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Climate change and eutrophication are among the main stressors of shallow freshwater ecosystems, and their effects on phytoplankton community structure and primary production have been studied extensively. However, their combined effects on the algal production of polyunsaturated fatty acids (PUFA), specifically, eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) are currently unresolved. Moreover, the proximate reasons for changes in phytoplankton EPA and DHA concentrations are unclear, i.e., the relative importance of ecological (changes in the community composition) vs. ecophysiological (within taxa changes in EPA and DHA levels) factors. We investigated the responses of phytoplankton EPA and DHA concentrations to warming (IPCC climate scenario) and nutrient additions in mesocosms which had been run continuously at varying temperature and nutrient levels for 15 years prior to this study. Nutrient treatment had a significant effect on phytoplankton EPA and DHA concentrations and about 59 % of the variation in EPA and DHA concentrations could be explained by changes in the phytoplankton community structure. Increased biomass of diatoms corresponded with high EPA and DHA concentrations, while cyanobacteria/chlorophyte dominated mesocosm had low EPA and DHA concentrations. Warming had only a marginal effect on the EPA and DHA concentrations in these mesocosms. However, a significant interaction was observed with warming and N:P ratio. Our findings indicate that direct nutrient/temperature effects on algal physiology and PUFA metabolism were negligible and the changes in EPA and DHA concentrations were mostly related to the phytoplankton community structure and biomass. These results also imply that in shallow temperate lakes eutrophication, leading to increased dominance of cyanobacteria, will probably be a greater threat to phytoplankton EPA and DHA production than warming. EPA and DHA are nutritionally important for upper trophic level consumers and decreased production may impair secondary production., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

We analysed changes in the abundance, biomass and cell size of the microbial food web community (bacteria, heterotrophic nanoflagellates, ciliates) at contrasting nutrient concentrations and temperatures during a simulated heat wave. We used 24 mesocosms mimicking shallow lakes in which two nutrient levels (unenriched and enriched by adding nitrogen and phosphorus) and three different temperature scenarios (ambient, IPCC A2 scenario and A2+%50) are simulated (4 replicates of each). Experiments using the mesocosms have been running un-interrupted since 2003. A 1-month heat wave was imitated by an extra 5 °C increase in the previously heated mesocosms (from 1st July to 1st August 2014). Changes in water temperature induced within a few days a strong effect on the microbial food web functioning, demonstrating a quick response of microbial communities to the changes in environment, due to their short generation times. Warming and nutrients showed synergistic effects. Microbial assemblages of heterotrophic nanoflagellates and ciliates responded positively to the heating, the increase being largest in the enriched mesocosms. The results indicate that warming and nutrients in combination can set off complex interactions in the microbial food web functioning., (Copyright © 2018 Elsevier GmbH. All rights reserved.)

Aquatic ecosystems are threatened by eutrophication from nutrient pollution. In lakes, eutrophication causes a plethora of deleterious effects, such as harmful algal blooms, fish kills and increased methane emissions. However, lake-specific responses to nutrient changes are highly variable, complicating eutrophication management. These lake-specific responses could result from short-term stochastic drivers overshadowing lake-independent, long-term relationships between phytoplankton and nutrients. Here, we show that strong stoichiometric long-term relationships exist between nutrients and chlorophyll a (Chla) for 5-year simple moving averages (SMA, median R² = 0.87) along a gradient of total nitrogen to total phosphorus (TN:TP) ratios. These stoichiometric relationships are consistent across 159 shallow lakes (defined as average depth < 6 m) from a cross-continental, open-access database. We calculate 5-year SMA residuals to assess short-term variability and find substantial short-term Chla variation which is weakly related to nutrient concentrations (median R² = 0.12). With shallow lakes representing 89% of the world's lakes, the identified stoichiometric long-term relationships can globally improve quantitative nutrient management in both lakes and their catchments through a nutrient-ratio-based strategy. Nutrient limitation is a well-known control of phytoplankton growth, but predicting specific responses in individual lakes based on nutrient data alone has proven challenging. Here, the authors show that long-term signals of chlorophyll-a dynamics in shallow lakes can be captured based on stoichiometric effects of N and P concentrations along a continuum of total N:total P ratios. [ABSTRACT FROM AUTHOR]

