Copepod grazing in a summer cyanobacteria bloom in the Gulf of Finland

Blooms of the cyanobacteria Nodularia spumigena and Aphanizomenon flos-aquae dominated the phytoplankton assemblages of the western Gulf of Finland and the eastern side of the northern Baltic Sea in late July–August, 1992. The bloom overlapped the peak seasonal contributions of the dominant mesozooplankton herbivores in the region, the copepods Acartia bifilosa and Eurytemora affinis and the cladoceran Bosmina longispina maritima. Using radio-labelling techniques; the copepods were offered one of the cyanobacteria, Nodularia, as well as the 10–54 µm fraction of the natural phytoplankton assemblage. In general, incorporation rates of the labelled phytoplankton into the copepods declined with increasing contributions of the cyanobacteria. For both copepods, incorporation was inversely related to total phytoplankton biomass, whether measured as chlorophyll, total cells or cyanobacteria biomass. The very low rates for Acartia (< 0.8 µl [copepod h]^–1) indicated that this copepod was likely starving in the cyanobacteria bloom, consistent with the generally poor condition of the animal observed in the laboratory. The other major mesozooplanktor, B. longispina maritima, ingested substantially more cyanobacterial biomass than the two copepods, based on HPLC-identified cyanobacteria-specific pigment echinenone in the gut. Bloom carbon provided < 1% and < 4% of the daily rations for Acartia and Eurytemora, respectively. Total copepod demand in the cyanobacteria blooms was trivial, < 1% of bloom biomass consumed daily. These results suggest that copepod herbivory is relatively unimportant in dissipating summer cyanobacteria blooms in the Gulf of Finland.     

Projected marine climate change: effects on copepod oxidative status and reproduction

Zooplankton are an important link between primary producers and fish. Therefore, it is crucial to address their responses when predicting effects of climate change on pelagic ecosystems. For realistic community‐level predictions, several biotic and abiotic climate‐related variables should be examined in combination. We studied the combined effects of ocean acidification and global warming predicted for year 2100 with toxic cyanobacteria on the calanoid copepod, Acartia bifilosa. Acidification together with higher temperature reduced copepod antioxidant capacity. Higher temperature also decreased egg viability, nauplii development, and oxidative status. Exposure to cyanobacteria and its toxin had a negative effect on egg production but, a positive effect on oxidative status and egg viability, giving no net effects on viable egg production. Additionally, nauplii development was enhanced by the presence of cyanobacteria, which partially alleviated the otherwise negative effects of increased temperature and decreased pH on the copepod recruitment. The interactive effects of temperature, acidification, and cyanobacteria on copepods highlight the importance of testing combined effects of climate‐related factors when predicting biological responses.     

Copepod reproductive effort and oxidative status as responses to warming in the marine environment

The marine ecosystems are under severe climate change‐induced stress globally. The Baltic Sea is especially vulnerable to ongoing changes, such as warming. The aim of this study was to measure eco‐physiological responses of a key copepod species to elevated temperature in an experiment, and by collecting field samples in the western Gulf of Finland. The potential trade‐off between reproductive output and oxidative balance in copepods during thermal stress was studied by incubating female Acartia sp. for reproduction rate and oxidative stress measurements in ambient and elevated temperatures. Our field observations show that the glutathione cycle had a clear response in increasing stress and possibly had an important role in preventing oxidative damage: Lipid peroxidation and ratio of reduced and oxidized glutathione were negatively correlated throughout the study. Moreover, glutathione‐s‐transferase activated in late July when the sea water temperature was exceptionally high and Acartia sp. experienced high oxidative stress. The combined effect of a heatwave, increased cyanobacteria, and decreased dinoflagellate abundance may have caused larger variability in reproductive output in the field. An increase of 7°C had a negative effect on egg production rate in the experiment. However, the effect on reproduction was relatively small, implying that Acartia sp. can tolerate warming at least within the temperature range of 9–16°C. However, our data from the experiment suggest a link between reproductive success and oxidative stress during warming, shown as a significant combined effect of temperature and catalase on egg production rate.     

