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.     

How does eutrophication affect the role of grazers in harmful algal bloom dynamics?

Population dynamics of harmful algal bloom species are regulated both from the “bottom-up” by factors that affect their growth rate and from the “top-down” by factors that affect their loss rates. While it might seem apparent that eutrophication would have the greatest impact on factors affecting growth rates of phytoplankton (nutrient supply, light availability) the roles of top-down controls, including grazers and pathogens, cannot be ignored in studies of harmful bloom dynamics. Lags between the growth of phytoplankton and zooplankton populations, or disruption of zooplankton populations by adverse environmental conditions may be important factors in the initiation of plankton blooms under eutrophic conditions. Grazers that avoid feeding on harmful species and actively graze on competing species may also play important roles in bloom initiation. Grazers that are not affected by phytoplankton toxins and have growth rates comparable to phytoplankton (e.g. protozoan grazers) may have the potential to control the initiation of blooms. If the inhibition of grazers varies with cell density for blooms of toxic phytoplankton, eutrophication may increase the chances of blooms reaching threshold densities for grazer inhibition. In addition, secondary effects of eutrophication, including hypoxia and change in pH may adversely affect grazer populations, and further release HAB species from top-down control. The Texas brown tide (Aureoumbra lagunensis) blooms provide evidence for the role of grazer disruption in bloom initiation and the importance of high densities of brown tide cells in continued suppression of grazers.     

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.     

Allelopathic effects of the Baltic cyanobacteria Nodularia spumdigena, Aphanizomenon flos-aquae and Anabaena lemmermannii on algal monocultures

Allelopathy, the release of extracellular compounds that inhibit the growth of other microorganisms, may be one factor contributing to the formation and/or maintenance of cyanobacterial blooms. We investigated the allelopathic effects of three cyanobacterial species (Nodularia spumigena, Aphanizomenon flos-aquae and Anabaena lemmermannii) that frequently form mass-occurrences in the Baltic Sea. We exposed monocultures of three phytoplankton species (Thalassiosira weissflogii, Rhodomonas sp. and Prymnesium parvum) to cell-free filtrates of the three cyanobacteria, and quantified allelopathic effects with cell counts. We also investigated the role of the growth phase of cyanobacteria in their allelopathy, by comparing the effects of an exponential and a stationary phase culture of N. spumigena. All tested cyanobacteria inhibited the growth of Rhodomonas sp., but none of them affected P. parvum. The effects on T. weissflogii were more variable, and they were amplified by repeated filtrate additions compared to a single filtrate addition. N. spumigena was more allelopathic in exponential than in stationary growth phase, whereas the culture filtrate was more hepatotoxic in stationary phase. Hepatotoxins were thus probably not involved in the allelopathic effects, which is also indicated by the allelopathic properties of the non-toxic A. flos-aquae and A. lemmermannii. The results demonstrate that the common Baltic cyanobacteria affect some coexisting phytoplankton species negatively. Allelopathy may therefore play a role in interspecific competition and contribute to cyanobacterial bloom maintenance.     

Assimilation of Diazotrophic Nitrogen into Pelagic Food Webs

The fate of diazotrophic nitrogen (N_D) fixed by planktonic cyanobacteria in pelagic food webs remains unresolved, particularly for toxic cyanophytes that are selectively avoided by most herbivorous zooplankton. Current theory suggests that N_D fixed during cyanobacterial blooms can enter planktonic food webs contemporaneously with peak bloom biomass via direct grazing of zooplankton on cyanobacteria or via the uptake of bioavailable N_D (exuded from viable cyanobacterial cells) by palatable phytoplankton or microbial consortia. Alternatively, N_D can enter planktonic food webs post-bloom following the remineralization of bloom detritus. Although the relative contribution of these processes to planktonic nutrient cycles is unknown, we hypothesized that assimilation of bioavailable N_D (e.g., nitrate, ammonium) by palatable phytoplankton and subsequent grazing by zooplankton (either during or after the cyanobacterial bloom) would be the primary pathway by which N_D was incorporated into the planktonic food web. Instead, in situ stable isotope measurements and grazing experiments clearly documented that the assimilation of N_D by zooplankton outpaced assimilation by palatable phytoplankton during a bloom of toxic Nodularia spumigena Mertens. We identified two distinct temporal phases in the trophic transfer of N_D from N. spumigena to the plankton community. The first phase was a highly dynamic transfer of N_D to zooplankton with rates that covaried with bloom biomass while bypassing other phytoplankton taxa; a trophic transfer that we infer was routed through bloom-associated bacteria. The second phase was a slowly accelerating assimilation of the dissolved-N_D pool by phytoplankton that was decoupled from contemporaneous variability in N. spumigena concentrations. These findings provide empirical evidence that N_D can be assimilated and transferred rapidly throughout natural plankton communities and yield insights into the specific processes underlying the propagation of N_D through pelagic food webs.     

