Cyanobacteria and cyanotoxins: the influence of nitrogen versus phosphorus

The importance of nitrogen (N) versus phosphorus (P) in explaining total cyanobacterial biovolume, the biovolume of specific cyanobacterial taxa, and the incidence of cyanotoxins was determined for 102 north German lakes, using methods to separate the effects of joint variation in N and P concentration from those of differential variation in N versus P. While the positive relationship between total cyanobacteria biovolume and P concentration disappeared at high P concentrations, cyanobacteria biovolume increased continually with N concentration, indicating potential N limitation in highly P enriched lakes. The biovolumes of all cyanobacterial taxa were higher in lakes with above average joint NP concentrations, although the relative biovolumes of some Nostocales were higher in less enriched lakes. Taxa were found to have diverse responses to differential N versus P concentration, and the differences between taxa were not consistent with the hypothesis that potentially N(2)-fixing Nostocales taxa would be favoured in low N relative to P conditions. In particular Aphanizomenon gracile and the subtropical invasive species Cylindrospermopsis raciborskii often reached their highest biovolumes in lakes with high nitrogen relative to phosphorus concentration. Concentrations of all cyanotoxin groups increased with increasing TP and TN, congruent with the biovolumes of their likely producers. Microcystin concentration was strongly correlated with the biovolume of Planktothrix agardhii but concentrations of anatoxin, cylindrospermopsin and paralytic shellfish poison were not strongly related to any individual taxa. Cyanobacteria should not be treated as a single group when considering the potential effects of changes in nutrient loading on phytoplankton community structure and neither should the N(2)-fixing Nostocales. This is of particular importance when considering the occurrence of cyanotoxins, as the two most abundant potentially toxin producing Nostocales in our study were found in lakes with high N relative to P enrichment.     

Deriving nutrient targets to prevent excessive cyanobacterial densities in U.S. lakes and reservoirs

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Increased risk of cyanobacterial blooms in northern high‐latitude lakes through climate warming and phosphorus enrichment

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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.     

Nutrient addition effects on chlorophyll a,phytoplankton biomass,and heterocyte formation in Lake Erie’s central basin during 2014–2017: Insights into diazotrophic blooms in high nitrogen water

1. Phosphorus (P) usually is the primary limiting nutrient of phytoplankton biomass, but attention towards nitrogen (N) and trace nutrients, such as iron (Fe), has surfaced. Additionally, N-fixing cyanobacterial blooms have been documented to occur in N-rich, P-poor waters, which is counterintuitive from the paradigm that low N and high P promotes blooms. For example, Lake Erie's central basin has Dolichospermum blooms when nitrate concentrations are high, which raises questions about which nutrient(s) are selecting for Dolichospermum over other phytoplankton and why an N-fixer is present in high N waters?
2. We conducted a 4-year (2014–2017) study in Lake Erie's central basin to determine which nutrient (P, N, or trace nutrients such as Fe, molybdenum [Mo], and boron [B]) constrained chlorophyll concentration, phytoplankton biovolume, and nitrate assimilation using nutrient enrichment bioassays. The enriched lake water was incubated in 1-L bottles in a growth chamber programmed at light and temperatures of in situ conditions for 4–7 days. We also quantified heterocytes when N-fixing cyanobacteria were present.
3. Compared to the non-enriched control, the P-enriched (+P) treatment had significantly higher chlorophyll and phytoplankton biovolume in c. 75% of experiments. Combination enrichments of P with ammonium-N, nitrate-N, Fe, Mo, and B were compared to the +P treatment to determine secondary limitations. +P and ammonium-N and +P nitrate-N resulted in higher chlorophyll in 50% of experiments but higher phytoplankton biovolume in only 25% of experiments. These results show that P was the primary limiting nutrient, but there were times when N was secondarily limiting.
4. Chlorophyll concentration indicated N secondary limitation in half of the experiments, but biovolume indicated only N secondary limitation in 25% of the experiments. To make robust conclusions from nutrient enrichment bioassays, both chlorophyll and phytoplankton biovolume should be measured.
5. The secondary effects of Fe, Mo, and B on chlorophyll were low (<26% of experiments), and no secondary effects were observed on phytoplankton biovolume and nitrate assimilation. However, +P and Fe resulted in more chlorophyll than +P in experiments conducted during Dolichospermum blooms, and +P and B significantly increased the number of heterocytes in Dolichospermum. These results indicate that low Fe availability might select for Dolichospermum, and low B constrains heterocyte formation in the central basin of Lake Erie. Furthermore, these results could apply to other lakes with high N and low P where diazotrophic cyanobacterial blooms occur.