Deposit-feeders accumulate the cyanobacterial toxin nodularin

Blooms of toxic cyanobacteria may potentially affect food web productivity and even be a human health hazard. In the Baltic Sea, regularly occurring summer blooms of nitrogen-fixing cyanobacteria are often dominated by Nodularia spumigena, which produces the potent hepatotoxin nodularin. Evidence of sedimentation of these blooms indicates that benthic fauna can be exposed to nodularin. In a one month experiment, we simulated the settling of a summer bloom dominated by N. spumigena in sediment microcosms with three species of sediment-dwelling, deposit-feeding macrofauna, the amphipods Monoporeia affinis and Pontoporeia femorata and the bivalve Macoma balthica, and analyzed nodularin in the animals by HPLC–ESI–MS (high-performance liquid chromatography–electrospray ionization–mass spectrometry). We found nodularin in quantities of 50–120 ng g⁻¹ DW. The results show that deposit-feeding macrofauna in the Baltic Sea may contribute to trophic transfer of nodularin.     

Secondary production of the brackish copepod communities and their contribution to the carbon fluxes in the Westerschelde estuary (The Netherlands)

The zooplankton community of the brackish part of the Westerschelde estuary (November 1989–October 1990) was dominated by two calanoid copepods, Eurytemora affinis and Acartia tonsa. Eurytemora was present during a longer period of the year and was much more important in terms of total abundances and biomasses than Acartia.The secondary production of these species was estimated by means of the growth rate method, using weight-specific growth rates obtained from laboratory cultures (Eurytemora) or from the literature (Acartia).Due to the substantially higher growth rates of Acartia compared to Eurytemora, total yearly productions of both communities were comparable, notwithstanding the large discrepancies in biomass. They amounted to about 5 and 6 g C m^–2 y^–1 by Acartia and Eurytemora respectively.The food needed to realise this production was estimated to be about 14 and 17 g C m^–2 y^–1 by Acartia and Eurytemora respectively. Provided that the copepods are able to selectively ingest the phytoplankton, in situ net production provides sufficient carbon for zooplankton demands for a short period of the year only. As phytoplankton standing stock is very low and net phytoplankton productivity is negative from late fall to early spring, nutritional demands of the copepods have to be fulfilled by other than algal food at least during this period of the year.Although the copepods in the brackish part can have an important impact on some food items, their contribution to total carbon fluxes in the brackish zone is negligible: each year some 6% of all consumed carbon in the brackish part of the estuary passes through the copepod food web.     

Mesozooplankton Grazing on Picocyanobacteria in the Baltic Sea as Inferred from Molecular Diet Analysis

Our current knowledge on the microbial component of zooplankton diet is limited, and it is generally assumed that bacteria-sized prey is not directly consumed by most mesozooplankton grazers in the marine food webs. We questioned this assumption and conducted field and laboratory studies to examine picocyanobacteria contribution to the diets of Baltic Sea zooplankton, including copepods. First, qPCR targeting ITS-1 rDNA sequence of the picocyanobacteria Synechococcus spp. was used to examine picocyanobacterial DNA occurrence in the guts of Baltic zooplankton (copepods, cladocerans and rotifers). All field-collected zooplankton were found to consume picocyanobacteria in substantial quantities. In terms of Synechococcus quantity, the individual gut content was highest in cladocerans, whereas biomass-specific gut content was highest in rotifers and copepod nauplii. Moreover, the gut content in copepods was positively related to the picocyanobacteria abundance and negatively to the total phytoplankton abundance in the water column at the time of sampling. This indicates that increased availability of picocyanobacteria resulted in the increased intake of this prey and that copepods may rely more on picoplankton when food in the preferred size range declines. Second, a feeding experiments with a laboratory reared copepod Acartia tonsa fed a mixture of the picocyanobacterium Synechococcus bacillaris and microalga Rhodomonas salina confirmed that copepods ingested Synechococcus, even when the alternative food was plentiful. Finally, palatability of the picocyanobacteria for A. tonsa was demonstrated using uptake of ¹³C by the copepods as a proxy for carbon uptake in feeding experiment with ¹³C-labeled S. bacillaris. These findings suggest that, if abundant, picoplankton may become an important component of mesozooplankton diet, which needs to be accounted for in food web models and productivity assessments.     

