limited by exposure to winds and wave action and (ii) the lower range of their distribution is limited by the depth of the thermocline or by the boundary of mud deposition. 3. The average depth distribution of E. complanata shifted to greater depths in larger lake basins. When comparing individual transects, maximum mussel density was found deeper at more exposed sites. Mussel size decreased with increasing depth and was larger, on average, in larger lake basins. These results suggest that physical forces limit the upper range of mussel distribution in lakes. 4. The maximum depth at which mussels were found in different lakes was closely related to thermocline depth. However, mussels were commonly observed below the predicted depth of the mud deposition boundary. The thermocline limits the lower range of mussel distribution in lakes, probably by limiting food availability and by determining water temperature. Substratum type does not limit the lower distribution of mussels. 5. These results suggest that unionid mussels are present in the deeper parts of the littoral zone, especially in large lakes. Therefore, comparisons of mussel populations between sites and between lakes would be biased unless the full depth distribution of these mussels is considered. These results also suggest that long‐term changes in the thermal structure of lakes could affect the range of unionid mussel populations and their functional role in littoral ecosystems.     

Stoichiometry of regenerated nutrients differs between native and invasive freshwater mussels with implications for algal growth

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Seston quality controls zebra mussel (Dreissena polymorpha ) energetics in turbid rivers

Feeding processes and energetic balance of zebra mussels were both related to the quantity and quality of natural seston. Filtration rate and pseudofeces production increased while clearance rate remained constant with increasing seston concentration. Ingestion rate, assimilation efficiency, and assimilation rate all increased with increasing food quality, measured as the ratio of organic to inorganic material in the seston. Respiration rate did not change with either food quantity or quality. As a result, scope for growth declined with decreasing food quality, and fell below 0 cal mg⁻¹ h⁻¹ at an organic:inorganic ratio of 0.5. The association between feeding processes and food quality appears related to a breakdown in the ability of zebra mussels to selectively ingest high-quality organic particles when the organic content of the seston is low. Ingestion, assimilation efficiency, assimilation rate and scope for growth were all higher when seston was amended with an addition of a natural assemblage of algae. Food quality may be a better indicator of environmental conditions suitable for growth than food quantity. These results suggest that the conditions of high suspended inorganic sediment concentrations in large turbid rivers represent a difficult growth environment for the zebra mussel. Received: 12 May 1997 / Accepted: 7 July 1998     

Effect of inorganic and organic nutrient addition on coral–algae assemblages from the Northern Red Sea

Previous studies in fringing reefs of the Northern Red Sea demonstrated that the in-situ competition of corals and algae in natural assemblages is highly variable between seasons displaying fast overgrowth of corals by benthic reef algae in fall that follows close to equilibrium between both groups of organisms in summer. This may be caused by up to 5-fold higher inorganic nutrient and 6-fold higher organic nutrient concentrations in fall and winter, thereby potentially promoting algae and cyanobacteria growth with concomitant phase shift. A long term mesocosm experiment (duration: 90 days) was conducted in order to study the effect of dissolved inorganic (ammonium, phosphate, nitrate, and mix of all three) and organic (glucose) nutrient addition onto the competitive process in the dominant coral–algae assemblages of the Northern Red Sea involving branching corals of the genus Acropora and a typical consortium of benthic turf algae. Nutrients were added in 3-fold higher concentrations compared to the annual averages, and the parameters algal growth, extension of bleached area on corals, tissue colour change and chlorophyll a concentrations were monitored at regular intervals over experimental duration. This revealed that elevated ammonium concentrations and elevated organic nutrient concentrations stimulate algal growth, while coral tissue pigmentation and chlorophyll a content were significantly decreased. But only in the elevated organic nutrient treatment all effects on corals were significantly pronounced when assembled with benthic turf algae. Supplementary logger measurements revealed that O₂ water concentrations were significantly lower in the elevated organic nutrient mesocosm compared to all other treatments, confirming side-effects on microbial activity. These findings indicate that organic nutrient input into coral reefs can affect physiology and metabolism of both corals and benthic turf algae. Reinforcing interaction between both groups of organisms along with involvement of microbes may facilitate phase shifts in coral reef ecosystems.     