The climate-driven changes in temperature, in combination with high inputs of nutrients through anthropogenic activities, significantly affect phytoplankton communities in shallow lakes. This study aimed to assess the effect of nutrients on the community composition, size distribution, and diversity of phytoplankton at three contrasting temperature regimes in phosphorus (P)–enriched mesocosms and with different nitrogen (N) availability imitating eutrophic environments. We applied imaging flow cytometry (IFC) to evaluate complex phytoplankton communities changes, particularly size of planktonic cells, biomass, and phytoplankton composition. We found that N enrichment led to the shift in the dominance from the bloom-forming cyanobacteria to the mixed-type blooming by cyanobacteria and green algae. Moreover, the N enrichment stimulated phytoplankton size increase in the high-temperature regime and led to phytoplankton size decrease in lower temperatures. A combination of high temperature and N enrichment resulted in the lowest phytoplankton diversity. Together these findings demonstrate that the net effect of N and P pollution on phytoplankton communities depends on the temperature conditions. These implications are important for forecasting future climate change impacts on the world's shallow lake ecosystems. [ABSTRACT FROM AUTHOR]

A machine learning approach was employed to detect and quantify Microcystis colonial morphospecies using FlowCAM-based imaging flow cytometry. The system was trained and tested using samples from a long-term mesocosm experiment (LMWE, Central Jutland, Denmark). The statistical validation of the classification approaches was performed using Hellinger distances, Bray–Curtis dissimilarity, and Kullback–Leibler divergence. The semi-automatic classification based on well-balanced training sets from Microcystis seasonal bloom provided a high level of intergeneric accuracy (96–100%) but relatively low intrageneric accuracy (67–78%). Our results provide a proof-of-concept of how machine learning approaches can be applied to analyze the colonial microalgae. This approach allowed to evaluate Microcystis seasonal bloom in individual mesocosms with high level of temporal and spatial resolution. The observation that some Microcystis morphotypes completely disappeared and re-appeared along the mesocosm experiment timeline supports the hypothesis of the main transition pathways of colonial Microcystis morphoforms. We demonstrated that significant changes in the training sets with colonial images required for accurate classification of Microcystis spp. from time points differed by only two weeks due to Microcystis high phenotypic heterogeneity during the bloom. We conclude that automatic methods not only allow a performance level of human taxonomist, and thus be a valuable time-saving tool in the routine-like identification of colonial phytoplankton taxa, but also can be applied to increase temporal and spatial resolution of the study. [ABSTRACT FROM AUTHOR]

Shallow lakes are a key component of the global carbon cycle. It is, therefore, important to know how shallow lake ecosystems will respond to the current climate change. Global warming affects not only average temperatures, but also the frequency of heat waves (HW). The impact of extreme events on ecosystems processes, particularly greenhouse gas ( GHG) emissions, is uncertain., Using the world's longest-running shallow lake experiment, we studied the effects of a simulated summer HW on the fluxes of carbon dioxide ( CO2), methane ( CH4) and nitrous oxide (N2O). The experimental mesocosms had been exposed to different temperature treatments and nutrient loading for 11 years prior to the artificial HW., In general, there was an increase in total GHG emissions during the 1-month artificial HW, with a significant increase in CO2, CH4 and N2O being observed in the shallow lake mesocosms. No significant effect of the HW on CO2 emissions could be traced, though, in the mesocosms with high nutrient levels. Furthermore, the data suggested that in addition to the direct effect of increased temperature on metabolic processes during the HW, biotic interactions exerted a significant control of GHG emissions. For example, at low nutrient levels, increased CO2 emissions were associated with low macrophyte abundance, whereas at high nutrient levels, decreased phytoplankton abundance was linked to increased emissions of CO2 and CH4., In contrast to the observable heat-wave effect, no clear general effect of the long-term temperature treatments could be discerned over the summer, likely because the potential effects of the moderate temperature increase, applied as a press disturbance, were overridden by biotic interactions. This study demonstrates that the role of biotic interactions needs to be considered within the context of global warming on ecosystem processes. [ABSTRACT FROM AUTHOR]