     

Temporal and spatial variability in the phytoplankton community of Myall Lakes,Australia, and influences of salinity

The variability in the phytoplankton communities of the Myall Lakes, a series of four interconnected coastal lakes on the lower north coast of New South Wales, was studied between 1999 and 2002. There was considerable spatial variability across the lake system. Bombah Broadwater experienced blooms of Anabaena in 1999 and early 2000, but these were replaced from late 2000 onwards by Chroococcus and a variety of eukaryotic taxa, particularly flagellates and diatoms. In comparison, the phytoplankton community of Myall Lake was dominated for much of the study period by Chroococcus, Merismopedia and chlorophyte taxa. The sites located midway between these two main lakes represent an ecotone, with elements of the phytoplanktonic flora of both main lakes being present. Changes in phytoplankton community composition in Bombah Broadwater occurred fairly frequently. In contrast, the phytoplankton community in Myall Lake changed little during the course of the study and can be considered as being at long-term equilibrium. The reasons for this lie in the morphology and hydrology of the lake system, which in turn create gradients in a number of physico-chemical water quality attributes. Bombah Broadwater is influenced by episodic and stochastic freshwater inflows from the upper Myall River catchment, and in times of drought by saline marine incursions via the lower Myall River. Myall Lake however represents a cul-de-sac, with only a small hydraulic connection to the remainder of the lake system. As it has little input from its small catchment, the limnological conditions within this lake remain relatively constant for long periods of time. Although no patterns of seasonal succession were discernable in any of the lakes, some longer-term (annual) changes did occur, and certain taxa displayed enhanced growth in summer. Salinity was found to be an important factor in determining phytoplankton community composition and abundance. Canonical Correspondence Analysis of phytoplankton and environmental data for all sites combined, showed ammonia, total nitrogen and salinity (measured as electrical conductivity) to have the most influence on the phytoplankton community composition and abundance. Anabaena growth was positively related to ammonia concentration and negatively related to conductivity.     

Lake responses to long‐term disturbances and management practices

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The role of environmental parameters in the structure of phytoplankton assemblages and cyanobacteria toxins in two hypereutrophic lakes

We evaluated the variability of cyanotoxins, water chemistry, and cyanobacteria communities in two hypereutrophic drowned river mouth lakes (Spring Lake and Mona Lake; summer 2006) in west Michigan, USA. Even with considerable geographical and watershed similarity, local variations in nutrient concentrations and environmental factors were found to influence the differences observed in cyanobacteria assemblages and cyanotoxins levels between the two lakes. Limnothrix sp. dominated the phytoplankton community in Spring Lake (82% of biovolume) and was negatively correlated with total phosphorus (TP) concentrations. Although Spring Lake was treated with alum during the previous year, Limnothrix sp. was able to bloom in the lower P environment. In contrast, the N₂-fixing cyanobacterium, Anabaena flos-aquae, dominated the phytoplankton in Mona Lake (64% of biovolume). N₂-fixing cyanobacteria dominance in Mona Lake was correlated with higher TP lower dissolved nitrogen levels. Cylindrospermopsis raciborskii was found in both systems; however, the toxin-producing polyketide synthetase gene was not present in either population. The higher TP in Mona Lake appeared to account for the 3-fold increase in cyanobacteria biovolume. Restoration plans for both lakes should include assessments of internal loading and continued phytoplankton monitoring to track the temporal distribution of cyanobacteria species and cyanotoxin concentrations.     

Phototrophic Picoplankton in Temperate Lakes: Seasonal Abundance and Importance along a Trophic Gradient

The seasonal development of autotrophic picoplankton was investigated in seven Danish lakes representing a eutrophication gradient. Highest cell abundance between 1.5 to 6 × 10⁵ cells ml⁻¹ were found in mid-summer. Minor peaks were observed in spring. In winter, densities were below 10³ ml⁻¹. The highest relative picoplankton contribution to total autotrophic biomass also occurred in mid-summer. In the eutrophic lakes and one humic lake the average seasonal contribution of picoplankton to total chlorophyll was below 1% increasing to 5-8% in the meso- and oligotrophic clear water lakes. During short periods the proportion of picoplankton did reach 25%. The higher relative importance of picoplankton in less productive lakes was not due to higher actual chlorophyll concentrations, but due to a much more pronounced response by larger algae at higher nutrient loading. Both cyanobacteria and eukaryote organisms were present as picoplankton. Only eukaryotes were found in one eutrophic lake and an acidic, humic lake. In the eutrophic lakes eukaryote picoplankton was dominant; both with respect to cell densities and biovolume, whereas cyanobacteria dominated the two meso-oligotrophic lakes. Autotrophic picoplankton were present in all lake types, however their importance seemed to be less in most eutrophic lakes than in less productive, meso-oligotrophic lakes.     