A review of the feedbacks between bivalve grazing and ecosystem processes

This paper gives an overview of interactions betweenbivalve grazing and ecosystem processes, that mayaffect the carrying capacity of ecosystems for bivalvesuspension feeders. These interactions consist of anumber of positive and negative feedbacks.Bivalve grazing can result in local food depletion,which may negatively influence bivalve growth. On alarger scale, it may induce a top-down control ofphytoplankton biomasss, and structural shifts inphytoplankton composition. In the case of harmfulalgal blooms, phytoplankton may negatively affectbivalve grazing rates.The processing of large amounts of particulate mattermay change nutrient cycling on the scale of estuaries,and can result in changes in the inorganic nutrientpool available for phytoplankton, through regenerationand reduced storage of nutrients in algal biomass.This can reduce nutrient limitation of thephytoplankton and stimulate algal growth rates.Observations from mesocosm studies suggest that apositive feedback from bivalve grazing onphytoplankton growth may also change the physiologicalstate of the algae and improve food quality. This revised version was published online in August 2006 with corrections to the Cover Date.     

Evidence of cannibalism and bentho-pelagic coupling within the life cycle of the mussel, Perna canaliculus

The feeding ecology of the green-lipped mussel, Perna canaliculus, was investigated within three intertidal mussel beds along Ninety Mile Beach, northern New Zealand, between August 2000 and March 2001. Adult mussels of different sizes (45-105 mm in shell length) were collected from the intertidal sites about 30 min after being submerged by the incoming tide for gut content analyses. Results of these analyses indicate that mussels consume a variety of phytoplankton, micro- and mesozooplankton, including mussel larvae and post-larvae. Cannibalism of juveniles of up to 620 μm was recorded for intertidal mussels, and conspecifics of up to 2.4 mm were found within the stomachs of additional mussels collected in August 2000 from a nearby subtidal site. For all three intertidal populations, mussel larvae and juveniles contribute about 70% of the food particle consumption during the spawning peak in August, while phytoplankton and other zooplankton constitute the majority of the food source (about 99%) in March, during gametogenesis. Larger intertidal mussels tended to have more food particles in their stomachs than smaller mussels within all three populations. Distinctive differences in food consumption among intertidal populations directly coincide with variations in total particulate matter (TPM), particulate organic matter (POM) and percent organic matter (OM) in the adjacent seawater.Separate experiments designed to test the feeding behavior of mussels feeding at different times during the incoming tide were conducted at one of the intertidal sites during August 2000 and March 2001. Results from these experiments indicate a marked shift in food consumption from bivalves to other mesozooplankton in August, and from phytoplankton to mesozooplankton in March. The observed combination of mussel predatory and grazing behavior over the incoming tide and through the year provides evidence for a strong food-web link between the benthic and pelagic life stages of this species. Furthermore, the high rate of cannibalism during some months of the year suggests that this source of food may significantly contribute to the energy budget of wild populations, with potential implications for evolutionary adaptive success.     

The role of environmental factors in the induction of oxidative stress in zebra mussel (<Emphasis Type="Italic">Dreissena polymorpha</Emphasis>)

The aim of this study was to investigate the influence of specific environmental factors, such as temperature, pH, oxygen concentration, and phosphate, nitrate, chloride, sodium, potassium, sulphate, magnesium and calcium ions concentration, as well as microcystins, on the seasonal variations in the activity of the antioxidant system of the zebra mussel. We examined changes in lipid peroxidation (LPO) levels, glutathione content and the catalase activity of mussels inhabiting the two ecosystems, which differ due to their trophic structure and the presence of toxic cyanobacteria. The results show a relationship between the activity of the antioxidant system of zebra mussels and the seasonal fluctuations of environmental parameters: the symptoms of oxidative stress were generally the highest during spring and the lowest during summer in both ecosystems. Our study also revealed that regardless of the study area the most important factors determining the activity of the antioxidant defences of mussels were the mineral composition (particularly magnesium and calcium ions concentrations) and physical parameters of the water (oxygen concentration and pH). However, factors resulting from the trophic status of studied ecosystems, such as limitations in food resources or high concentration of microcystins during cyanobacterial blooms, were periodically responsible for increased level of LPO in the tissues of zebra mussel. These findings may indicate a limited tolerance of the zebra mussel to the local environmental conditions.     

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.