Distribution of Sediment Phosphorus Fractions in Hypertrophic Strongly Stratified Lake Verevi

Lake Verevi is a hypertrophic and strongly stratified (partly meromictic) small temperate lake. Vertical distribution of sediment phosphorus fractions as well as iron, manganese, organic matter and calcium carbonate of the deep bottom sediment was determined. The study focused on the ecologically important layer of the sediment [<20(45) cm]. In the uppermost layers of the sediment, NaOH-NRP (organic P) dominated while HCl-RP (apatite-P) became dominant in some deeper layers below 7 cm. Extremely high concentrations of labile phosphorus fraction (NH₄Cl-RP) indicated the low binding capacity of phosphorus by lake sediment. Due to sediment and hypolimnion anoxia, the internal load of phosphorus in this lake is most likely. Potentially mobile phosphorus fractions (NH₄Cl-RP, BD-RP, NaOH-NRP) formed 301 kg in upper 10 cm thick sediment layer of hypolimnetic bottom sediment (40% of lake bottom area).     

Laboratory-determined Phosphorus Flux from Lake Sediments as a Measure of Internal Phosphorus Loading

Eutrophication is a water quality issue in lakes worldwide, and there is a critical need to identify and control nutrient sources. Internal phosphorus (P) loading from lake sediments can account for a substantial portion of the total P load in eutrophic, and some mesotrophic, lakes. Laboratory determination of P release rates from sediment cores is one approach for determining the role of internal P loading and guiding management decisions. Two principal alternatives to experimental determination of sediment P release exist for estimating internal load: in situ measurements of changes in hypolimnetic P over time and P mass balance. The experimental approach using laboratory-based sediment incubations to quantify internal P load is a direct method, making it a valuable tool for lake management and restoration.Laboratory incubations of sediment cores can help determine the relative importance of internal vs. external P loads, as well as be used to answer a variety of lake management and research questions. We illustrate the use of sediment core incubations to assess the effectiveness of an aluminum sulfate (alum) treatment for reducing sediment P release. Other research questions that can be investigated using this approach include the effects of sediment resuspension and bioturbation on P release.The approach also has limitations. Assumptions must be made with respect to: extrapolating results from sediment cores to the entire lake; deciding over what time periods to measure nutrient release; and addressing possible core tube artifacts. A comprehensive dissolved oxygen monitoring strategy to assess temporal and spatial redox status in the lake provides greater confidence in annual P loads estimated from sediment core incubations.     

Eutrophication control strategies for three shallow Vecht lakes in the province of North Holland

Like many shallow surface waters in the Netherlands the North Holland Vecht lakes, formerly known for their rich variety of flora and fauna, now face a serious eutrophication problem. Nutrient enrichment has been mainly in the form of (treated) wastewater discharges, and the continuing ingress of nutrient-loaded water from the river Vecht. Yet, this water has to be supplied in order to compensate for water shortages resulting from (i) changes in the groundwater flow pattern due to reclamation of the deep polder Horstermeer, (ii) extensive groundwater extraction in the Gooi hills, and (iii) extensive drainage for agricultural purposes.The present policy of eutrophication abatement and restoration of the Ankeveen and Kortenhoef lakes ecosystems is focused on eliminating wastewater discharges and Vecht water supply. It also allows for additional dredging measures. Because of the un-suitable major ion composition of the Vecht, the aim is to compensate for this water supply by (i) partial restoration of the original groundwater flow from the Gooi hills and (ii) periphere additional supply with fresh seepage water from the skirts of the Horstermeer polder. However, uncertainty exists about the amounts of water needed.Water balances and phosphorus budgets have been established to ascertain the water demands of the lakes and to gain a detailed insight into the nutrient fluxes through the lakes. A groundwater flow model is used to assess the beneficial effects of the proposed measures.The results obtained, question the current unilateral restoration objectives. Calculations reveal that, both in the present situation and after (total) reduction of groundwater extractions in the future, the available quantity of fresh seepage water from the skirts of the polder Horstermeer is not sufficient to replace the inlet from the river Vecht into the Kortenhoef lakes. Additional supply options are available but the ones favoured from an ecological viewpoint are either the most expensive or less favoured from a social point of view. Although the sediments of the lakes appear to be a major source of eutrophication, the possibility of dredging the lakes will be considered only after reviewing results of a pilot-dredging project in the Hollands Ankeveen lakes in 1991.     