Browning affects pelagic productivity in northern lakes by surface water warming and carbon fertilization

Global change impacts important environmental drivers for pelagic gross primary production (GPP) in northern lakes, such as temperature, light, nutrient, and inorganic carbon availability. Separate and/or synergistic impacts of these environmental drivers on pelagic GPP remain largely unresolved. Here, we assess key drivers of pelagic GPP by combining detailed depth profiles of summer pelagic GPP with environmental and climatic data across 45 small and shallow lakes across northern Sweden (20 boreal, 6 subarctic, and 19 arctic lakes). We found that across lakes summer pelagic GPP was strongest associated with lake water temperatures, lake carbon dioxide (CO₂) concentrations impacted by lake water pH, and further moderated by dissolved organic carbon (DOC) concentrations influencing light and nutrient conditions. We further used this dataset to assess the extent of additional DOC-induced warming of epilimnia (here named internal warming), which was especially pronounced in shallow lakes (decreasing 0.96°C for every decreasing m in average lake depth) and increased with higher concentrations of DOC. Additionally, the total pools and relative proportion of dissolved inorganic carbon and DOC, further influenced pelagic GPP with drivers differing slightly among the boreal, subarctic and Arctic biomes. Our study provides novel insights in that global change affects pelagic GPP in northern lakes not only by modifying the organic carbon cycle and light and nutrient conditions, but also through modifications of inorganic carbon supply and temperature. Considering the large-scale impacts and similarities of global warming, browning and recovery from acidification of lakes at higher latitudes throughout the northern hemisphere, these changes are likely to operate on a global scale.     

Estimating microcystin levels at recreational sites in western Lake Erie and Ohio

Cyanobacterial harmful algal blooms (cyanoHABs) and associated toxins, such as microcystin, are a major global water-quality issue. Water-resource managers need tools to quickly predict when and where toxin-producing cyanoHABs will occur. This could be done by using site-specific models that estimate the potential for elevated toxin concentrations that cause public health concerns. With this study, samples were collected at three Ohio lakes to identify environmental and water-quality factors to develop linear-regression models to estimate microcystin levels. Measures of the algal community (phycocyanin, cyanobacterial biovolume, and cyanobacterial gene concentrations) and pH were most strongly correlated with microcystin concentrations. Cyanobacterial genes were quantified for general cyanobacteria, general Microcystis and Dolichospermum, and for microcystin synthetase (mcyE) for Microcystis, Dolichospermum, and Planktothrix. For phycocyanin, the relations were different between sites and were different between hand-held measurements on-site and nearby continuous monitor measurements for the same site. Continuous measurements of parameters such as phycocyanin, pH, and temperature over multiple days showed the highest correlations to microcystin concentrations. The development of models with high R² values (0.81–0.90), sensitivities (92%), and specificities (100%) for estimating microcystin concentrations above or below the Ohio Recreational Public Health Advisory level of 6 μg L⁻¹ was demonstrated for one site; these statistics may change as more data are collected in subsequent years. This study showed that models could be developed for estimates of exceeding a microcystin threshold concentration at a recreational freshwater lake site, with potential to expand their use to provide relevant public health information to water resource managers and the public for both recreational and drinking waters.     

Effect of Environmental Factors on Cyanobacteria Richness in Some Agricultural Soils

The physicochemical variations of soil, such as temperature, pH, nutrients, and the type of plant cultivation, affect the diversity of cyanobacteria, whether heterocystous or not. The aim of this study was to identify the species of cyanobacteria in a soil and the effect of environmental characteristics on cyanobacteria. Soil samples collected from six different agricultural sites in Al Diwaniyah city/Iraq during September 2016 in the autumn season were analyzed, and the physicochemical characteristics of the samples were measured using approved methods.