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.     

The Thiamine Content of Phytoplankton Cells Is Affected by Abiotic Stress and Growth Rate

Thiamine (vitamin B₁) is produced by many plants, algae and bacteria, but by higher trophic levels, it must be acquired through the diet. We experimentally investigated how the thiamine content of six phytoplankton species belonging to five different phyla is affected by abiotic stress caused by changes in temperature, salinity and photon flux density. Correlations between growth rate and thiamine content per cell were negative for the five eukaryotic species, but not for the cyanobacterium Nodularia spumigena. We demonstrate a high variability in thiamine content among phytoplankton species, with the highest content in N. spumigena. Salinity was the factor with the strongest effect, followed by temperature and photon flux density, although the responses varied between the investigated phytoplankton species. Our results suggest that regime shifts in phytoplankton community composition through large-scale environmental changes has the potential to alter the thiamine availability for higher trophic levels. A decreased access to this essential vitamin may have serious consequences for aquatic food webs.     

Bloom-Forming Cyanobacteria Support Copepod Reproduction and Development in the Baltic Sea

It is commonly accepted that summer cyanobacterial blooms cannot be efficiently utilized by grazers due to low nutritional quality and production of toxins; however the evidence for such effects in situ is often contradictory. Using field and experimental observations on Baltic copepods and bloom-forming diazotrophic filamentous cyanobacteria, we show that cyanobacteria may in fact support zooplankton production during summer. To highlight this side of zooplankton-cyanobacteria interactions, we conducted: (1) a field survey investigating linkages between cyanobacteria, reproduction and growth indices in the copepod Acartia tonsa; (2) an experiment testing relationships between ingestion of the cyanobacterium Nodularia spumigena (measured by molecular diet analysis) and organismal responses (oxidative balance, reproduction and development) in the copepod A. bifilosa; and (3) an analysis of long term (1999–2009) data testing relationships between cyanobacteria and growth indices in nauplii of the copepods, Acartia spp. and Eurytemora affinis, in a coastal area of the northern Baltic proper. In the field survey, N. spumigena had positive effects on copepod egg production and egg viability, effectively increasing their viable egg production. By contrast, Aphanizomenon sp. showed a negative relationship with egg viability yet no significant effect on the viable egg production. In the experiment, ingestion of N. spumigena mixed with green algae Brachiomonas submarina had significant positive effects on copepod oxidative balance, egg viability and development of early nauplial stages, whereas egg production was negatively affected. Finally, the long term data analysis identified cyanobacteria as a significant positive predictor for the nauplial growth in Acartia spp. and E. affinis. Taken together, these results suggest that bloom forming diazotrophic cyanobacteria contribute to feeding and reproduction of zooplankton during summer and create a favorable growth environment for the copepod 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.     

121 Influence of Benthic vs. Pelagic Seeding on Coastal Diatom Bloom Development

Diatom blooms may be initiated by cells that have survived in the plankton or germinated resting stages from the sediments. However, it is not well understood how these different inocula contribute to bloom development. We followed diatom community development in twenty‐liter microcosms given different inocula. Surface sediment and phytoplankton were collected in Gullmar Fjord, Sweden. Replicate microcosms were then dosed with local sediment and/or plankton and incubated in situ in Gullmar Fjord. We also followed the concurrent development of the phytoplankton community in the fjord. Experiments run in both spring and fall 2002 showed that bloom development in the microcosms was significantly faster when seeded by planktonic cells. However, addition of sediment may have stimulated planktonic growth and also provided additional propagules. The type of inoculum used strongly influenced the diatom composition in the microcosms. Sediment additions, through germination of resting stages, resulted in communities dominated by Detonula confervacea and Thalassiosira minima in spring, and Skeletonema costatum in fall. Planktonic inocula resulted in blooms of T. nordenskioeldii and Chaetoceros debilis in spring, and S. costatum and several Chaetoceros spp. in fall. Microcosms dosed with both plankton and sediment showed a mixed species assemblage. Comparisons between the microcosms and fjord phytoplankton suggest an important role for benthic seeding of diatom blooms.     

