Grid-generated turbulence in a mesocosm experiment

In two mesocosm experiments designed to study the effect of nutrients and turbulence on trophic levels below fish larvae in the ecosystem, turbulence was created by oscillating grids. Utilizing two different frequencies, two levels of turbulence (`high' and `low') were generated in a total of eight mesocosms in both experiments. In each mesocosm a two-layer density structure was created. We succeeded in generating turbulence that mimics turbulence found in natural ecosystems, with turbulent eddies on larger scales than those relevant for organism interactions. Energy dissipation rates () were calculated from the frequency spectrums of the measured turbulent velocities. In the second experiment the average energy dissipation rates in the upper layer were 1.9×10^–7 W kg^–1 and 5.5×10^–8 W kg^–1 in the `high' and `low' enclosures, respectively. Finding `background' turbulence levels between 5.3×10^–9 and 3×10 ^–8 W kg^–1, we suggest that future experiments consider turbulent measurements as a standard variable.     

Development of Baltic Sea zooplankton in the presence of a toxic cyanobacterium: a mesocosm approach

Cyanobacteria blooms are common in the Baltic Sea and are consideredto be a poor food source and sometimes toxic to zooplankton.Most experiments demonstrating harmful effects have been short-termincubations with monocultures or simple mixtures of food. Inthis study, a mesocosm approach was used to examine zooplanktonresponses over generation timescales. A toxic strain of thecyanobacterium Nodularia spumigena was added to bag enclosuresof ambient water. The initial mesozooplankton concentrationwas either reduced by prescreening the water or enriched withlocally caught zooplankton. Experiments ran for 15 days, longenough to monitor reproductive success and development of thenext mesozooplankton generations. There was no major harmfuleffect on the zooplankton assemblage, even though the concentrationof the toxin nodularin was in the upper range of field observations.The copepod Eurytemora affinis, rotifers Synchaeta spp. andthe cladoceran Bosmina longispina maritima were able to developand reproduce successfully in the presence of N. spumigena.The only species showing impaired recruitment was the copepodAcartia bifilosa. The general lack of population level effectsfrom N. spumigena in this study can be reconciled with previousobservations of adverse effects. Cyanobacteria alone may bepoor food and toxic to zooplankton, but in the mesocosms a richassemblage of microbiota developed, similar to that associatedwith blooms in the field. We suggest that, in the context ofotherwise food-depleted summer situations in the open BalticSea, zooplankton can derive benefit from cyanobacteria bloomassemblages.     

Immigration and zooplankton community responses to nutrient enrichment: a mesocosm experiment

Dispersal can be an important determinant of local diversity and species composition, but evidence for effects of the regional species pool on local zooplankton communities has been mixed. Theory and experiments suggest that immigration will be necessary for maintenance of community diversity and functioning during periods of environmental change; conversely, fluctuating resource levels may increase the likelihood of invasion success. We conducted a factorial-design mesocosm experiment to test the effects of a nutrient pulse and weekly immigration from other lakes on the diversity and composition of a pelagic zooplankton community. Contrary to expectations, there were no interactive effects of nutrient enrichment and immigration on any measure of diversity, and the initial shift in community composition in response to the nutrient pulse did not depend on the introduction of new species or genotypes from more productive lakes. Although immigration increased species richness in enclosures, success of most colonising species was poor. However, the dispersal treatment appears to have enabled a stronger predator response to increased herbivore numbers in nutrient-pulsed enclosures, leading to an eventual decline in the abundance of some herbivorous species in response to immigration. We conclude that community invasibility was not influenced by productivity, and that dispersal limitation did not strongly constrain the response of the zooplankton community to our applied disturbance. This indicates an unexpected resistance to change in species composition and diversity in spite of disturbance, and suggests that, in our study system, changes in the abundance of resident species are more important than introductions of new species in the community response to short-term environmental change.Electronic Supplementary Material Supplementary material is available to authorized users in the online version of this article at .     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.     

