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Sediment phosphorus (P) recycling is one of the key issues in lake water quality management. We studied sediment P mobility in Võrtsjärv, a large shallow lake in Estonia using both sorption experiments and long-term (1985-2020) monitoring data of the lake. Over the years studied, the lake has undergone a decline in external phosphorus loading (EL), while no improvement in phytoplankton indicators was observed. The results of the sorption experiments revealed that it may be successfully used as a tool to determine P forms involved in P retention, as up to 100% of the P from the water column was detected in sediments. Incubation of wet sediment is preferred to dry because of the sensitivity of organic P to desiccation. In the sediments of Võrtsjärv, the labile P (Lab-P) and iron bound (Fe-P) fractions are the major forms of the mobile pool that supply internal P load as sediment released P. The internal P load calculated from summer total P (TP) increases (IL in situ ) in the water column was on average 42%, but could reach 240% of EL at extreme environmental conditions. IL in situ was correlated with the active area, which resembles the area involved in redox-related P release in polymictic lakes, and with the mean bottom shear stress in summer. IL in situ showed a similar decreasing pattern as the external P load over the years 1985-2020, and was likely driven by the decrease of the pool of releasable P. Similarly, the decreases in sediment loading by P retention in our P sorption experiment were associated with decreases in the concentration of the potentially mobile P forms (mainly Lab-P and Fe-P). These results show that changes in external P loading can successfully control internal P loading and are useful in water quality management of large lakes., 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.)

Photoautotrophic picoplankton (0.2-2 μm) can be a major contributor to primary production and play a significant part in the ecosystem carbon flow. However, the understanding about the dynamics of both eukaryotic and prokaryotic components of picoplankton in shallow eutrophic freshwater environments is still poor. Very few studies in these ecosystems reveal the taxonomic composition of picoeukaryotes. The main objective of this study was to investigate the seasonal dynamics of phototrophic picoplankton with the emphasis on the eukaryote community composition in a large shallow, eutrophic lake of the northern temperate zone (Lake Võrtsjärv). Phytoplankton pigments were employed to determine the taxonomic composition of photoautotrophic picoplankton. We found out that photoautotrophic picoplankton constitutes an important part of the phytoplankton community in Lake Võrtsjärv and its contribution can be highly variable (from ~9.3% to ~39%) in different years. The eukaryotic photoautotrophic picoplankton was dominated by diatoms followed by chrysophytes and other minor groups. Picoeukaryotes were prevailing in low-light conditions and low temperatures as their predominance in the picoplankton community was tightly linked to the presence or absence of ice cover. Ice cover strongly suppressed the growth of picocyanobacteria. Total phosphorus, turbidity and metazooplankton abundance had a clear relationship with photoautotrophic picoplankton chlorophyll a., Competing Interests: The authors have declared that no competing interests exist.

Reconstructions of past food web dynamics are necessary for better understanding long-term impacts of climate change on subarctic lakes. We studied elemental and stable isotopic composition of sedimentary organic matter, photosynthetic pigments and carbon stable isotopic composition of Daphnia (Cladocera; Crustacea) resting eggs (δ 13 C Clado ) in a sediment record from a small subarctic lake. We examined how regional climate and landscape changes over the last 5800 years affected the relative importance of allochthonous and autochthonous carbon transfer to zooplankton. Overall, δ 13 C Clado values were well in line with the range of theoretical values of aquatic primary producers, confirming that zooplankton consumers in subarctic lakes, even in the long-term perspective, are mainly fuelled by autochthonous primary production. Results also revealed greater incorporations of benthic algae into zooplankton biomass in periods that had a warmer and drier climate and clearer water, whereas a colder and wetter climate and lower water transparency induced higher contributions of planktonic algae to Daphnia biomass. This study thus emphasizes long-term influence of terrestrial-aquatic linkages and in-lake processes on the functioning of subarctic lake food webs.