Nitrogen Fixed By Cyanobacteria Is Utilized By Deposit-Feeders

Benthic communities below the photic zone depend for food on allochthonous organic matter derived from seasonal phytoplankton blooms. In the Baltic Sea, the spring diatom bloom is considered the most important input of organic matter, whereas the contribution of the summer bloom dominated by diazotrophic cyanobacteria is less understood. The possible increase in cyanobacteria blooms as a consequence of eutrophication and climate change calls for evaluation of cyanobacteria effects on benthic community functioning and productivity. Here, we examine utilization of cyanobacterial nitrogen by deposit-feeding benthic macrofauna following a cyanobacteria bloom at three stations during two consecutive years and link these changes to isotopic niche and variations in body condition (assayed as C:N ratio) of the animals. Since nitrogen-fixing cyanobacteria have δ¹⁵N close to -2‰, we expected the δ¹⁵N in the deposit-feeders to decrease after the bloom if their assimilation of cyanobacteria-derived nitrogen was substantial. We also expected the settled cyanobacteria with their associated microheterotrophic community and relatively high nitrogen content to increase the isotopic niche area, trophic diversity and dietary divergence between individuals (estimated as the nearest neighbour distance) in the benthic fauna after the bloom. The three surface-feeding species (Monoporeia affinis, Macoma balthica and Marenzelleria arctia) showed significantly lower δ¹⁵N values after the bloom, while the sub-surface feeder Pontoporeia femorata did not. The effect of the bloom on isotopic niche varied greatly between stations; populations which increased niche area after the bloom had better body condition than populations with reduced niche, regardless of species. Thus, cyanobacterial nitrogen is efficiently integrated into the benthic food webs in the Baltic, with likely consequences for their functioning, secondary production, transfer efficiency, trophic interactions, and intra- and interspecific competition.     

The Diversity of Harmful Algal Bloom-Triggering Mechanisms and the Complexity of Bloom Initiation

Mechanisms influencing initiation of harmful algal blooms (HABs) are diverse, and are not likely to be mutually exclusive. Rather, initiation of HABs is a result of interactions between processes, which result in biological, physical, and chemical conditions optimal for a bloom. Due to the complexity of some bloom initiation processes, bloom-preventative management may be possible. Results from a modeling exercise and a laboratory experiment indicated that a phytoplankton bloom could be circumvented through manipulation of the nutrient-loading mode, i.e., pulsed vs. continuous loading. These findings, should they prove consistent in more robust field experiments, may provide insights for the development of new management approaches for some HABs. Optimal bloom conditions, however, vary between HAB species. Consequently, it is unlikely that a single management solution will exist. Preventative management efforts will require early warning of HAB initiation, perhaps even before the appearance of an HAB species. An indicator based on the dynamic nature of phytoplankton succession events and phytoplankton species diversity may prove useful for this purpose. Applying this index to an existing plankton data set showed that Microcystis blooms might have been predicted months before the start of the bloom.     

Resolving phytoplankton taxa based on high-throughput sequencing during brown tides in the Bohai Sea,China

Large-scale blooms formed by pico-sized phytoplankton, which strongly inhibited feeding activity and growth of cultured scallops, have been recorded along the coast of Qinhuangdao in the Bohai Sea since 2009. Based on pigment profiles and clone library analysis of phytoplankton samples during the blooms, the major bloom-forming species was primarily identified as pelagophyte Aureococcus anophagefferens Hargraves et Sieburth, the causative species of intensive brown tides in the United States and South Africa. Due to the indistinct morphological features of the bloom-forming microalgae and limited knowledge on the composition of phytoplankton communities, there were still disputes concerning the causative species of the blooms. In this study, the method of high-throughput sequencing targeted 18S rDNA V4 region was used to study the composition of pico- and nano-sized phytoplankton communities in 2013 and 2014. A total of 18 groups of eukaryotic microalgae at the class level and more than 2000 operational taxonomic units (OTUs) were identified in phytoplankton samples collected from the brown-tide zone in the Qinhuangdao coastal waters. For both years, A. anophagefferens was the most dominant species during the bloom period and its maximum relative abundance exceeded 60 percent. Along with other evidence, the results further confirm that A. anophagefferens is the major causative species of the pico-sized phytoplankton blooms in the Bohai Sea. The outbreak of brown tide exhibited a strong inter-annual variation between 2013 and 2014, and an increasing dominance of dinoflagellates could be observed in the Qinhuangdao coastal waters.     