A two-decade (1989-2008) time series of lake phyto-and zooplankton, water characteristics and climate in 17 Danish lakes was analysed to examine the long term changes and the effects of lake restoration efforts. The analyses of the pair-wise correlations across time series revealed a strong synchrony in climatic variables among the lakes. A significant, but weak increase in air temperature was observed and resulted in a corresponding increase in surface water temperature only in summer. Lake physico-chemical variables had weaker synchrony than climatic variables. Synchrony in water temperature and stratification was stronger than lake chemistry as the former is mostly affected by atmospheric energy flux. Synchrony in the taxonomic richness of the plankton groups and phytoplankton biomass was apparent, to a similar degree as observed for lake chemistry. The synchrony and the temporal trends in lake chemistry and plankton were more pronounced for the lakes with strong re-oligotrophication. Phytoplankton biomass decreased and plankton richness increased in these lakes, with a shift from Chlorophyta dominance towards more heterogeneous phytoplankton communities. Notably, a widespread significant positive trend in plankton richness was observed not only in lakes with strong re-oligotrophication but across all lakes. The widespread increase in plankton richness coincided with widespread decrease in phosphate and total nitrogen concentrations, as well as with the trends in climate indicating a likely joint effect of nutrient reduction and climate in driving lake plankton. However, temporal changes and synchrony as well as the recovery of richness and composition of lake plankton more coherently corresponded with the nutrient loading reduction across the Danish landscape, while the role of climate control of the lake plankton was less pronounced. [ABSTRACT FROM AUTHOR]

Lakes can be sources or sinks of carbon, depending on local conditions. Recent studies have shown that the CO efflux increases when lakes recover from eutrophication, mainly as a result of a reduction in phytoplankton biomass, leading to less uptake of CO by producers. We hypothesised that lake restoration by removal of coarse fish (biomanipulation) or invasion of mussels would have a similar effect. We studied 14-22 year time series of five temperate Danish lakes and found profound effects on the calculated CO efflux of major shifts in ecosystem structure. In two lakes, where limited colonisation of submerged macrophytes occurred after biomanipulation or invasion of zebra mussels ( Dreissena polymorpha), the efflux increased significantly with decreasing phytoplankton chlorophyll a. In three lakes with major interannual variation in macrophyte abundance, the efflux declined with increasing macrophyte abundance in two of the lakes, while no relation to macrophytes or chlorophyll a was found in the third lake, likely due to high groundwater input to this lake. We conclude that clearing water through invasive mussels or lake restoration by biomanipulation may increase the CO efflux from lakes. However, if submerged macrophytes establish and form dense beds, the CO efflux may decline again. [ABSTRACT FROM AUTHOR]

