The past and future of phytoplankton in the UK's largest lake,Lough Neagh

Lough Neagh is the largest lake in the UK and has been extensively monitored since 1974. It has suffered from considerable eutrophication and toxic algal blooms. The lake continues to endure many of the symptoms of nutrient enrichment despite improvements in nutrient management throughout the catchment, in particular a permanently dominant crop of the cyanobacterium Planktothrix agardhii. This study examines the historical changes in the Lough, and uses the PROTECH lake model to predict how the phytoplankton community may adapt in response to potential future changes in air temperature and nutrient load. PROTECH was calibrated against 2008 observations, with a restriction on the maximum simulated mixed depth to reflect the shallow nature of the lake and the addition of sediment released phosphorus throughout the mixed water column between 1 May and 1 October (with an equivalent in-lake concentration of 2.0 mg m⁻³). The historical analysis showed that phytoplankton biomass (total chlorophyll a) experienced a steady decline since the mid-1990s. During the same period the key nutrients for phytoplankton growth in the lake have shown contrasting trends, with increases in phosphorus concentrations and declines in nitrate concentrations. The modelled future scenarios which simulated a temperature increase of up to 3 °C showed a continuation of those trends, i.e. total chlorophyll a and nitrate concentrations declined in the surface water, while phosphorus concentrations increased and P. agardhii dominated. However, scenarios which simulated a 4 °C increase in air temperature showed a switch in dominance to the cyanobacteria, Dolichospermum spp. (formerly Anabaena spp.). This change was caused by a temperature related increase in growth driving nutrient consumption to a point where nitrate was limiting, allowing the nitrogen-fixing Dolichospermum spp. to gain sufficient advantage. These results suggest that in the long term, one nuisance cyanobacteria bloom may only be replaced by another unless the in-lake phosphorus concentration can be greatly reduced.     

Modelling the effects of changing retention time on abundance and composition of phytoplankton species in a small lake

1. The phytoplankton community model, PROTECH, was used to model the algal response to changing annual mean retention time in a small lake. 2. Simulations of short retention time with a fixed nutrient load resulted in a reduced chlorophyll concentration. A similar relationship was observed when the simulations were repeated but with inflowing nutrients increased in proportion to river discharge. 3. Longer retention time caused the spring bloom to start earlier and the autumn bloom to persist longer. 4. Changes in discharge of the inflowing river also caused a change in the thermal structure of the lake. This change in thermal structure, in turn, influenced the magnitude and composition of the phytoplankton population, particularly those in the CS‐functional group, such as Aphanizomenon.     

Invertebrate food sources for waterbirds provided by the reconstructed wetland of Nyirkai-Hany,northwestern Hungary

Detailed studies of the macroinvertebrate benthos and zooplankton communities in Loch Leven, the largest shallow lowland lake in Scotland, UK, were carried out from 1966 to 1973 as part of the International Biological Programme (IBP). The results revealed a reduction in species diversity that was attributed to increasing eutrophication. This work provides a baseline against which the response of the invertebrate communities to subsequent changes in management can be assessed. This article compares macroinvertebrate benthos and zooplankton data from the IBP study with the post-IBP era during which changes at Loch Leven included a 60% reduction in the phosphorus input from external sources and variations in fish stocking rates. Only in recent years has there been evidence of ecological recovery by the invertebrate communities: the number of macroinvertebrate and zooplankton taxa has increased (including taxa considered to be sensitive to nutrient enrichment) and invertebrate abundances have declined. These changes appear to reflect the improvements in water quality and habitat conditions at Loch Leven that have occurred as a result of the recent reduction in nutrient loads, albeit with a substantial delay before any ecological response could be detected. This time lag in recovery has important implications for assessing improvements in the ecological status of other lake systems, as is required by the EU Water Framework Directive.     

The seasonal sensitivity of Cyanobacteria and other phytoplankton to changes in flushing rate and water temperature