Nutrient limitation of Microcystis aeruginosa in northern California Klamath River reservoirs

Nutrient limitations were investigated in Copco and Iron Gate Reservoirs, on the Klamath River in California, where blooms of the toxin-producing cyanobacterium Microcystis aeruginosa were first reported in 2005. Nutrient enrichment experiments conducted in situ in June and August, 2007 and 2008, determined responses in phytoplankton biomass, Microcystis abundance and microcystin concentration to additions of phosphorus and different forms of nitrogen (NH₄⁺, NO₃, and urea). Microcystis abundance was determined using quantitative PCR targeting the phycocyanin intergenic spacer cpcBA.Total phytoplankton biomass increased with additions of N both before and during Microcystis blooms, with no primary effects from P, suggesting overall N limitation for phytoplankton growth during the summer season. NH₄⁺ generally produced the greatest response in phytoplankton growth, while Microcystis abundance increased in response to all forms of N. Microcystis doubling time in the in situ experiments was 1.24–1.39 days when N was not limiting growth. The results from this study suggest availability of N during the summer is a key growth-limiting factor for the initiation and maintenance of toxic Microcystis blooms in Copco and Iron Gate Reservoirs in the Klamath River.     

Long term nutrient loads and chlorophyll dynamics in a large temperate Australian lagoon system affected by recurring blooms of cyanobacteria

While rare globally, blooms of the toxic cyanobacteria Nodularia spumigena are a recurring problem in a few estuaries, such as the Baltic Sea and several southern Australian estuaries. Here, we document recurring Nodularia spumigena Mertens blooms in the Gippsland Lakes, S.E. Australia; a temperate lagoon system with episodic, winter-spring dominated catchment inflows. Physico-chemical conditions exerted a strong influence over bloom development, with blooms consistently occurring at surface water salinities between 9 and 20 (average?=?15), inorganic nitrogen concentrations <0.4?μM, and inorganic nitrogen to reactive phosphorus ratios <5. There was a positive correlation between average annual chlorophyll a and total phosphorus (TP) load in years when there was no Nodularia bloom, but this relationship broke down in Nodularia bloom years, even though there was a strong correlation between in-lake TP and chlorophyll a during these years; this highlights the importance of internal sources of phosphorus to bloom development. Large catchment derived nitrate and nitrite (NO_x) inputs following wildfires and floods in 2007, led to high concentrations of NO_x within the surface waters of the Gippsland Lakes through the second half of 2007 and the start of 2008. We hypothesise that these high NO_x concentrations were a key factor leading to an unprecedented Synechococcus sp. bloom that developed in the austral summer of 2007–2008, despite conditions that would otherwise favour a Nodularia bloom.     

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.     

Fungal Parasitism: Life Cycle,Dynamics and Impact on Cyanobacterial Blooms

Many species of phytoplankton are susceptible to parasitism by fungi from the phylum Chytridiomycota (i.e. chytrids). However, few studies have reported the effects of fungal parasites on filamentous cyanobacterial blooms. To investigate the missing components of bloom ecosystems, we examined an entire field bloom of the cyanobacterium Anabaena macrospora for evidence of chytrid infection in a productive freshwater lake, using a high resolution sampling strategy. A. macrospora was infected by two species of the genus Rhizosiphon which have similar life cycles but differed in their infective regimes depending on the cellular niches offered by their host. R. crassum infected both vegetative cells and akinetes while R. akinetum infected only akinetes. A tentative reconstruction of the developmental stages suggested that the life cycle of R. crassum was completed in about 3 days. The infection affected 6% of total cells (and 4% of akinètes), spread over a maximum of 17% of the filaments of cyanobacteria, in which 60% of the cells could be parasitized. Furthermore, chytrids may reduce the length of filaments of Anabaena macrospora significantly by “mechanistic fragmentation” following infection. All these results suggest that chytrid parasitism is one of the driving factors involved in the decline of a cyanobacteria blooms, by direct mortality of parasitized cells and indirectly by the mechanistic fragmentation, which could weaken the resistance of A. macrospora to grazing.     

Abiotic factors influencing cyanobacterial bloom development in the Gulf of Finland (Baltic Sea)

Blooms of cyanobacteria are a recurrent phenomenon in the Baltic Sea, including the Gulf of Finland. The spatial extension, duration, intensity and species composition of these blooms varies widely between years. Alg@line data collected regularly from ferries as well as weather service and marine monitoring data from 1997 to 2005 are analysed to determine the main abiotic factors influencing the intensity and species composition of cyanobacterial blooms in the Gulf of Finland. It is demonstrated that the development of the Nodularia spumigena Mertens bloom is highly dependent on weather conditions such as photosynthetically active radiation and water temperature. Nutrient conditions, especially the surplus of phosphorus (according to Redfield ratio) related to the pre-bloom upwelling events in the Gulf, affect the intensity of Aphanizomenon sp. (L.) Ralfs blooms. Differences in bloom timing and duration indicate that, if the preconditions (like nutrient ratio/concentration and weather conditions) for bloom formation are favourable, then the Aphanizomenon bloom starts earlier, the overall bloom period is longer and the Nodularia peak might appear in a wider time window. Handling editor: K. Martens     

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.