Effects of Food Availability and Predation on a Marine Zooplankton Community — a Study on Copepods in the Baltic Sea

The importance of food resources versus predatory effects in determining the abundance pattern of zooplankton varies over time and space. Here, we evaluate the population dynamics of dominant calanoid copepods (Acartia spp. bifilosa/A. longiremis and Eurytemora affinis) in a reference area and a eutrophied area in the northern Baltic Sea Proper. We base the paper on a combination of statistical analyses of a long term (12 years) data set on population biomass dynamics, laboratory experiments on the feeding biology of these species and literature data of their susceptibility to predation. Overall, copepods were equally abundant at both sites, but Eurytemora was more abundant in the eutrophied area while Acartia spp. dominated in the non-eutrophied reference area. There was, however, no significant difference between the two areas in biomass dynamics (biomass specific rate of change) of the two groups of copepods. However, in spring and early summer the population biomass increased faster for Eurytemora as compared to Acartia spp. This is consistent with our findings from feeding and respiration experiments. Eurytemora exhibited generally higher ingestion rates and had a higher growth efficiency (ratio of ingestion to respiration rates), which suggests an advantage over Acartia spp. when food availability increases. Both species generally showed a rapid population decline starting in late summer, most likely caused by intensive predation by fish and mysid shrimps. In late autumn and winter, biomasses of both genera continued to decrease, but the decline was fastest for Eurytemora. During this period of the year, when the food availability is sparse, Acartia spp. may have an advantage over Eurytemora, as we found that the former ingests a broader size range of food particles and it has also been reported to be able to switch from filter feeding to raptorial feeding. This study shows that dominant copepods in the net-zooplankton community in our study area were structured both by food availability and predation. The relative importance of these effects varied seasonally.     

Short communication. Carbon:nitrogen ratios of Baltic Sea copepods-indication of mineral limitation?

The carbon (C) and nitrogen (N) content and the C:N ratio of two common calanoid copepods, Eurytemora affinis and Acartia bifilosa were measured during spring and summer at the SW coast of Finland, northern Baltic Sea. The C:N ratio of both copepod species was slow and stable (4-4.5), irrespective of sampling time, which implies N limitation at least during intermediate to high food concentrations in spring and early summer. In addition, experiments were conducted to reveal whether the diet of copepods affects their C and N content. Adding green algae Brachiomonas submarina in concentrations of 50-500 g C l^-1 to <100-m-filtered sea water did not significantly increase copepod C or N content in 3 day experiments. However, the C and N contents of E.affinis eggs were 2 times higher in high food concentration, which may increase survival of eggs and nauplii.      

Proteomic profiling of the Baltic Sea cyanobacterium Nodularia spumigena strain AV1 during ammonium supplementation

The cyanobacterium Nodularia spumigena dominates the annual, toxic summer blooms in the Baltic Sea. Although Nodularia has been receiving attention due to its production of the hepatotoxin nodularin, molecular data regarding the regulation of nitrogen fixation is lacking. We have previously reported that N. spumigena strain AV1, unlike model filamentous cyanobacteria, differentiates heterocysts in the absence of detectable nitrogen fixation activity. To further analyze the uncoupling between these two linked processes, we assessed the impact of ammonium ions on the N. spumigena metabolism using a proteomic approach. Proteomic profiling was performed at three different times during ammonium supplementation using quantitative 2-dimensional gel electrophoresis followed by MS/MS analysis. Using this approach, we identified 34 proteins, 28 of which were unique proteins that changed successively in abundance during growth on ammonium. Our results indicate that N. spumigena generally exhibits lower energy production and carbon fixation in the presence of ammonium and seems to be inefficient in utilizing ammonium as an external nitrogen source. The possibility of ammonium toxicity due to PSII damage was investigated and the results are discussed. Our findings have implications in regard to the strategies considered to manage the cyanobacterial blooms in the Baltic Sea.     

Settling cyanobacterial blooms do not improve growth conditions for soft bottom meiofauna