Multi-indicator palaeoecological studies have become increasingly popular over the last decade as the need for a more complete understanding of lake ecological histories has increased. However, the true potential of the full biological record for assessing the potential drivers of observed ecological shifts in lake sediment records has rarely been demonstrated., Here, we examine the remains of a range of food-web components including algae (diatoms), macrophytes (plant macrofossils), zooplankton (chitinous and ephippial Cladocera remains), invertebrates (including chironomids, bryozoans, Mollusca) and fish (fish scales and fish leech egg cocoons) in multiple sediment cores from Groby Pool, an enriched English shallow lake, to assess whole-ecosystem response to eutrophication over the last two centuries. We focus on three striking changes in the palaeorecord, namely the post-1900 increase in Daphnia spp., the post-1840 decline in Cristatella mucedo and the post-1940 increase in Cocconeis placentula, and utilise the multi-indicator palaeoecological data to evaluate possible explanations for these patterns., Principal curves analysis revealed marked and broadly simultaneous changes in the plant macrofossils, cladocerans, diatoms and chironomids (as well as in other animal remains such as bryozoans and Mollusca), indicating an early period of enrichment most likely associated with land-use change in the late 18th century, followed by a more recent eutrophication phase coincident with the discharge of sewage effluent to the lake from 1935., Ecological change, resulting from eutrophication, was shown to have progressed slowly and steadily and to have occurred at all trophic levels with a shift from a relatively diverse 'mesotrophic' macrophyte assemblage, dominance by benthic diatoms and plant-associated Chydoridae and chironomids towards a relatively species-poor, 'eutrophic' macrophyte community with dominance by planktonic algae (e.g. Cyclostephanoid diatom taxa), planktonic Cladocera ( Bosmina, Daphnia) and a chironomid fauna dominated by mud-associated taxa. The inferred shift in the macrophyte community from charophyte to fine-leaved pondweed and Callitriche truncata suggests a reduction in the seasonal duration of plant dominance., The multi-indicator analysis indicates that a combination of increased phytoplankton biomass and low zooplanktivorous fish predation is likely to explain the recent increases in Daphnia spp., while loss of plant habitat and increased competition for food appear to be the most likely causes of the observed decline in C. mucedo, and resistance to increased grazing pressure from invertebrates is the most probable driver of the C. placentula increase., Our study illustrates the potential of using the full array of fossil groups preserved in lake sediment cores to infer ecosystem dynamics over centennial timescales and to contribute to our understanding of the mechanisms that may link biological assemblages to a range of forcing factors. Further, this paper provides methodological guidance by demonstrating the ability of amalgamated records from three cores to reveal a strong sequence of events and coherent patterns. [ABSTRACT FROM AUTHOR]

Ponds have significant conservation value due to their potentially high contribution to local and regional diversity. However, most ponds are located in anthropogenically influenced areas, and their biodiversity is constantly threatened by human activities. Thus, knowledge of the effect of pond management on biodiversity is essential for designing effective conservation strategies., Here, we study the main drivers of diversity of three functional groups of primary producers (phytoplankton, filamentous green algae and submerged macrophytes) in 87 ponds distributed across a large region (c. 90000 km2) in Southern Spain. We hypothesised that spatial effects would increase with increasing propagule size. However, given the regional extent of the study, we anticipated that both spatial and environmental controls would be significant., We determined α -, β - and γ-components of biodiversity for each functional group and pond type (embankment ponds, excavated ponds and artificial ponds) and assessed the influence of environmental and spatial drivers on diversity with generalised additive models ( GAMs). Redundancy analyses ( RDAs) with variation partitioning were used to determine the relative contribution of environmental and spatial predictors of the community assembly. Spatial variables were calculated by applying distance-based Moran's eigenvector maps (db- MEM)., Both α - and β -diversities of phytoplankton and filamentous green algae varied significantly with pond type. Generally, environmental predictors of diversity were more important than spatial variables. Assemblage structure was controlled by water chemistry and eutrophication, with a marked influence of pond type. Spatial variables included broad-scale variation for the three groups of primary producers, which were also strongly influenced by the management regime., Limited management activities, as occurs at embankment ponds, promoted the local richness of phytoplankton and contributed importantly to the regional diversity of macrophytes. Moreover, the relative contribution of environmental and spatial variables was similar between embankment ponds and natural ecosystems, that is, dispersal limitation increasing with propagule size. Excavated and artificial ponds also contributed importantly to regional diversity by enhancing phytoplankton and filamentous green algal β-diversity. However, spatial patterns in the latter pond type did not meet our expectations, most likely due to the intensive disturbance from pond management hampering the development of macrophytes. [ABSTRACT FROM AUTHOR]