With increasing primary productivity, ciliates may become the most important members of the microbial loop and form a central linkage in the transformation of microbial production to upper trophic levels. How metazooplankters, especially copepods, regulate ciliate community structure in shallow eutrophic waters is not completely clear. We carried out mesocosm experiments with different cyclopoid copepod enrichments in a shallow eutrophic lake to examine the responses of ciliate community structure and abundance to changes in cyclopoid copepod biomass and to detect any cascading effects on bacterioplankton and edible phytoplankton. Our results indicate that an increase in copepod zooplankton biomass favours the development of small-sized bacterivorous ciliates. This effect is unleashed by the decline of predaceous ciliate abundance, which would otherwise graze effectively on the small-sized ciliates. The inverse relationship between crustacean zooplankton and large predaceous ciliates is an important feature adjusting not only the structure of the ciliate community but also the energy transfer between meta- and protozooplankton. Still we could not detect any cascading effects on bacterio- or phytoplankton that would be caused by the structural changes in the ciliate community., (Copyright © 2015 Elsevier GmbH. All rights reserved.)

Pigment-based chemotaxonomy and CHEMTAX software have proven to be a valuable phytoplankton monitoring tool in marine environments, but are yet underdeveloped to determine algal assemblages in freshwater ecosystems. The main objectives of this study were (1) to compare the results of direct microscopy and CHEMTAX in describing phytoplankton community composition dynamics in a large, shallow and eutrophic lake; (2) to analyze the efficiency of the pigment-based method to detect changes in phytoplankton seasonal dynamics and during rapid bloom periods; (3) to assess the suitability of specific marker pigments and available marker pigment:chlorophyll a ratios to follow seasonal changes in eutrophic freshwater environment. A 5-year (2009-2013) parallel phytoplankton assessment by direct microscopy and by CHEMTAX was conducted using published marker pigment:chlorophyll a ratios. Despite displaying some differences from microscopy results, the pigment-based method successfully described the overall pattern of phytoplankton community dynamics during seasonal cycle in a eutrophic lake. Good agreement between the methods was achieved for most phytoplankton groups - cyanobacteria, chlorophytes, diatoms and cryptophytes. The agreement was poor in case of chrysophytes and dinoflagellates. Our study shows clearly that published marker pigment:chlorophyll a ratios can be used to describe algal class abundances, but they need to be calibrated for specific freshwater environment. Broader use of this method would enable to expand monitoring networks and increase measurement frequencies of freshwater ecosystems to meet the goals of the Water Framework Directive.

We used HPLC to identify and quantify pigments in a Holocene sediment record from large, shallow Lake Peipsi, Estonia. The aim of our study was to track the influence of long-term climate change (i.e. temperature fluctuations) on past dynamics of aquatic primary producers. Sedimentary pigments were separated and quantified in 182 samples that span the last ca. 10,000 years. There was an increasing trend in sedimentary pigment concentrations from basal to upper sediment layers, suggesting a gradual increase in lake trophic status through time. Using additive models, our results suggested that primary producer dynamics in Lake Peipsi were closely related to temperature fluctuations. We, however, identified two periods (early Holocene and after ca. 2.5 cal ka BP) when the relationship between primary producer composition and temperature was weak, suggesting the influence of additional drivers on the primary producer community. We postulate that: (a) the increase of primary producer biomass in the early Holocene could have been caused by input of allochthonous organic matter and nutrients from the flooded areas when water level in Lake Peipsi was increasing, and (b) changes in the abundance and structure of primary producer assemblages since ca. 2.5 cal ka BP was related to widespread agricultural activities in the Lake Peipsi catchment. These results suggest that human activities can disrupt the relationship between the primary producer community and temperature in large, shallow lakes. [ABSTRACT FROM AUTHOR]