Summer blooms of the toxin-producing cyanobacteria Nodularia spumigena are frequent in the Baltic Sea and recent findings suggest that they may be an important food source for the benthos below the euphotic zone. To investigate the effects of settling spring and summer phytoplankton blooms on meiofaunal growth, we assayed concentrations of nucleic acids in three ostracod species (Candona neglecta; Heterocyprideis sorbyana and Paracyprideis fennica) and one genus of nematodes (Paracanthonchus spp.) after incubation in sediments with the one of the following food additions: (1) diatoms, (2) the cyanobacterium Nodularia spumigena, (3) Tetraphyll® as a known high-quality food source, (4) lignin as a refractory artificial food, and (5) control (no added organic matter). The ribosomal ribonucleic acid (RNA) content and RNA:DNA ratios of the tested organisms were lower in the cyanobacteria treatment than in the diatom treatment, with the difference in RNA:DNA ratios being statistically significant for all species except C. neglecta. Moreover, individuals incubated with N. spumigena showed RNA:DNA levels similar to those found in the lignin and control treatments. Furthermore, N. spumigena had lower concentrations of both enzymatically hydrolysable amino acids (EHAA) and eicosapentaenoic acid (EPA) than diatoms suggesting lower nutritional quality for consumers. These results indicate that recently settled summer blooms of N. spumigena are nutritionally poor and do not improve conditions for meiofaunal growth in Baltic sediments. In contrast, input of diatoms to the sediments during spring is crucial for meiofaunal growth.     

Feeding interactions of the copepods Eurytemora affinis and Acartia bifilosa with the cyanobacteria Nodularia sp.

We measured ingestion and clearance rates of two Baltic Seacalanoid copepods, Eurytemora affinis and Acartia bifilosa,on toxic and non-toxic cyanobacteria Nodularia sp. using theisotope technique. Eurytemora affinis fed actively on the non-toxicstrain and moderately actively on the toxic strain, whereasA.bifilosa totally avoided feeding on both strains. This suggeststhat A.bifilosa rejected cyanobacterial filaments due to theirnutritional inadequacy or difficult manageability. The differentresponse of E.affinis to the non-toxic and toxic strains, inturn, shows that this copepod species was able to sense thepresence of the toxin in cyanobacterial filaments and thereforefed less on the toxic strain. The interaction between A.bifilosaand Nodularia sp. was further examined (with the particle countingmethod) by measuring the clearance rates of A.bifilosa on ediblegreen flagellates in the presence of cyanobacteria. The presenceor concentration of toxic Nodularia sp. did not affect grazingrates of A.bifilosa on Brachiomonas submarina. Since earlierstudies have shown that ingestion of Nodularia sp. decreasesegg production and increases mortality in E.affinis, we suggestthat the occurrence of Nodularia sp. blooms in the Baltic Seamay favour individuals of copepod species capable of selectivefeeding, such as A.bifilosa.     

Carbon, nitrogen and phosphorus stoichiometry of crustacean zooplankton in the Baltic Sea: implications for nutrient recycling

The carbon (C), nitrogen (N) and phosphorus (P) contents (%of dry weight) of some crustacean zooplankton were studied inthe Baltic Sea. The copepod Acartia sp. had a stable C and Ncontent (48.3 ± 0.8% C, 12.4 ± 0.2% N, C:N ratio4.5 ± 0.1). The P content was variable (1–2%),probably depending on developmental stage and season. Copepodsaccumulating fat, like Pseudocalanus minutus elongatus, hadhigher and more variable C content (50–60%), and lowerN and P content (7–12% N, 0.6–1.5% P). The highestC and lowest N and P contents were found in adult Limnocalanusmacrurus. However, the N:P ratio was apparently independentof fat content and between 14 and 27 for all copepods. The cladoceransBosmina longispina maritima and Evadne nordmanni had lower Ncontent (9.3–10.8%) and higher C:N ratio (5.1–5.7)than Acartia sp. The P content (1.2–1.4%) was similarto Acartia sp. and the N:P ratios (16–19) were in thelower range of that found for the copepods. The N:P ratio wasgenerally somewhat higher in the copepods than in seston, whichmost of the year had nearly Redfield C:N:P ratios. Potentially,nutrient recycling from crustacean zooplankton could enhanceN limitation of phytoplankton, but small stoichiometric differencessuggest that this effect is probably weak. The extent is dependenton the structure of the zooplankton community and the grossgrowth efficiencies. Acartia copepodites, which had nearly RedfieldN:P ratios, would have the opposite effect and enhance P limitationin late summer when seston N:P ratios increased.     