Lake restoration using a reed swamp to remove nutrients from non-point sources

The reed swamp adjacent to Lake Glumsø was partly separated from the lake by a dam and the inflow to the reed swamp controlled by pumping from the tributary. An investigation of the nutrient balances for the reed swamp showed an average daily denitrification rate of 252 mg/m2 from August 1988 to March 1989. Mineralization of the peat took place simultaneously, in sufficient amounts to supply carbon for the denitrification process. Ammonia and phosphorus were released in the ratio 7:1, corresponding to 111 mg N and 16 mg P/m²/ day. Most of the ammonia was nitrified and denitrified (included in the above mentioned denitrification rate). The net release of phosphorus implies that the method only can be applied satisfactorily in situations where nitrogen is the limiting nutrient. This is the case for Lake Glumso during the summer period, but not during the spring.This paper was presented at the INTECOL IV International Wetlands Conference in Columbus, Ohio, 1992, as part of a session organized by Prof. S. E. Jørgensen and sponsored by the International Lake Environment Committee.Corresponding Editor: Prof K. R. Reddy     

Biomanipulation as an Application of Food-Chain Theory: Constraints, Synthesis, and Recommendations for Temperate Lakes

The aim of this review is to identify problems, find general patterns, and extract recommendations for successful biomanipulation. An important conclusion is that the pelagic food chain from fish to algae may not be the only process affected by a biomanipulation. Instead, this process should be viewed as the “trigger” for secondary processes, such as establishment of submerged macrophytes, reduced internal loading of nutrients, and reduced resuspension of particles from the sediment. However, fish reduction also leads to a high recruitment of young-of-the-year (YOY) fish, which feed extensively on zooplankton. This expansion of YOY the first years after fish reduction is probably a major reason for less successful biomanipulations. Recent, large-scale biomanipulations have made it possible to update earlier recommendations regarding when, where, and how biomanipulation should be performed. More applicable recommendations include (1) the reduction in the biomass of planktivorous fish should be 75% or more; (2) the fish reduction should be performed efficiently and rapidly (within 1–3 years); (3) efforts should be made to reduce the number of benthic feeding fish; (4) the recruitment of YOY fish should be reduced; (5) the conditions for establishment of submerged macrophytes should be improved; and (6) the external input of nutrients (phosphorus and nitrogen) should be reduced as much as possible before the biomanipulation. Recent biomanipulations have shown that, correctly performed, the method also achieves results in large, relatively deep and eutrophic lakes, at least in a 5-year perspective. Although repeated measures may be necessary, the general conclusion is that biomanipulation is not only possible, but also a relatively inexpensive and attractive method for management of eutrophic lakes, and in particular as a follow-up measure to reduced nutrient load. Received 14 April 1998; accepted 31 August 1998     

Support of benthic invertebrates by detrital resources and current autochthonous primary production: results from a whole-lake 13C addition

1. Secondary production of benthic invertebrates in lakes is supported by current autochthonous primary production, and by detritus derived from a combination of terrestrial inputs and old autochthonous production from prior seasons. We quantified the importance of these two resources for the dominant benthic insects in Crampton Lake, a 26 ha, clear-water system.
2. Daily additions of NaH¹³CO₃ to the lake caused an increase in the stable carbon isotope ratios ( δ ¹³C) of the current primary production of phytoplankton and periphyton. We measured the response of four insect groups (taxon-depth combinations) to this manipulation, quantifying their current autochthony (% reliance on current autochthonous primary production) by fitting dynamic mixing models to time series of insect δ ¹³C.
3. The δ ¹³C of all four groups increased in response to the manipulation, although the magnitude of response differed by taxon and by depth, indicating differences in current autochthony. Odonate larvae (Libellulidae and Corduliidae) collected at 1.5 m depth derived 75% of their C from current autochthonous primary production. Chironomid larvae collected at 1.5, 3.5 and 10 m depths derived, respectively, 43%, 39% and 17% of their C from current autochthonous primary production.
4. Both taxon-specific diet preferences and depth-specific differences in resource availability may contribute to differences in current autochthony. Our results demonstrate significant but incomplete support of insect production by current autochthony, and indicate that allochthonous inputs and old autochthonous detritus support a substantial fraction (25–83%) of insect production.     