Abstract: A palaeolimnological study, covering the last c. 12,000 years, was conducted in a small subalpine lake located in the Alps to study climate change impacts on carbon flows through food webs in small lakes. We used analysis of sedimentary pigments and carbon stable isotopic composition of chironomid remains (δ13CHC) to reconstruct past dynamics of phytoplankton community and carbon sources sustaining benthic consumers. Chironomid biomass was sustained by a combination of allochthonous, autochthonous and CH4‐derived organic matters, and their relative contributions were correlated to changes in temperature. Relatively high terrestrial contributions to chironomid biomass were observed during period of the Holocene when in‐lake production was low. Relatively high incorporation of CH4‐derived carbon to chironomid biomass was found during anoxic events co‐occurring with the Holocene thermal maximum. Results were then compared with those collected in a small boreal lake in Estonia. We tested the hypothesis that responses in carbon flows through benthic food web to past climate change would be similar between these lakes. We found a negative correlation between δ13CHC values of both lakes and inferred air temperature, suggesting that temperature was the major driver to different food sources being incorporated into chironomid biomass. Our study demonstrated that air temperature was the principal driver of the energy flows through benthic food web in the studied small lakes. We conjectured that carbon cycling in food webs of small lakes might be strongly sensitive to climate change. [ABSTRACT FROM AUTHOR]

Small-bodied cladocerans and cyclopoid copepods are becoming increasingly dominant over large crustacean zooplankton in eutrophic waters where they often coexist with cyanobacterial blooms. However, relatively little is known about their algal diet preferences. We studied grazing selectivity of small crustaceans (the cyclopoid copepods Mesocyclops leuckarti, Thermocyclops oithonoides, Cyclops kolensis, and the cladocerans Daphnia cucullata, Chydorus sphaericus, Bosmina spp.) by liquid chromatographic analyses of phytoplankton marker pigments in the shallow, highly eutrophic Lake Võrtsjärv (Estonia) during a seasonal cycle. Copepods (mainly C. kolensis) preferably consumed cryptophytes (identified by the marker pigment alloxanthin in gut contents) during colder periods, while they preferred small non-filamentous diatoms and green algae (identified mainly by diatoxanthin and lutein, respectively) from May to September. All studied cladoceran species showed highest selectivity towards colonial cyanobacteria (identified by canthaxanthin). For small C. sphaericus, commonly occuring in the pelagic zone of eutrophic lakes, colonial cyanobacteria can be their major food source, supporting their coexistence with cyanobacterial blooms. Pigments characteristic of filamentous cyanobacteria and diatoms (zeaxanthin and fucoxanthin, respectively), algae dominating in Võrtsjärv, were also found in the grazers’ diet but were generally avoided by the crustaceans commonly dominating the zooplankton assemblage. Together these results suggest that the co-occurring small-bodied cyclopoid and cladoceran species have markedly different algal diets and that the cladocera represent the main trophic link transferring cyanobacterial carbon to the food web in a highly eutrophic lake. [ABSTRACT FROM AUTHOR]

We applied a multi-proxy palaeolimnological approach to provide insights into the natural variability and human-mediated trends of two interconnected temperate large shallow lakes, Peipsi and Võrtsjärv, during the twentieth century. The history of the lakes was assessed on the basis of age-related changes in the sediment main constituents (water, organic matter and carbonate), sub-fossil pigments, diatom assemblages and organic matter dissolved in pore water. The temporal changes in the palaeodata indicate an increase of the in-lake biological production in both lakes from about the 1960s, suggesting enhanced nutrient inputs. In subsequent decades, the gradual increase of autochthonous organic matter becomes more obvious, indicating progressive eutrophication of the lakes. Palaeolimnological indicators from the sediment record of Lake Peipsi indicate a slight recession of the lake's eutrophication in the 1990s but not for Lake Võrtsjärv. The results of the study also suggest that after the lakes became eutrophied, the climatically induced water-level fluctuations ceased to be the main driver determining the abundance of phytoplankton. Responses of the lakes to human-induced impacts are better recorded in the sediments of Lake Peipsi than in those of Lake Võrtsjärv, which is shallower of the two and where the wave-induced resuspension of deposits markedly smooths or erases the signals of environmental changes. The results of the investigation expand the knowledge on how large shallow lakes respond to human-mediated and natural perturbations, including those in the lake catchment areas and the capability of the lakes to store the chronology and sequence of these changes. [ABSTRACT FROM AUTHOR]