The results showed significant correlation and differences between cyanobacteria composition, distribution, and physicochemical factors among soil sites. The Agricultural soil was slightly alkaline and moderately saline and contained abundant nutrients, cations and a high percentage of organic matter. All these characteristics influenced the distribution and diversity of cyanobacteria. Ninety-six species were identified, including four heterocystous species represented by Anabaena, Calothrix, Cylnidrospermum, and Nostoc. However, the non-heterocystous were represented by 13 species: Aphanocapsa, Aphanothece, Arthrospira, Chroococcus, Gloeocapsa, Lyngbya, Merismopedia, Microcystis, Microcoleus, Oscillatoria, Phormidium, Schizothrix, and Spirulina. The dominant species of cyanobacteria was Oscillatoria, followed by Phormidium, Chroococcus, Gleocapsa and Lyngbya. The highest value of Shannon’s and Simpson’s diversities were registered in the Ghammas site, which is a paddy field, but the lowest was registered in the Afak site, cultivated with the alfalfa plant. Soil was classified as finely textured with silty clayey characterization, favorable for cyanobacteria growth.     

The ecology of the planktonic diatom Cyclotella and its implications for global environmental change studies

The fossil record of diatoms in lake sediments can be used to assess the effects of climate variability on lake ecosystems if ecological relationships between diatom community structure and environmental parameters are well understood. Cyclotella sensu lato taxa are a key group of diatoms that are frequently dominant members of phytoplankton communities in low‐ to moderate‐productivity lakes. Their relative abundances have fluctuated significantly in palaeolimnological records spanning over a century in arctic, alpine, boreal and temperate lakes. This suggests that these species are sensitive to environmental change and may serve as early indicators of ecosystem effects of global change. Yet patterns of change in Cyclotella species are not synchronous or unidirectional across, or even within, regions, raising the question of how to interpret these widespread changes in diatom community structure. We suggest that the path forward in resolving seemingly disparate records is to identify clearly the autecology of Cyclotella species, notably the role of nutrients, dissolved organic carbon and light, coupled with better consideration of both the mechanisms controlling lake thermal stratification processes and the resulting effects of changing lake thermal regimes on light and nutrients. Here we begin by reviewing the literature on the resource requirements of common Cyclotella taxa, illustrating that many studies reveal the importance of light, nitrogen, phosphorus, and interactions among these resources in controlling relative abundances. We then discuss how these resource requirements can be linked to shifts in limnological processes driven by environmental change, including climate‐driven change in lakewater temperature, thermal stratification and nutrient loading, as well as acidification‐driven shifts in nutrients and water clarity. We examine three case studies, each involving two lakes from the same region that have disparate trends in the relative abundances of the same species, and illustrate how the mechanisms by which these species abundances are changing can be deciphered. Ultimately, changes in resource availability and water clarity are key factors leading to shifts in Cyclotella abundances. Tighter integration of the autecology of this important group of diatoms with environmental change and subsequent alterations in limnological processes will improve interpretations of palaeolimnological records, and clarify the drivers of seemingly disparate patterns in fossil records showing widespread and rapid changes across the northern hemisphere.     

Light availability as a possible regulator of cyanobacteria species composition in a shallow subtropical lake

1. Variations in the relative biovolumes of dominant cyanobacterial taxa were evaluated in the context of environmental conditions using canonical correlation analysis (CCorrA) and Redundancy Analysis (RDA). The objective was to test a conceptual model in which underwater irradiance determines dominance by bloom-forming (high light adapted) or non-blooming (low light adapted) taxa. 2. The data set consisted of 404 contiguous observations, collected over a 3-year period at eight pelagic sites, in shallow Lake Okeechobee, Florida, U.S.A. Data included species biovolumes, total phosphorus (TP), total nitrogen (TN), dissolved oxygen (DO) and chlorophyll a concentrations, as well as two indices: underwater irradiance (Secchi depth) and the ratio of Secchi:total depth. 3. The first environmental canonical variable was strongly correlated with the two light-related indices, and negatively correlated with TP. This reflects the predominant role of resuspended P-rich lake sediments in controlling underwater irradiance in the shallow lake. The first species canonical variable displayed a strong negative correlation with Lyngbya limnetica and L. contorta, and positive correlations with Anabaena circinalis, Aphanizomenon flos aquae and Microcystis spp. The results support the conceptual model; the first pair of canonical variables explained 55% of the variation in the species–environmental data set. RDA results provided further support for the hypothesis that irradiance was the major force controlling community structure. 4. One unexpected result was a positive association between Oscillatoria spp. dominance and indicators of high irradiance. This conflicts with past research indicating that Oscillatoria is a low light adapted taxon, and the finding that it is the most abundant taxon in Lake Okeechobee. This may reflect the fact that the two Lyngbya taxa were more strongly associated with low light conditions than Oscillatoria. CCorrA results indicated that Oscillatoria densities are strongly controlled by water temperature. There is a need for more detailed studies of cyanobacteria ecophysiology in order to explain fully the seasonality of phytoplankton in this and other shallow subtropical lakes.     