Resilience of Alternative Stable States during the Recovery of Shallow Lakes from Eutrophication: Lake Veluwe as a Case Study

In this paper we analyze a long-term dataset on the recovery from eutrophication of Lake Veluwe (The Netherlands). Clear hysteresis was observed in a number of ecosystem variables: the route to recovery differed significantly from the route that led to loss of clear water. The macrophyte dominated state disappeared in the late 1960s at TP above 0.20 mg l⁻¹, whereas its return occurred at less than 0.10 mg TP l⁻¹. Several regime shifts resulting in the occurrence of three alternative stable states were observed over a period of 30 years. The turbid state showed resistance to change, despite a strong and prompt reduction in Chl-a following reduction of external P-loading. The most important component that determined hysteresis in the return to clear water was not internal P-loading, but a high level of nonalgal light attenuation (through sediment resuspension) maintained by the interaction between wind and benthivorous fish. Although Chara was able to re-colonize the most shallow parts of the lake, recovery stalled and for a number of years clear (above charophyte beds) and turbid (deeper parts of the lake) water co-existed, as a separate alternative state on route to full recovery. Lake-wide clear water was re-established after bream density had been reduced substantially. This allowed a return of zebra mussels to the lake, whose high filtration capacity helped in maintaining clear water. In this study, we were able to identify the main drivers of hysteresis and regime shifts, although formal demonstration of cause and effect was not possible on the basis of field data alone. We argue that resilience of the present clear water state of Lake Veluwe very much depends on sizable populations of a few keystone species, especially Chara (stoneworts) and Dreissena (zebra mussels), and that careful management of these species is equally important as control of nutrients. Lake management should strive to maintain and strengthen resilience of the ecosystem, and this should offer protection against a renewed collapse of the clear state.     

The Proportion of Empneuston and Total Atmospheric Inputs of Carbon,Nitrogen and Phosphorus in the Nutrient Budget of a Small Mesotrophic Lake (Piburger See,Austria)

Litter and wet traps were employed to determine the inputs of coarse organic matter (empneuston), total organic carbon, total nitrogen and phosphorus through dry fallout and precipitation to a soft-water, mesotrophic lake. The dispersal of airborne material over the lake surface was investigated, and a method for the calculation of total input was developed. The component of the particulate matter larger than 1 mm contained the largest proportion of organic carbon, while the dissolved and the fine particle (smaller than 1 mm) fractions made greater contributions to the inputs of nitrogen and phosphorus compounds. The importance of the large particle fraction of the airborne organic matter in the lake's nutrient budget during the autumnal fall of litter was confirmed. However, the maximum total airborne input seems to occur during spring and summer. The input of phosphorus compounds through the atmosphere to small forest lakes is large compared to that from other sources. Airborne material accounts for 39% of the total phosphorus loading.     