Abstract: Analytical method based on HPLC has been used to characterize aquatic dissolved organic matter (DOM) from sediment core of Lake Võrtsjärv, South- Estonia. High-performance size exclusion chromatography (HPSEC) as separation method was coupled with diode-array detection (DAD) and separated molecular fractions of DOM were subject to qualitative and semi-quantitative analysis. Qualitative analysis based on UV- spectra revealed the presence of proteins and humic constituents in separated high molecular weight fraction and aromatic constituents in low molecular weight fraction. Statistical data treatment methods enabled clustering sediment layers into 4 periods according to sediment depth and age. The upper 0-30 cm sediment DOM had statistically relevant differences in comparison to other periods as revealed by lower total peak and humic substances (HS) fraction areas, and molecular weights. Samples from 80-120 cm depth differed from others by decreased low molecular weight (LMW) fraction content. The observed down-core trends suggest polymerization of LMW organic constituents and increasing humification. The statistical analyses revealed that some chromatographic and spectrometric parameters can be used to differentiate between sediment layers and to evaluate environmental changes. [Copyright &y& Elsevier]

We investigated stratigraphic changes in fossil pigments and the molecular structure of the UV-absorbing fraction of pore-water dissolved organic matter in a sedimentary record from Lake Peipsi (Estonia/Russia) temporally covering the 20th century. The aims of the study were to define the onset of eutrophication in the lake and to track its course. An attempt was also made to reconstruct lake conditions before the intensive nutrient loading began. Fossil pigment analysis indicated that the eutrophication of the lake started in the 1960s and accelerated in the 1970s. Sedimentary pigments also indicate a continuing tendency of the lake ecosystem towards eutrophy in the 1980s and 1990s. However, changes in the molecular size structure of pore-water dissolved organic matter indicated that the contribution of autochthonous matter to the organic pool of the lake ecosystem had already started to increase around the end of the 1930s. We conclude that this rise was generated by a coincidence of several anthropogenic and natural factors. The pore-water data also show that a slight relative reduction in the autochthonous organic matter took place in the 1990s. A discordance in the paleodata obtained for the beginning of the 20th century complicates clear conclusions about earlier conditions in the lake. On the one hand, the qualitative characteristics of pore-water dissolved organic matter and the low concentration of chlorophyll a indicate that the phytoplankton biomass was low in Lake Peipsi during that period. On the other hand, the concentrations of marker pigments of specific phytoplankton groups are high, comparable with the values in the recent sediments. Possible reasons for the high levels of these pigments in the early 1900s sediments, such as a shift in the preservation conditions of organic substances and their transport from the lake’s catchment, are discussed. [ABSTRACT FROM AUTHOR]