Trade-Offs between Predation Risk and Growth Benefits in the Copepod Eurytemora affinis with Contrasting Pigmentation

Intraspecific variation in body pigmentation is an ecologically and evolutionary important trait; however, the pigmentation related trade-offs in marine zooplankton are poorly understood. We tested the effects of intrapopulation phenotypic variation in the pigmentation of the copepod Eurytemora affinis on predation risk, foraging, growth, metabolic activity and antioxidant capacity. Using pigmented and unpigmented specimens, we compared (1) predation and selectivity by the invertebrate predator Cercopagis pengoi, (2) feeding activity of the copepods measured as grazing rate in experiments and gut fluorescence in situ, (3) metabolic activity assayed as RNA:DNA ratio in both experimental and field-collected copepods, (4) reproductive output estimated as egg ratio in the population, and (5) total antioxidant capacity. Moreover, mitochondrial DNA (mtDNA) COI gene variation was analysed. The pigmented individuals were at higher predation risk as evidenced by significantly higher predation rate by C. pengoi on pigmented individuals and positive selection by the predator fed pigmented and unpigmented copepods in a mixture. However, the antioxidant capacity, RNA:DNA and egg ratio values were significantly higher in the pigmented copepods, whereas neither feeding rate nor gut fluorescence differed between the pigmented and unpigmented copepods. The phenotypic variation in pigmentation was not associated with any specific mtDNA genotype. Together, these results support the metabolic stimulation hypothesis to explain variation in E. affinis pigmentation, which translates into beneficial increase in growth via enhanced metabolism and antioxidant protective capacity, together with disadvantageous increase in predation risk. We also suggest an alternative mechanism for the metabolic stimulation via elevated antioxidant levels as a primary means of increasing metabolism without the increase in heat absorbance. The observed trade-offs are relevant to evolutionary mechanisms underlying plasticity and adaptation and have the capacity to modify strength of complex trophic interactions.     

The paradox high zooplankton biomass-low vegetal particulate organic matter in high turbidity zones: What way for energy transfer?

Estuarine ecosystems are characterized by high zooplanktonic biomasses, essentially constituted by copepods and mysids whose nutritional requirements are mainly provided by phytoplankton, an easily available carbon form. The Gironde estuary is characterized by high turbidities which limit light penetration in the water column and therefore primary production. Consequently, primary production is low and its availability for higher trophic level is very limited. The main goal of this study was to characterize the total vegetal particulate organic matter (POM) in high turbidity zones of the Gironde estuary during summer (a critical period characterized by high heterotrophic bacterial degradation and high zooplanktonic biomasses) and to analyse its utilization by zooplankton, using prey/predator experiments and trophic biomarkers (fatty acids). The specific goals were to define (i) how vegetal POM was exploited by the different zooplanktonic groups (protozoa, copepods and mysids) and (ii) which alternative preys could be used when vegetal POM was not sufficient to ensure their nutritional requirements.Chlorophyll biomass was very low in the MTZ during summer 2002 (0.48 ± 0.03 mg m^− 3). Total zooplankton grazing was low (19% d^− 1) probably due to a large contribution of detritus originating from terrestrial plants in vegetal POM compared to phytoplankton. The highest grazing pressure was exercised by the mysid Mesopodopsis slabberi due to its high abundances and by its almost entirely herbivorous diet (phytoplankton and small terrestrial detritus). Grazing rates (19.7 ± 4.2 and 9.6 μgC cop^− 1 d^− 1 for juveniles and adults, respectively) seemed to be sufficient to satisfy their daily carbon requirement. Grazing rate of the copepod Eurytemora affinis (139 ngC cop^− 1 d^− 1) seemed to be insufficient to cover its nutritional requirements and the copepods probably needed to complete a great part of their diet from protozoa. Grazing rates of the mysid Neomysis integer (24.7 ± 0.01 and 20.89 ± 8.45 μgC cop^− 1 d^− 1 for juveniles and adults, respectively) were higher than those of M. slabberi when feeding only on phytoplankton. However, when other preys were introduced in its environment, N. integer only fed on the copepod E. affinis with a preference for nauplii. The study revealed the great importance of protozoa and bacteria in the trophic transfers between vegetal POM and zooplankton in the MTZ during summer, despite the low protozoa grazing pressure on vegetal POM (3.1%). The detritic food chain probably implies various trophic transfers with little direct relationships between vegetal POM and zooplankton.     