Impact of turbulence on riverine zooplankton: a mesocosm experiment

1. With increases in river discharge over time and space, zooplankton generally encounter increased turbulence, turbidity, hydraulic forces, downstream advection and food limitations, all of which should affect species diversity and densities. Of these factors, the role of turbulence on the distribution of zooplankton is least known along longitudinal and lateral dimensions in river networks. 2. We tested the factorial effects of turbulence and grazing level on Ohio River potamoplankton in spring and summer using twelve 1600‐L, outdoor mesocosms. Turbulence was calculated using the Froude number for equal depths but with current velocities of 0.064 and 0.32 m s^?1. Grazing levels corresponded to a high density treatment (=ambient river densities of rotifers, copepods and cladocerans) and a low density treatment (initially no zooplankton >64 μm). All tanks had the same water residence time, and hydraulic stress was minimized by circular flow patterns. 3. Zooplankton densities and population growth rates were significantly affected by turbulence level and season. In general, rotifer populations grew faster in high turbulence tanks (though Keratella and Brachionus populations flourished in both treatments in summer) and microcrustaceans thrived better in low turbulence environments. The larger, calanoid copepods handled more turbulent conditions much better than cyclopoids or nauplii. Zooplankton had no detectable effects on particulate organic carbon concentrations in either month (values were higher in spring), but rotifers reduced chlorophyll concentrations in both months. 4. The relative importance of turbulence in controlling potamoplankton is probably to vary not only on a longitudinal basis in river networks but also with both the hydrogeomorphic complexity of river reaches and the type and amount of river regulation. Plans for river rehabilitation and management should incorporate non‐turbulent habitats in large rivers as a means of enhancing zooplankton populations and providing an important food web component for planktivores.     

Horizontal distribution of photo- and heterotrophic microorganisms on the coastal area of the northern Baltic Sea--a case study

The horizontal distribution of bacteria (BACT), bacterial production(FDC), picoplanktonic cyanobacteria (CYANO), and photosynthetic(PNF) as well as heterotrophic (HNF) nanoflagellates was studiedat the end of June on the SW coast of Finland, the Baltic Sea.Samples were taken at 0 and 10 m depths from three transects500 and 800 m apart. The distance between sampling points was20 m. Cell counts of BACT, CYANO, PNF, HNF and the FDC showed3- to 5-fold deviations in the whole data set. Differences betweenthe duplicate counts within the subsamples were 10–20%.Within the transects, the maximum values were 1.2–3.0times higher than the minimum values. Differences between thetransects were statistically significant for all variables exceptFDC. Deviations in cell numbers between the surface and 10 mwere quite irregular. but often statistically significant. Inmost cases, the organism groups did not correlate significantlywith each other. The horizontal distribution of pelagic microbeswas sufficiently uniform, on a meter scale. for one samplingpoint to give a reliable estimate of microbial densities inthe area under investigation. The differences between the surfaceand the lower stratum of the mixed layer (10 m) were in mostcases statistically significant.     

Temperature-induced increase in methane release from peat bogs: a mesocosm experiment

Peat bogs are primarily situated at mid to high latitudes and future climatic change projections indicate that these areas may become increasingly wetter and warmer. Methane emissions from peat bogs are reduced by symbiotic methane oxidizing bacteria (methanotrophs). Higher temperatures and increasing water levels will enhance methane production, but also methane oxidation. To unravel the temperature effect on methane and carbon cycling, a set of mesocosm experiments were executed, where intact peat cores containing actively growing Sphagnum were incubated at 5, 10, 15, 20, and 25°C. After two months of incubation, methane flux measurements indicated that, at increasing temperatures, methanotrophs are not able to fully compensate for the increasing methane production by methanogens. Net methane fluxes showed a strong temperature-dependence, with higher methane fluxes at higher temperatures. After removal of Sphagnum, methane fluxes were higher, increasing with increasing temperature. This indicates that the methanotrophs associated with Sphagnum plants play an important role in limiting the net methane flux from peat. Methanotrophs appear to consume almost all methane transported through diffusion between 5 and 15°C. Still, even though methane consumption increased with increasing temperature, the higher fluxes from the methane producing microbes could not be balanced by methanotrophic activity. The efficiency of the Sphagnum-methanotroph consortium as a filter for methane escape thus decreases with increasing temperature. Whereas 98% of the produced methane is retained at 5°C, this drops to approximately 50% at 25°C. This implies that warming at the mid to high latitudes may be enhanced through increased methane release from peat bogs.     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