The phytoplankton lake community model PROTECH (Phytoplankton RespOnses To Environmental CHange) was applied to the eutrophic lake, Esthwaite Water (United Kingdom). It was validated against monitoring data from 2003 and simulated well the seasonal pattern of total chlorophyll, diatom chlorophyll and Cyanobacteria chlorophyll with respective R²‐values calculated between observed and simulated of 0.68, 0.72 and 0.77 (all P<0.01). This simulation was then rerun through various combinations of factorized changes covering a range of half to double the flushing rate and from ?1 to +4 °C changes in water temperature. Their effect on the phytoplankton was measured as annual, spring, summer and autumn means of the total and species chlorophyll concentrations. In addition, Cyanobacteria mean percentage abundance (%Cb) and maximum percentage abundance (Max %Cb) was recorded, as were the number of days that Cyanobacteria chlorophyll concentration exceed two World Health Organization (WHO) derived risk thresholds (10 and 50 mg m^?3). The phytoplankton community was dominated in the year by three of the eight phytoplankton simulated. The vernal bloom of the diatom Asterionella showed little annual or seasonal response to the changing drivers but this was not the case for the two Cyanobacteria that also dominated, Anabaena and Aphanizomenon . These Cyanobacteria showed enhanced abundance, community dominance and increased duration above the highest WHO risk threshold with increasing water temperature and decreasing flushing rate: this effect was greatest in the summer period. However, the response was ultimately controlled by the availability of nutrients, particularly phosphorus and nitrogen, with occasional declines in the latter's concentration helping the dominance of these nitrogen‐fixing phytoplankton.     

Modelling the effects on phytoplankton communities of changing mixed depth and background extinction coefficient on three contrasting lakes in the English Lake District

1. The process‐based phytoplankton community model, PROTECH, was used to model the response of algal biomass to a range of mixed layer depths and extinction coefficients for three contrasting lakes: Blelham Tarn (eutrophic), Bassenthwaite Lake (mesotrophic) and Ullswater (oligotrophic). 2. As expected, in most cases biomass and diversity decreased with decreasing light availability caused by increasing the mixed depth and background extinction coefficient. The communities were generally dominated by phytoplankton tolerant of low light. Further, more novel, factors were identified, however. 3. In Blelham Tarn in the second half of the year, biomass and diversity did not generally decline with deeper mixing and the community was dominated by nitrogen‐fixing phytoplankton because that nutrient was limiting to growth. 4. In Bassenthwaite Lake, changing mixed depth influenced the retention time so that, as the mixed depth declined, the flushing rate in the mixed layer increased to the point that only fast‐growing phytoplankton could dominate. 5. In the oligotrophic Ullswater, changing the mixed depth had a greater effect through nutrient supply rather than light availability. This effect was observed when the mixed layer was relatively shallow (<5.5 m) and the driver for this was that the inflowing nutrients were added to a smaller volume of water, thus increasing nutrient concentrations and algal growth. 6. Therefore, whilst changes in mixed depth generally affect the phytoplankton via commonly recognized factors (light availability, sedimentation rate), it also affected phytoplankton growth and community composition through other important factors such as retention time and nutrient supply.     

The impacts of changing nutrient load and climate on a deep,eutrophic, monomictic lake

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Changes in shallow lake functioning: response to climate change and nutrient reduction

Hydrobiologia - Nutrient, phytoplankton and zooplankton dynamics have been monitored intensively at Loch Leven for 34 years. The data collected reveal a decline in phosphorus concentrations,...     

The importance of nutrient source in determining the influence of retention time on phytoplankton: an explorative modelling study of a naturally well-flushed lake

Two models were used to examine the relationship between hydraulic retention time, nutrient source and total chlorophyll in a shallow lake (Bassenthwaite Lake, UK). The first model was a derivation of the Vollenweider model and the second was the phytoplankton community model, PROTECH. The adapted Vollenweider model produced two different responses to changing the retention time that were phosphorus source dependent. If the phosphorus was totally from a point source, then annual mean chlorophyll steadily declined with increasing flushing rate. However, when a diffuse source was used, the chlorophyll changed little and even increased with short retention times (retention time <40 days). The PROTECH model produced some similar responses but they were more season dependent. Winter mean chlorophyll always declined with decreasing retention time, regardless of nutrient source, but the summer mean curves were source dependent and similar to those produced by the adapted Vollenweider model. Further simulations with PROTECH using a standardized flow regime provided strong evidence as to the mechanisms behind these responses. Analysis showed that the decline in chlorophyll with decreasing retention time was the prevalent response of the PROTECH simulations due to flushing loss of both nutrients and algae. Furthermore, the curve formed an asymptote at long retention times because other factors (e.g. light) limited growth; retention times >100 days had little effect on chlorophyll. However, with a diffuse phosphorus source and short retention times, an increase in biomass was observed when the nutrient was limiting for growth. Handling editor: Luigi Naselli-Flores     

Identifying from recent sediment records the effects of nutrients and climate on diatom dynamics in Loch Leven