1. Seasonal relationships between macrophyte and phytoplankton populations may alter considerably as lakes undergo eutrophication. Understanding of these changes may be key to the interpretation of ecological processes operating over longer (decadal-centennial) timescales. 2. We explore the seasonal dynamics of macrophytes (measured twice in June and August) and phytoplankton (measured monthly May–September) populations in 39 shallow lakes (29 in the U.K. and 10 in Denmark) covering broad gradients for nutrients and plant abundance. 3. Three site groups were identified based on macrophyte seasonality; 16 lakes where macrophyte abundance was perennially low and the water generally turbid (‘turbid lakes’); 7 where macrophyte abundance was high in June but low in August (‘crashing’ lakes); and 12 where macrophyte abundance was high in both June and August (‘stable’ lakes). The seasonal behaviour of the crashing and turbid lakes was extremely similar with a consistent increase in nutrient concentrations and chlorophyll- a over May–September. By contrast in the stable lakes, seasonal changes were dampened with chlorophyll- a consistently low (<10–15 μg L−1) over the entire summer. The crashing lakes were dominated by one or a combination of Potamogeton pusillus, Potamogeton pectinatus and Zannichellia palustris, whereas Ceratophyllum demersum and Chara spp. were more abundant in the stable lakes. 4. A long-term loss of macrophyte species diversity has occurred in many shallow lakes affected by eutrophication. One common pathway is from a species-rich plant community with charophytes to a species-poor community dominated by P. pusillus, P. pectinatus and Z. palustris. Such compositional changes may often be accompanied by a substantial reduction in the seasonal duration of plant dominance and a greater tendency for incursions by phytoplankton. We hypothesise a slow-enacting (10–100 s years) feedback loop in nutrient-enriched shallow lakes whereby increases in algal abundance are associated with losses of macrophyte species and hence different plant seasonal strategies. In turn such changes may favour increased phytoplankton production thus placing further pressure on remaining macrophytes. This study blurs the distinction between so-called turbid phytoplankton-dominated and clear plant-dominated shallow lakes and suggests that plant loss from them may be a gradual process. [ABSTRACT FROM AUTHOR]

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long-term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple ( n = 5) chronologically correlated sediment cores for plant macro-remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton ( c. pre-1900), to Ceratophyllum–Chara–Potamogeton ( c. 1900–1960) and finally to Zannichellia–Potamogeton ( c. post-1960). Macrophyte species richness declined from at least 10 species pre-1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi-benthic and phytoplankton-based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant-covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication-induced reductions in macrophyte species richness, especially if the number of plant-seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self-perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss. [ABSTRACT FROM AUTHOR]

Phytoplankton usually responds directly and fast to environmental fluctuations, making them useful indicators of lake ecosystem changes caused by various stressors. Here, we examined the phytoplankton community composition before, during, and after a simulated 1-month heat wave in a mesocosm facility in Silkeborg, Denmark. The experiment was conducted over three contrasting temperature scenarios (ambient (A0), Intergovernmental Panel on Climate Change A2 scenario (circa +3 °C, A2) and A2+ %50 (circa +4.5 °C, A2+)) crossed with two nutrient levels (low (LN) and high (HN)) with four replicates. The facility includes 24 mesocosms mimicking shallow lakes, which at the time of our experiment had run without interruption for 11 years. The 1-month heat wave effect was simulated by increasing the temperature by 5 °C (1 July to 1 August) in A2 and A2+, while A0 was not additionally heated. Throughout the study, HN treatments were mostly dominated by Cyanobacteria, whereas LN treatments were richer in genera and mostly dominated by Chlorophyta. Linear mixed model analyses revealed that high nutrient conditions were the most important structuring factor, which, regardless of temperature treatments and heat waves, increased total phytoplankton, Chlorophyta, Bacillariophyta, and Cyanobacteria biomasses and decreased genus richness and the grazing pressure of zooplankton. The effect of temperature was, however, modest. The effect of warming on the phytoplankton community was not significant before the heat wave, yet during the heat wave it became significant, especially in LN-A2+, and negative interaction effects between nutrient and A2+ warming were recorded. These warming effects continued after the heat wave, as also evidenced by Co-inertia analyses. In contrast to the prevailing theory stating that more diverse ecosystems would be more stable, HN were less affected by the heat wave disturbance, most likely because the dominant phytoplankton group cyanobacteria is adapted to high nutrient conditions and also benefits from increased temperature. We did not find any significant change in phytoplankton size diversity, but size evenness decreased in HN as a result of an increase in the smallest and largest size classes simultaneously. We conclude that the phytoplankton community was most strongly affected by the nutrient level, but less sensitive to changes in both temperature treatments and the heat wave simulation in these systems, which have been adapted for a long time to different temperatures. Moreover, the temperature and heat wave effects were observed mostly in LN systems, indicating that the sensitivity of phytoplankton community structure to high temperatures is dependent on nutrient availability. [ABSTRACT FROM AUTHOR]