In the 1990s a sharp decrease in nitrogen loading occurred in Estonian rivers, bringing about a reduction of the nitrogen-to-phosphorus ratio (N:P ratio) in the large shallow lakes, Peipsi (3,555 km2, mean depth 7.1 m) and Võrtsjärv (270 km2, 2.8 m). The average mass ratio of total nitrogen (TN) and total phosphorus (TP) in Võrtsjärv (45) was about twice as high as that in Peipsi (22). In Peipsi, the N2-fixing Gloeotrichia echinulata, Aphanizomenon flos-aquae and Anabaena species prevailed in the summer phytoplankton, while in Võrtsjärv the dominant cyanobacteria were Limnothrix planktonica, L. redekei and Planktolyngbya limnetica, which cannot fix N2; the main N2-fixing taxa Aphanizomenon skujae and Anabaena sp. seldom gained dominance. In May–October the critical TN:TP mass ratio, below which N2-fixing cyanobacteria (Nfix) achieved high biomasses, was ∼40 in Võrtsjärv and ∼30 in Peipsi. The percentages of both total cyanobacteria (CY) and Nfix (CY% and Nfix%) in Peipsi achieved their maximum values at an N:P mass ratio at or below 20 for both TN:TP and Nmin:SRP. In Võrtsjärv, the TN:TP supporting a high Nfix% was between 30 and 40 and the Nmin:SRP supporting this high percentage was in the same range as that in Peipsi (<20), though the maximum Nfix% values in Võrtsjärv (69%) were much lower than in Peipsi (96%). The Nmin:SRP ratio explained 77% of the variability in Nfix% in May–October. The temperature dependence of Nfix% approximated to the maximum function type, with an upper limiting value at a certain water temperature, and this was most distinct in May–October. The critical TN:TP ratios obtained from our study (roughly 30 for Peipsi and 40 for Võrtsjärv) are much higher than the Redfield N:P mass ratio routinely considered (7). Our results represent valuable guidelines for creating effective management strategies for large shallow lakes. They provide a basis for stressing the urgent need to decrease phosphorus loading and to keep the in-lake P concentration low, and not to implement nitrogen reduction measures without a simultaneous decrease of phosphorus concentration. [ABSTRACT FROM AUTHOR]

Stratified lakes often receive more attention from limnologists although most of the world's lakes are indeed shallow. This study describes ecological processes in three shallow lakes, one dominated by macrophytes, one by phytoplankton, and one with both of them. Investigations focused on abiotic (water chemistry) as well as biotic factors such as phytoplankton, bacteria, zooplankton, and macrophytes. The macrophyte- and plankton-dominated lakes differed clearly in water transparency, which was far greater in macrophyte-dominated lakes due to phytoplankton setback in summer. Similar nutrient and zooplankton dynamics occurred in both types of lakes, suggesting that an alternative mechanism may account for the observed phytoplankton decline in macrophyte lakes. In contrast, herbivorous ciliates dominated only in the macrophyte lake. More intensive ciliate grazing could be one factor causing a decrease in phytoplankton numbers. In the plankton-dominated lake bacteria were mostly related to phytoplankton production but that was not the case in the macrophyte-dominated lake. In the plankton-dominated lake the grazing rate of ciliates on bacteria was higher than in the macrophyte-dominated one. However, when grazing on small algae was considered, the highest grazing rates were found in the macrophyte-dominated lake. This suggests that the microbial loop was weaker in the macrophyte-dominated lake than in the plankton-dominated lake. [ABSTRACT FROM AUTHOR]

The dynamics of different nitrogen compounds and nitrification in diverse habitats of a stratified Lake Verevi (Estonia) was investigated in 2000–2001. Also planktonic N2-fixation (N2fix) was measured in August of the observed years. The nitrogen that accumulated in the hypolimnion was trapped in the non-mixed layer during most of the vegetation period causing a concentration of an order of magnitude higher than in the epilimnion. The ammonium level remained low in the epilimnion (maximum 577 mgN m−3, average 115 mgN m−3) in spite of high concentrations in the hypolimnion (maximum 12223 mgN m−3, average 4807 mgN m−3). The concentrations of NO and NO remained on a low level both in the epilimnion (average 0.94 and 9.09 mgN m−3, respectively) and hypolimnion (average 0.47 and 5.05 mgN m−3, respectively). N2fix and nitrification ranged from 0.30 to 2.80 mgN m−3 day−1 and 6.0 to 107 mgN m−3 day−1, respectively; the most intensive processes occurred in 07.08.00 at depths of 2 and 5 m, accordingly. The role of N2fix in the total nitrogen budget of Lake Verevi (in August 2000 and 2001) was negligible while episodically in the nitrogen-depleted epilimnion the N2fix could substantially contribute to the pool of mineral nitrogen. Nitrification was unable to influence nitrogen dynamics in the epilimnion while some temporary coupling with ammonium dynamics in the hypolimnion was documented. [ABSTRACT FROM AUTHOR]