1. Changes in nutrients and climate have occurred over approximately the same timescales in many European lake catchments. Here, we attempt to interpret the sedimentary diatom record of a large shallow lake, Loch Leven, in relation to these pressures using information gained from analysis of long‐term data sets of water quality, climate and planktonic diatoms. 2. The core data indicate the enrichment of Loch Leven starting in c. 1800–1850, most likely from agricultural practices in the catchment, with a more marked phase since c. 1940–1950 caused by increased phosphorus inputs from sewage treatment works, land drainage and a woollen mill. 3. While the recent diatom plankton remains are dominated by taxa associated with nutrient‐rich conditions, an increase in Aulacoseira subarctica relative to Stephanodiscus taxa since the mid‐1980s suggests that reductions in external catchment sources of nutrients (since 1985) may have resulted in partial recovery. This observation accords well with the long‐term monitoring series of water chemistry and phytoplankton. 4. On a decadal‐centennial scale, the eutrophication signal in the sediment record outweighs any evidence of climate as a control on the diatom community. However, at an inter‐annual scale, while the diatom data exhibit high variability, there are several changes in species composition in the recent fossil record that may be attributed to climatic controls. 5. The study highlights the value of a palaeolimnological approach, particularly when coupled with long‐term data sets, for developing our understanding of environmental change at a range of temporal scales. The diatom record in the sediment can be used effectively to track recovery from eutrophication, but requires greater understanding of contemporary ecology to fully interpret climate impacts. 6. The study illustrates the complexity of ecosystem response to synchronous changes in nutrients and climate, and the difficulty of disentangling the effects of these multiple, interacting pressures.     

The effect of increased nitrogen load on phytoplankton in a phosphorus‐limited lake

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A seasonal and ecological study of the phytoplankton of Loch Lomond

The seasonal succession of phytoplankton in Loch Lomond has been studied both quantitatively and from observations on net plankton. From a qualitative investigation of net phytoplankton Loch Lomond may be described as diatom-desmid in nature although from the quantitative studies the general domination of the population by diatoms becomes clear. During the vernal phytoplankton increase diatoms contributed more than 90% of the population throughout the loch. The bulk of the phytoplankton production occurred in the southern region, where the standing crop was considerably greater throughout the year than in other regions of the loch.Patterns of seasonal succession are described for all species which make an important contribution to the productivity of the loch and the observed successions related to measured physical and chemical changes in the water. Explanations are put forward to account for the absence of a second (autumnal) pulse ofMelosira in Loch Lomond. The appearance ofAnabaena circinalis is a possible indication of the changing trophic status of the southern region of the loch.     

Increases in lake phytoplankton biomass caused by future climate‐driven changes to seasonal river flow

For the many lakes world‐wide with short residence times, changes to the rate of water throughput may have important effects on lake ecology. We studied relationships between current and predicted residence times and phytoplankton biomass using a eutrophic lake in the north‐west of England with an annual residence time averaging about 20 days, as a test case. Using 32 years of recent hydrological flow data for Bassenthwaite Lake, multiple sets of scaled flow for each year, and the process‐based phytoplankton response model, PROTECH, we modelled the effects of changing river flow on phytoplankton biomass in the lake. The impact on biomass was shown to depend on seasonal changes in flow rather than annual changes. Furthermore, there was a qualitative difference in impact depending on whether the nutrient loading to the lake came principally from flow‐independent sources, or from flow‐dependent ones. Predictions for changes in river flow under future climate scenarios in the north‐west of England have suggested that, despite little change in the annual flow magnitude, there will be a shift to greater flow in the winter and lesser flow in the summer. Applying these flow predictions to our modelling of Bassenthwaite Lake revealed that, with flow‐independent nutrient loading, and no overall increase in nutrient load, phytoplankton abundance in the summer could increase by up to 70%, including an increased proportion of Cyanobacteria. Conversely, were the loading completely dependent on the flow, the biomass would fall. In many parts of the world, river flow is expected to decrease in the summer even more than in England, suggesting these areas may expect substantial changes to seasonal phytoplankton biomass as a result of climate‐driven changes to seasonal river flow. Such changes would be in addition to any other changes owing to warming effects or eutrophication.     

Strong spatial differentiation of N and P deficiency,primary productivity and community composition between Nyanza Gulf and Lake Victoria (Kenya,East Africa) and the implications for nutrient management

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Nutrient limitation of bacterioplankton and phytoplankton in humic lakes in northern Sweden

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L^–1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), respectively. A third lake was unfertilized and served as a control. In addition to this lake fertilization experiment, data from different regional surveys were used to assess the role of different limiting factors.
2. The P fertilization had no effects on bacterioplankton or phytoplankton, while phytoplankton were significantly stimulated by N fertilization. Inorganic nutrient limitation of bacterioplankton was a function of DOC concentration in water of the investigated region and nutrient-limited bacteria were found only in lakes with DOC concentrations less than around 15 mg L^–1
3. The fertilization experiments demonstrated that the DOC-rich experimental lakes contained a bioavailable pool of P that was not utilized to its full potential under natural conditions. The overall mobilization of energy (bacterioplankton plus phytoplankton) in the experimental lakes was restricted by lack of inorganic N.     

Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.     

Growth response of four freshwater algal species to dissolved organic nitrogen of different concentration and complexity

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Long-term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Müggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Müggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N₂‐fixing species Aphanizomenon flos‐aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re‐colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Müggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.     

Changes in a deep lake following sewage diversion – a challenge to the orthodoxy of external phosphorus control as a restoration strategy?

1. The restoration of deep lakes has traditionally focused on reducing the external phosphorus loading. 2. Following the diversion of sewage effluent, that led to marked reductions in nutrient concentrations in its main inflow, Rostherne Mere has shown no reduction in phosphorus or chlorophyll a concentrations. A shallow lake upstream (Little Mere), however, has shown a marked response to effluent diversion. 3. Nutrient budgets for Rostherne Mere reveal that sewage effluent was by far the most significant external source of total phosphorus and that diffuse drainage from the catchment was the most significant external source of dissolved inorganic nitrogen. Phosphorus loads from groundwater and a bird roost were insignificant. Internal sources of phosphorus were, however, considerable and were largely responsible for the observed delay in recovery. 4. Phosphorus limitation of phytoplankton biomass may never be attainable because of substantial internal and diffuse sources of phosphorus, combined with a long retention time. Nitrogen is likely to be more important in limiting phytoplankton biomass. Control of diffuse nitrogen sources may therefore be more effective in the restoration of the deeper lakes of this region.     

Testing the Sensitivity of Phytoplankton Communities to Changes in Water Temperature and Nutrient Load, in a Temperate Lake

Freshwater lakes are biologically sensitive to changes in the surrounding environment and the impacts that such changes have on their water quality are of considerable ecological, recreational and economic importance. In this study the phytoplankton community model, PROTECH, was used to experiment with the effects of elevated temperatures and increased nutrient load on phytoplankton succession and productivity. The response of a phytoplankton community to combined incremental changes in these drivers was analysed, in order to elucidate the resulting ecological changes. Annual mean phytoplankton biomass increased with increases in temperature and nutrient loading, although the latter had the larger effect. The phenology of the dominant phytoplankton taxa changed with increasing water temperature; the three spring blooming species all peaked earlier in the year. The simulated summer bloom of Anabaena became earlier in the year and the Chlorella bloom later. The increased phytoplankton biomass was largely dominated by the cyanobacterium Anabaena, which was especially prevalent during the summer bloom. This resulted in a progressive loss of phytoplankton biodiversity with increasing water temperature and nutrient supply. Model experimentation showed that whilst both factors greatly affected the community, the changes to nutrient loading generally had the greater effect and that at low nutrient levels the effect of water temperature change was reduced considerably. Finally, the model predicted that cyanobacteria have the potential to dominate the phytoplankton community, with clear consequences for water quality, and that this dominance was at its greatest when high water temperatures were combined with high nutrient loads.     

Hypolimnetic phosphorus retrieval by diel vertical migrations of lake phytoplankton

SUMMARY. I. Movement of ³³P from hypolimnion to epilimnion in a small, dystrophic lake was investigated using small-diameter experimental tubes enclosing thermally stratified water columns. This approach was made possible by the extremely sharp stratification found in such lakes, in which the euphotic zone closely coincides with the epilimnion.
2. The vertical distribution of inorganic phosphorus in the lake showed a sharp increase across the thermocline so that enhanced concentrations were available to phytoplankton just below the thermocline. Inorganic nitrogen concentrations did not show such a marked relation to thermal stratification.
3. One abundant motile alga ( Cryptomonas marssonii ) showed striking and regular vertical migrations in the lake, moving below the thermocline at night and returning to the surface waters in early morning. These migrations took cells across a 10°C temperature gradient. Non-motile phytoplankton showed constant vertical distributions.
4. In the experimental tubes an upward movement of phosphorus took place from hypolimnion to epilimnion which was only attributable to transport by phytoplankton cells undertaking active vertical migrations. No equivalent movement of phosphorus occurred in control tubes from which algae were absent.
5. The possible significance of such nutrient retrieval is discussed with reference to plankton phosphorus budgets and competition between phytoplankton species.