[Display omitted] • Microcystis morphospecies respond to pH, temperature, and nutrients differently. • NCSC positively correlate with M. ichtyoblabe , M. aeruginosa , and M. smithii. • Colonial sheath abundances positively correlate with M.wesenbergii abundances. • Colonial sheaths with remaining cells suited for Microcystis dispersal. • Imaging flow cytometry are available for early indication of Microcystis detection and dispersal. The alarming increase in the frequency of blooms of Microcystis in freshwater lakes and reservoirs occurs worldwide, with major implications for their ecosystem functioning and water quality. The dominance of Microcystis is tightly related to colonial formation by Microcystis. However, studies of the colonial development of Microcystis morphospecies are rare. This research applied FlowCAM-based imaging flow cytometry to analyze the development of Microcystis morphospecies in the mesocosms mimicking eutrophic shallow lakes and the effect of temperature changes. A significant positive association was found between M. ichtyoblabe , M. aeruginosa , and M. smithii colonies, particularly in the high-temperature tanks, suggesting that these morphospecies belong to one ecocluster, which supports the hypothesis of the central transition pathways of colonial Microcystis. The small colonial clusters of Microcystis cells represented an important stage in the sequence of Microcystis bloom and were associated with the development of colonial forms. A correlation analysis showed that the higher pH was positively correlated with the abundance of M.wesenbergii independently of temperature changes. The colonial sheaths' abundances increased following a maximum of M.wesenbergii abundance, reaching significant numbers (thousands), and a majority of sheaths contained at least some Microcystis cells. We hypothesize that colonial sheaths may be crucial at Microcystis spp. dispersal and represent an obligatory stage of colonies development. The sheaths may protect Microcystis cells against environmental stress factors, improve cell survival at low nutrient levels, and participate in Microcystis dispersal and spreading. Our findings can be applicable to early CyanoHAB detection and management of Microcystis dispersal. [ABSTRACT FROM AUTHOR]

Abstract: Eutrophication resulting from high nutrient loading has been the paramount environmental problem for lakes world-wide for the past four decades. Efforts are being made in many parts of the world to reduce external nutrient loading via improved wastewater treatment or diversion of nutrient-rich inflows. However, even after a reduction of the external phosphorus loading, the effects obtained may be unsatisfactory. This may reflect an insufficient reduction in the external nutrient loading to effectively limit phytoplankton growth. However, the lack of success may also be due to chemical or biological within-lake inertia preventing or delaying improvements. To overcome the resilience and thereby reinforce recovery, a number of physico-chemical and biological restoration methods have been developed. In this chapter, we describe recent developments of biological restoration methods related to eutrophication, their short-term and long-term effects, and discuss the possibility of using combined physico-chemical and biological methods to improve the long-term stability of restoration and to reduce restoration costs. As comprehensive reviews of the effect of fish manipulation in cold temperate lakes are numerous, for these waterbodies, we highlight recent results, including effects on biodiversity and metabolism, and present new approaches of biomanipulation. Our particular focus is, however, directed at biomanipulation in warm lakes and on combined treatments which are far less well described in the literature. [Copyright &y& Elsevier]