Response of epiphytic algae to nutrient loading and fish density in a shallow lake: a mesocosm experiment

Studies on the effect of eutrophication on the ecology of shallow lakes, usually pay scant attention to changes within the epiphytic algal community, though the contribution of this to the ecosystem dynamics is transcendental. In order to test the influence of nutrient loadings and fish densities in the structure of algal epiphyton in a shallow lake, an experiment was performed using in situ mesocosms. Nutrient additions were related to significant decreases in the total epiphyton biovolume and that of bacillariophyceans and zygnematophyceans, but with increases in the abundance of cyanobacteria. The different response of algal groups at the higher nutrient concentrations (increases or decreases in their abundance and/or biovolume) can be related to their ecophysiological constraints such as different resistance to toxicant ammonium accumulation. Plant-associated macroinvertebrates numbers were positively correlated with total numbers of epiphyton. The presence of planktivorous fish enhanced the abundance and biovolume of all algal groups, except cyanobacteria. Fishes enhanced the abundance of plant-associated animals and of total epiphyton. Fish indirect effects (e.g., nutrients release) and their dietary particularities were among the factors that together with nutrients influenced epiphyton growth. The role of indirect effects of fishes and the importance of their dietary particularities are stressed as key factors to understand the processes controlling epiphyton ecology and the food web structure of shallow lakes. Handling editor: D. Ryder.     

Arctic Sea ice biota: design and evaluation of a mesocosm experiment

A mesocosm experiment (enclosure volume 220 l) was designed such that sea ice inhabited by Arctic Sea ice organisms was formed and maintained under natural conditions at 66°N in Rovaniemi, Finland. The experiment was run from natural freezing in December 1994 to melting in April 1995. The ice was inhabited by diatoms, chlorophyceae, heterotrophic flagellates, ciliates, nematodes and turbellarians. Biomass in the ice, expressed as Chlorophyll a concentration, was 20–110 μg l⁻¹; total cell densities varied from 5 × 10⁶ to 35 × 10⁶ cells l⁻¹. Amongst phototrophic organisms, a succession from a flagellate-dominated community (Chlamydomonas sp.) to a multi-species diatom-dominated community was observed. Typical Arctic species such as Nitzschia frigida and Melosira arctica were present in the ice. Bacterial concentration varied between 2 × 10⁸ and 7 × 10⁸ cells l⁻¹. At least two trophic levels were present in the ice. Received: 3 April 1997 / Accepted: 9 September 1997     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.     

Environmental influences on Vibrio populations in northern temperate and boreal coastal waters (Baltic and Skagerrak Seas)