Inverse relationship of epilithic algae and pelagic phosphorus in unproductive lakes: Roles of N2 fixers and light

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Increased nutrient loading and rapid changes in phytoplankton expected with climate change in stratified South European lakes: sensitivity of lakes with different trophic state and catchment properties

We hypothesised that increasing winter affluence and summer temperatures, anticipated in southern Europe with climate change, will deteriorate the ecological status of lakes, especially in those with shorter retention time. We tested these hypotheses analysing weekly phytoplankton and chemistry data collected over 2 years of contrasting weather from two adjacent stratified lakes in North Italy, differing from each other by trophic state and water retention time. Dissolved oxygen concentrations were higher in colder hypolimnia of both lakes in the second year following the cold winter, despite the second summer was warmer and the lakes more strongly stratified. Higher loading during the rainy winter and spring increased nutrient (N, P, Si) concentrations, and a phytoplankton based trophic state index, whilst the N/P ratio decreased in both lakes. The weakened Si limitation in the second year enabled an increase of diatom biovolumes in spring in both lakes. Chlorophyll a concentration increased in the oligo-mesotrophic lake, but dropped markedly in the eutrophic lake where the series of commonly occurring cyanobacteria blooms was interrupted. The projected increase of winter precipitation in southern Europe is likely to increase the nutrient loadings to lakes and contribute to their eutrophication. The impact is proportional to the runoff/in-lake concentration ratio of nutrients rather than to the retention time, and is more pronounced in lakes with lower trophy.     

Lake responses to reduced nutrient loading – an analysis of contemporary long-term data from 35 case studies

1. This synthesis examines 35 long‐term (5–35 years, mean: 16 years) lake re‐oligotrophication studies. It covers lakes ranging from shallow (mean depth <5 m and/or polymictic) to deep (mean depth up to 177 m), oligotrophic to hypertrophic (summer mean total phosphorus concentration from 7.5 to 3500 μg L^?1 before loading reduction), subtropical to temperate (latitude: 28–65°), and lowland to upland (altitude: 0–481 m). Shallow north‐temperate lakes were most abundant. 2. Reduction of external total phosphorus (TP) loading resulted in lower in‐lake TP concentration, lower chlorophyll a (chl a) concentration and higher Secchi depth in most lakes. Internal loading delayed the recovery, but in most lakes a new equilibrium for TP was reached after 10–15 years, which was only marginally influenced by the hydraulic retention time of the lakes. With decreasing TP concentration, the concentration of soluble reactive phosphorus (SRP) also declined substantially. 3. Decreases (if any) in total nitrogen (TN) loading were lower than for TP in most lakes. As a result, the TN : TP ratio in lake water increased in 80% of the lakes. In lakes where the TN loading was reduced, the annual mean in‐lake TN concentration responded rapidly. Concentrations largely followed predictions derived from an empirical model developed earlier for Danish lakes, which includes external TN loading, hydraulic retention time and mean depth as explanatory variables. 4. Phytoplankton clearly responded to reduced nutrient loading, mainly reflecting declining TP concentrations. Declines in phytoplankton biomass were accompanied by shifts in community structure. In deep lakes, chrysophytes and dinophytes assumed greater importance at the expense of cyanobacteria. Diatoms, cryptophytes and chrysophytes became more dominant in shallow lakes, while no significant change was seen for cyanobacteria. 5. The observed declines in phytoplankton biomass and chl a may have been further augmented by enhanced zooplankton grazing, as indicated by increases in the zooplankton : phytoplankton biomass ratio and declines in the chl a : TP ratio at a summer mean TP concentration of <100–150 μg L^?1. This effect was strongest in shallow lakes. This implies potentially higher rates of zooplankton grazing and may be ascribed to the observed large changes in fish community structure and biomass with decreasing TP contribution. In 82% of the lakes for which data on fish are available, fish biomass declined with TP. The percentage of piscivores increased in 80% of those lakes and often a shift occurred towards dominance by fish species characteristic of less eutrophic waters. 6. Data on macrophytes were available only for a small subsample of lakes. In several of those lakes, abundance, coverage, plant volume inhabited or depth distribution of submerged macrophytes increased during oligotrophication, but in others no changes were observed despite greater water clarity. 7. Recovery of lakes after nutrient loading reduction may be confounded by concomitant environmental changes such as global warming. However, effects of global change are likely to run counter to reductions in nutrient loading rather than reinforcing re‐oligotrophication.     