In Chara-dominated shallow eutrophic Lake Prossa (Estonia), the collapse of spring phytoplankton community occurred in late May after which both primary production (PP) and phytoplankton biomass (B&pinf;) stayed at a very low level. By mid-June the Secchi depth had increased up to 2.6 m indicating the achievement of the `clear water phase', which persisted thoughout the rest of the vegetation period. The biomass of `edible' phytoplankton formed on average 53% of the total phytoplankton biomass, and the share of herbivorous zooplankton was on average 61% of the total zooplankton biomass. In spring zooplankton removed daily 27% of the total B&pinf; and 29% of PP by grazing while in summer these values rarely exceeded 5%. Zooplankton grazing was responsible for the decrease of `edible' (<31 μm) phytoplankton after its spring peak as well as for maintaining its biomass at a very low level during the whole vegetation period. Depletion of mineral forms of nitrogen and phosphorus that occurred most probably because of the development of charophytes by the end of May supported the collapse of the whole phytoplankton community and kept the water clear throughout the summer and autumn. [ABSTRACT FROM AUTHOR]

The hydrochemical regime and the biota in two lakes of Vooremaa landscape protection area, Central Estonia, were studied in 2000–2001 within the frames of the EC project ECOFRAME aimed to work out water quality criteria for shallow lakes in Europe. Lake Prossa is a macrophyte-dominated lake with an area of 33 ha and a mean depth of 2.2 m. Most of the bottom in this lake is covered by a thick mat of charophytes all the year round. Lake Kaiavere located at a distance of 10 km from the former is a much larger (250 ha, mean depth 2.8 m) plankton-dominated lake. Despite the different area and the Chara meadows in Lake Prossa, the nutrient dynamics were very similar in both lakes. The first vernal phytoplankton peak was expressed in reduced Secchi depth in both lakes. After that peak, the water became clear in L. Prossa but remained turbid in Lake Kaiavere. In the Chara-lake the mean individual weight of zooplankton reached its maximum in both years in June. In the year 2000 it coincided with the peak of the zooplankton/phytoplankton ratio and with the minimum of soluble reactive phosphorus both contributing to the initiation of the clear-water period. Towards autumn, the individual mean weight of zooplankton decreased in the Chara-lake and remained smaller than in the plankton-dominated lake. Hence, zooplankton grazing could initiate the clear-water phase in the Chara-lake but other factors were needed for its maintenance. Another parameter that showed a clear difference between these two lakes was the carbonate alkalinity. It was rather stable or even increasing during spring in the plankton-dominated lake, while it decreased by nearly 50% between April and July in the Chara-lake. The calculated saturation level of calcite remained still lower in the Chara-lake showing that the co-precipitation of algae with autochthonous calcite could not explain the observed increase in water transparency above Chara meadows. The reduced sediment resuspension and the possible allelopathic influence of charophytes on phytoplankton remain the main explanations for the maitenance of the extensive clear-water period in the Chara-lake. [ABSTRACT FROM AUTHOR]

The fixation of molecular nitrogen (N2fix) by cyanobacteria in situ and in PO4-P enrichment experiments was investigated in large shallow Lake Võrtsjärv in 1998–2000. In this lake, N2fix started when TN/TP mass ratio was about 20, which is much higher than Redfield mass ratio 7. The rate of N2fix varied between 0.81 and 2.61 μgN l-1 d-1 and maximum rate (2.61 μgN l-1 d-1) was measured in 15.08.2000. In L. Võrtsjärv a lag period of a couple of weaks occurred between the set-up of favourable conditions for N2fix as the appearance of N2-fixing species and depletion of mineral nitrogen, and the real N2fix itself. However, if the favorable conditions for N2fix occurred in the lake, N2fix started after enrichment with PO4-P in mesocosms even then when no N2fix was detected in the lake. N2fix in mesocosms was also more intensive than in lake water. In our experiments PO4-P concentrations higher than 100 μgP l-1started to inhibit N2fix. [ABSTRACT FROM AUTHOR]