Even if many Vibrio spp. are endemic to coastal waters, their distribution in northern temperate and boreal waters is poorly studied. To identify environmental factors regulating Vibrio populations in a salinity gradient along the Swedish coastline, we combined Vibrio-specific quantitative competitive PCR with denaturant gradient gel electrophoresis-based genotyping. The total Vibrio abundance ranged from 4 x 10(3) to 9.6 x 10(4) cells liter(-1), with the highest abundances in the more saline waters of the Skagerrak Sea. Several Vibrio populations were present throughout the salinity gradient, with abundances of single populations ranging from 5 x 10(2) to 7 x 10(4) cells liter(-1). Clear differences were observed along the salinity gradient, where three populations dominated the more saline waters of the Skagerrak Sea and two populations containing mainly representatives of V. anguillarum and V. aestuarianus genotypes were abundant in the brackish waters of the Baltic Sea. Our results suggest that this apparent niche separation within the genus Vibrio may also be influenced by alternate factors such as nutrient levels and high abundances of dinoflagellates. A V. cholerae/V. mimicus population was detected in more than 50% of the samples, with abundances exceeding 10(3) cells liter(-1), even in the cold (annual average water temperature of around 5 degrees C) and low-salinity (2 to 4 per thousand) samples from the Bothnian Bay (latitude, 65 degrees N). The unsuspected and widespread occurrence of this population in temperate and boreal coastal waters suggests that potential Vibrio pathogens may also be endemic to cold and brackish waters and hence may represent a previously overlooked health hazard.     

Response of planktonic communities to calcium hydroxide addition in a hardwater eutrophic lake: results from a mesocosm experiment

The main focus of this study was to investigate the effects of lime treatment with Ca(OH)₂ on a hypereutrophic hardwater lake (Lake Alserio, Northern Italy), focusing on its impact on plankton communities. We performed a mesocosm experiment using two large enclosures, one for treatment and one control, in the lake. The addition of Ca(OH)₂ proved to be effective in reducing phosphorus concentration, but the effect was limited in time. Redissolution of the precipitating calcium carbonate as it came into contact with the deep water determined a phosphorus increase about 2 weeks after the liming. The liming had a strong impact on the plankton assemblage, as was demonstrated by the sharp decrease in phytoplankton and zooplankton biomass and density, and 1 month after the treatment we observed a different taxonomic composition inside the enclosure from that of the lake. The result of the experiment showed that liming with Ca(OH)₂ can be regarded as a suitable treatment for accelerating recovery of the lake. Multiple treatments will be necessary to improve the trophic level over a longer period.     

Zooplankton response to shading effects of free-floating plants in shallow warm temperate lakes: a field mesocosm experiment

Dense mats of free floating plants (FFP) often produce severe underwater light attenuation and strong oxygen depletion in the water column. In this study, we experimentally assessed the zooplankton response to artificial shading using field mesocosms. During 30 days, we simulated three different light scenarios by mimicking the persistence, absence, and fluctuation of FFP typically encountered in vegetated shallow subtropical lakes. We used dark meshes to simulate the abiotic effects engineered by FFP. Both in the permanently covered and fluctuating situations, anoxia impaired zooplankton development. Anoxia constituted a major driving force in shaping the zooplankton response, whereas the feeding resource availability (phytoplankton) seemed to play a minor role; no top down effect on phytoplankton occurred in anoxic situations. In the fluctuating cover regime (periodic darkness and anoxia), the temporal variation of nanophytoplankton was not affected by zooplankton; once again oxygen availability seemed the main force shaping the zooplankton dynamics. Either periodical or permanent shading, associated to anoxic conditions, impaired the success of small herbivores. Large herbivores and microphytoplankton were negatively affected only under persistent shade and anoxia. In contrast, when neither light nor oxygen limitation occurred, such as in the scenario without shading, top-down control occurred. This study highlights the importance that the oxygen dynamics driven by the presence of FFP exert on the structure and dynamics of zooplankton assemblages and on the top down cascading effects on phytoplankton in warm temperate or subtropical shallow lakes.     