The impact of small chironomid grazers on epiphytic algal abundance and dispersion

• 1 A combination of field and laboratory experiments was used to assess the impact of chironomid grazers on taxonomic composition, abundance and dispersion of epiphytic algal assemblages.
• 2 In the laboratory, Psectrodadius sp. reduced the biovolume of algal species preferred as food and increased the degree of clumping of non-preferred species. Thienemanniella cf. fusca had both positive and negative effects (depending on the algal species) on the biovolumes of algal species preferred as food and increased the degree of clumping of non-preferred species.
• 3 In field exclosures, no effect of removal of chironomid larvae from the grazer assemblage could be detected in autumn or winter experiments. A third, longer removal experiment, conducted in summer, resulted in increased biovolumes of edible Cosmarium spp. and Aphanocapsa spp., preferred foods of chironomid larvae. Biovolumes of Lyngbya sp., Eulbochaete spp. and Oedogortium spp., filamentous taxa used extensively in larval case construction, also increased. Chironomid larvae had no effect on total algal biovolume or biovolume of large unicellular algae.
• 4 Chironomid larvae can influence epiphytic algal assemblages through selective grazing by reducing the biovolumes of preferred foods and through case-building activity by reducing the biovolumes of construction materials.

     

Influence of bacterial density and water temperature on the grazing activity of two freshwater ciliates

1. The influences of bacterial density and water temperature on the grazing activity of the ciliates Uronema sp. and Colpoda inflata were studied. The conditions assayed were two prey densities (10⁶ and 4 × 10⁷ bacteria ml^?1) and three water temperatures (10, 15 and 22 °C). 2. The response of the ciliates was measured from changes in protistan biovolumes and specific clearance rates. At high prey density, both ciliates showed lower biovolumes as water temperature increased, while at low prey density this tendency was minimized. 3. At the intermediate temperature of 15 °C both ciliates filtered ten times more body volume when bacteria were scarce; however, the ingested bacteria were fewer than at high prey density. At low prey density, a decrease from 15 to 10 °C evidenced different strategies of the two ciliates, which led to a similar ingestion of bacteria: C. inflata reduced its specific clearance rates and increased its biovolume, while Uronema sp. did not show changes. At high prey density, an increase from 15 to 22 °C caused lower biovolumes and a noticeable increase in specific clearance rates in both ciliates, indicating opportunist behaviour.     

Quantitative responses of lake phytoplankton to eutrophication in Northern Europe

Based on the currently largest available dataset of phytoplankton in lakes in northern Europe, we quantified the responses of three major phytoplankton classes to eutrophication. Responses were quantified by modelling the proportional biovolumes of a given group along the eutrophication gradient, using generalized additive models. Chlorophyll-a (Chl-a) was chosen as a proxy for eutrophication because all classes showed more consistent responses to Chl-a than to total phosphorus. Chrysophytes often dominate in (ultra-) oligotrophic lakes, and showed a clear decrease along the eutrophication gradient. Pennate diatoms were found to be most abundant at moderate eutrophication level (spring-samples). Cyanobacteria often dominate under eutrophic conditions, especially in clearwater lakes at Chl-a levels >10 μg l⁻¹ (late summer samples). We compare the relationships among types of lakes, based on the lake typology of the northern geographic intercalibration group, and among countries sharing common lake types. Significant differences were found especially between humic and clearwater lakes, and between low- and moderately alkaline lakes, but we could not identify significant differences between shallow and deep lakes. Country-specific differences in response curves were especially pronounced between lakes in Norway and Finland, while Swedish lakes showed an intermediate pattern, indicating that country-specific differences reflect large-scale geographic and climatic differences in the study area.