Feeding dynamics of the copepod Diacyclops thomasi before, during and following filamentous cyanobacteria blooms in a large, shallow temperate lake

Cyanobacteria blooms are an increasing problem in temperate freshwater lakes, leading to reduced water quality and in some cases harmful effects from toxic cyanobacteria species. To better understand the role of zooplankton in modulating cyanobacteria blooms, from 2008 to 2010 we measured water quality and plankton abundance, and measured feeding rates and prey selectivity of the copepod Diacyclops thomasi before, during and following summertime cyanobacteria blooms in a shallow, eutrophic lake (Vancouver Lake, Washington, USA). We used a combined field and experimental approach to specifically test the hypothesis that copepod grazing was a significant factor in establishing the timing of cyanobacteria bloom initiation and eventual decline in Vancouver Lake. There was a consistent annual succession of zooplankton taxa, with cyclopoid copepods (D. thomasi) dominant in spring, followed by small cladocerans (Eubosmina sp.). Before each cyanobacteria bloom, large cladocerans (Daphnia retrocurva, Daphnia laevis) peaked in abundance but quickly disappeared, followed by brief increases in rotifers. During the cyanobacteria blooms, D. thomasi was again dominant, with small cladocerans abundant in autumn. Before the cyanobacteria blooms, D. thomasi substantially consumed ciliates and dinoflagellates (up to 100% of prey biomass per day), which likely allowed diatoms to flourish. A shift in copepod grazing toward diatoms before the blooms may have then helped to facilitate the rapid increase in cyanobacteria. Copepod grazing impact was the highest during the cyanobacteria blooms both years, but focused on non-cyanobacteria prey; copepod grazing was minimal as the cyanobacteria blooms waned. We conclude that cyclopoid copepods may have an indirect role (via trophic cascades) in modulating cyanobacteria bloom initiation, but do not directly contribute to cyanobacteria bloom decline.     

Seasonal abundance, reproduction and development of four key copepod species on the Faroe shelf

The copepod community on the Faroe shelf is dominated by Calanus finmarchicus, Temora longicornis, Acartia longiremis and Pseudocalanus spp. The species composition, abundance and development of the copepod community varied considerably during the season 2004. These variations reflected to a large extent the different life strategies of the copepods. Both nauplii and copepodites of C. finmarchicus were most abundant during spring and early summer. The two neritic copepods T. longicornis and A. longiremis were present in low numbers during spring but dominated the copepod community later during the productive period. Pseudocalanus spp., on the other hand, occurred throughout the year, but showed no clear numerical response in abundance to the spring bloom. The egg production measurements of C. finmarchicus and T. longicornis showed some pre-bloom egg production, but as the spring bloom started the egg production rate increased significantly, especially for C. finmarchicus. There seemed to be a substantial loss of nauplii and copepods from the shelf ecosystem during the productive season. It was, however, not possible to determine whether this was mainly due to mortality or advective loss.     

Glutathione transferase activity and oocyte development in copepods exposed to toxic phytoplankton

Organisms present a series of cellular mechanisms to avoid the effects of toxic compounds. Such mechanisms include the increase in activity of detoxification enzymes [e.g., 7-ethoxyresorufin-O-deethylase (EROD) and glutathione S-transferase (GST)], which could explain the low retention of ingested toxins generally observed in copepods. In addition, decreasing gross growth efficiency (GGE) of copepods with increasing concentration of toxic diets could be caused either by a high expenditure coping with toxins (e.g., increase in the activity of detoxification enzymes) or by a deterioration of reproductive tissues. To assess the effect of toxic phytoplankton on the activity of detoxification enzymes and on oocyte maturation of Acartia tonsa and Temora longicornis, feeding and egg production experiments were carried out with a variety of toxic diets and an adequate non-toxic food control (Rhodomonas spp.) all provided as single species diets. Toxic diets included the nodularin-producing cyanobacterium Nodularia spumigena, the dinoflagellates Alexandrium minutum, and A. tamarense, which contained Paralytic Shellfish Poisoning (PSP) toxins, the dinoflagellate Prorocentrum lima with Diarrhetic Shellfish Poisoning (DSP) toxins and the haptophyte Prymnesium parvum, which produces ichtyotoxins with haemolytic activity. Feeding on toxic diets was lower than on Rhodomonas spp., except for A. minutum and A. tamarense. In addition, toxic diets negatively affected reproduction in both copepod species with the production of oocytes and oocyte development impaired with A. minutum and N. spumigena. While the negative effect of N. spumigena seemed to be connected to gonad atresia likely caused by severe food limitation (starvation), the negative effect of A. minutum could have been either caused by a direct effect of saxitoxins or nutritional inadequacy on oocyte production. We could not detect EROD activity in the copepods, while the activity of GST was generally higher with the non-toxic food control and positively related to the feeding and egestion rates, suggesting relation to feeding conditions rather than to exposure to toxic diets. No relationship was found between GGE and GST activity. Our results refute the hypothesis that toxic diets, provided at ecologically relevant levels, would induce cellular mechanisms in copepods regarding GST activity. GST activity thus seems to play no role in detoxification of copepods confronted with toxic phytoplankton. Toxin detoxification and its cost for copepods still remain an open question.     