Combined effects of fish and macroinvertebrate predation on zooplankton in a littoral mesocosm experiment

Whether macrophytes offer an effective refuge for zooplankton in all shallow lakes is subject to debate. To explore potential constraints between different predator threats and the related habitat choice by zooplankton, we conducted a mesocosm experiment in 12 large-sized pools mimicking the nearshore environment with part of its length being covered by submersed macrophytes (Egeria densa) and holding a mixed zooplankton community. Four treatments were used: (i) young zooplanktivorous fish (3 silverside, Odontesthes bonariensis) in the “open-water” zone; (ii) macroinvertebrate predator (31 grass shrimp, Palaemonetes argentinus) in the vegetated zone; (iii) both, fish in the open-water and shrimp in the vegetated zones; and (iv) control with no predators. Our results show specific effects of each predator on the abundance, composition, and size of cladocerans. Regarding distribution, in control and shrimp mesocosms, no differences were found between the two zones, while cladocerans were clearly more abundant in the vegetated side in the presence of fish. When both fish and shrimp were present, cladocerans preferred the vegetated zone too, but in a smaller proportion, and their abundance was less. The presence of predatory macroinvertebrates in vegetated littoral zone reduces the refuge value of this habitat, at least for cladocerans.

     

Effects of water-level fluctuations on the littoral benthic fish community in lakes: a mesocosm experiment

Changes in the water level of lakes, either natural or man-made,are important environmental perturbations for eulittoral benthicfish communities. In outdoor mesocosm experiments, we testedthe effects of decreasing shelter availability due to autumnlake-level decrease on the behavior and the growth of two littoralbenthic dwellers, the juvenile burbot and the stone loach. Thetwo species showed significantly different changes in behaviorwhen shelter availability decreased. Burbot built up a distincthierarchy when shelter became sparse, with larger fish beingsignificantly more successful in competing for suitable shelterthan smaller conspecifics. The hierarchy, however, ceased whenshelter availability decreased below a certain level. The largestfish then increasingly abandoned shelter use, while smallerfish persisted with their sheltering behavior. Stone loach,in contrast, showed no hierarchical order or size-related shelteruse. Only burbot showed a significant relationship between theability to occupy the preferred shelter and the somatic growth.These two species provide an example of different strategiesto deal with environmental perturbations like lake-level decrease,with the stenoecious, dominant strategy of the burbot and theeuryoecious, evasive strategy of stone loach. The results demonstratethe importance of conducting behavioral studies under as naturalconditions as possible.     

Reproductive characteristics of the male herring in the northern Baltic Sea

The gonad weight and gonadosomatic index of the male Baltic herring Clupea harengus membras L., were highest at the beginning of the reproductive season (April/May), the values decreasing towards the end of it (July/August) during 1988–1991. The decline could not be explained by fish size but may have been due to fish condition. A high individual variation was typical for both gonad weights and gonadosomatic indices in fish of the same size and maturity stage. The mean density of sperm cells was significantly higher in June (34·9 × 10⁹ ml⁻¹) than in July (19·2 × 10⁹ ml⁻¹, Mann-Whitney U= 17; P<0·05), the variation among the males being high in both groups. Electron microscope analysis showed a severe disruption of the mitochondrial elements in males spawning at 22°C.     

Effects of zebra mussels on phytoplankton and ciliates: a field mesocosm experiment

Many observational studies in North American lakes have documenteddecreases in phytoplankton abundance after the invasion of thezebra mussel (Dreissena polymorpha). However, few field experimentshave examined in detail the effect of zebra mussels on phytoplanktonabundance and species composition over an extended period. Replicatedin situ mesocosms were used to evaluate the impact of naturaldensities of zebra mussels on phytoplankton and ciliate biovolume,and algal species composition over a 5-week period in a habitatthat lacked extant mussel populations. Mussel biomass used inthe experiment was determined using a regression model basedon a data analysis that predicts zebra mussel biomass from totalphosphorus concentration. Within 1 week, zebra mussels decreasedphytoplankton biovolume by 53% and ciliate biovolume by 71%.The effect of zebra mussels on ciliate biovolume was sustainedthroughout the study. However, the effect of zebra mussels onphytoplankton abundance gradually waned over the remaining 4weeks of the experiment, such that the declining effect of zebramussels could not be explained by a shift towards less edibleand/or faster growing algal species. The mussels’ decliningcondition could help to explain the effect observed over thecourse of the experiment.