Zooplankton feeding ecology: does a diet of Phaeocystis support good copepod grazing, survival, egg production and egg hatching success?

The bloom-forming alga Phaeocystis is ingested by a varietyof zooplankton grazers, but is thought to be a poor source offood. We examined copepod grazing on solitary Phaeocystis cellsby adult females of Temora stylifera, and survival, fecal pelletproduction, egg production and egg hatching success in Calanushelgolandicus and T. stylifera over periods of 15 consecutivedays. Phaeocystis cell concentrations were high (1.2–3.6x 10⁴ cells ml^–1 for C. helgolandicus and 2.5–7.9x 10⁴ cells ml^–1 for T. stylifera), but within the rangeof maxima recorded for natural blooms. Both copepods survivedwell and continuously produced fecal pellets (indicating continuousgrazing) on a diet of Phaeocystis. However, egg production ratesfor both copepods were low, even though hatching success ofthe few eggs produced was high. Clearance rates for T. styliferawere higher than for most previous measurements of other copepodsfeeding on Phaeocystis solitary cells at lower cell concentrations.We conclude that even though copepods feed well upon Phaeocystis,resulting poor fecundity on this diet may inhibit copepod populationincreases during blooms, thereby contributing to the perpetuationof blooms. However, the high egg hatching success on this dietargues against Phaeocystis containing chemical compounds thatact as mitotic inhibitors reducing copepod egg viability, suchas those found in some other phytoplankters.     

Sedimentation of Nodularia spumigena and distribution of nodularin in the food web during transport of a cyanobacterial bloom from the Baltic Sea to the Kattegat

Nodularia spumigena is a toxic cyanobacteria that blooms in the Baltic Sea every year. In the brackish water of the Baltic Sea, its toxin, nodularin, mainly affects the biota in the surface water due to the natural buoyancy of this species. However, the fate of the toxin is unknown, once the cyanobacteria bloom enters the more saline waters of the Kattegat. In order to investigate this knowledge gap, a bloom of N. spumigena was followed during its passage, carried by surface currents, from the Baltic Sea into the Kattegat area, through the Öresund strait. N. spumigena cells showed an increased cell concentration through the water column during the passage of the bloom (up to 130 10³ cells ml⁻¹), and cells (4.2 10³ cells ml⁻¹) could be found down to 20 m depth, below a pycnocline. Sedimentation trap samples from below the pycnocline (10–12 m depth) also showed an increased sedimentation of N. spumigena filaments during the passage of the bloom. The toxin nodularin was detected both in water samples (0.3–6.0 μg l⁻¹), samples of sedimenting material (a toxin accumulation rate of 20 μg m^-2 day⁻¹), zooplankton (up to 0.1 ng ind.⁻¹ in copepods), blue mussels (70–230 μg kg⁻¹ DW), pelagic and benthic fish (herring (1.0–3.4 μg kg⁻¹ DW in herring muscle or liver) and flounder (1.3-6.2 μg kg⁻¹ DW in muscle, and 11.7-26.3 μg kg⁻¹ DW in liver). A laboratory experiment showed that N. spumigena filaments developed a decreased buoyancy at increased salinities and that they were even sinking with a rate of up to 1,7 m day⁻¹ at the highest salinity (32 PSU). This has implications for the fate of brackish water cyanobacterial blooms, when these reach more saline waters. It can be speculated that a significant part of the blooms content of nodularin will reach benthic organisms in this situation, compared to blooms decaying in brackish water, where most of the bloom is considered to be decomposed in the surface waters.