Abundance and production of bacteria, and relationship to phytoplankton production, in a large tropical lake (Lake Tanganyika)

1. Abundance and bacterial production (BP) of heterotrophic bacteria (HBact) were measured in the north and south basins of Lake Tanganyika, East Africa, during seasonal sampling series between 2002 and 2007. The major objective of the study was to assess whether BP can supplement phytoplankton particulate primary production (particulate PP) in the pelagic waters, and whether BP and particulate PP are related in this large lake. HBact were enumerated in the 0–100 m surface layer by epifluorescence microscopy and flow cytometry; BP was quantified using ³H‐thymidine incorporation, usually in three mixolimnion layers (0–40, 40–60 and 60–100 m). 2. Flow cytometry allowed three subpopulations to be distinguished: low nucleic acid content bacteria (LNA), high nucleic acid content bacteria (HNA) and Synechococcus‐like picocyanobacteria (PCya). The proportion of HNA was on average 67% of total bacterial abundance, and tended to increase with depth. HBact abundance was between 1.2 × 10⁵ and 4.8 × 10⁶ cells mL⁻¹, and was maximal in the 0–40 m layer (i.e. roughly, the euphotic layer). Using a single conversion factor of 15 fg C cell⁻¹, estimated from biovolume measurements, average HBact biomass (integrated over a 100‐m water column depth) was 1.89 ± 1.05 g C m⁻². 3. Significant differences in BP appeared between seasons, especially in the south basin. The range of BP integrated over the 0–100 m layer was 93–735 mg C m⁻² day⁻¹, and overlapped with the range of particulate PP (150–1687 mg C m⁻² day⁻¹) measured in the same period of time at the same sites. 4. Depth‐integrated BP was significantly correlated to particulate PP and chlorophyll‐a, and BP in the euphotic layer was on average 25% of PP. 5. These results suggest that HBact contribute substantially to the particulate organic carbon available to consumers in Lake Tanganyika, and that BP may be sustained by phytoplankton‐derived organic carbon in the pelagic waters.     

Recovery of three arctic stream reaches from experimental nutrient enrichment

1. Nutrient enrichment and resulting eutrophication is a widespread anthropogenic influence on freshwater ecosystems, but recovery from nutrient enrichment is poorly understood, especially in stream environments. We examined multi‐year patterns in community recovery from experimental low‐concentration nutrient enrichment (N + P or P only) in three reaches of two Arctic tundra streams (Kuparuk River and Oksrukuyik Creek) on the North Slope of Alaska (U.S.A.). 2. Rates of recovery varied among community components and depended on duration of enrichment (2–13 consecutive growing seasons). Biomass of epilithic algae returned to reference levels rapidly (within 2 years), regardless of nutrients added or enrichment duration. Aquatic bryophyte cover, which increased greatly in the Kuparuk River only after long‐term enrichment (8 years), took 8 years of recovery to approach reference levels, after storms had scoured most remnant moss in the recovering reach. 3. Multi‐year persistence of bryophytes in the Kuparuk River appeared to prevent recovery of insect populations that had either been positively (e.g. the mayfly Ephemerella, most chironomid midge taxa) or negatively (e.g. the tube‐building chironomid Orthocladius rivulorum) affected by this shift in dominant primary producer. These lags in recovery (of >3 years) were probably driven by the persistent effect of bryophytes on physical benthic habitat. 4. Summer growth rates of Arctic grayling (both adults and young‐of‐year) in Oksrukuyik Creek (fertilised for 6 years with no bryophyte colonisation), which were consistently increased by nutrient addition, returned to reference rates within 1–2 years. 5. Rates of recovery of these virtually pristine Arctic stream ecosystems from low‐level nutrient enrichment appeared to be controlled largely by duration of enrichment, mediated through physical habitat shifts caused by eventual bryophyte colonisation, and subsequent physical disturbance that removed bryophytes. Nutrient enrichment of oligotrophic Arctic stream ecosystems caused by climate change or local anthropogenic activity may have dramatic and persistent consequences if it results in the colonisation of long‐lived primary producers that alter physical habitat.     

Establishment of Microbial Eukaryotic Enrichment Cultures from a Chemically Stratified Antarctic Lake and Assessment of Carbon Fixation Potential

Lake Bonney is one of numerous permanently ice-covered lakes located in the McMurdo Dry Valleys, Antarctica. The perennial ice cover maintains a chemically stratified water column and unlike other inland bodies of water, largely prevents external input of carbon and nutrients from streams. Biota are exposed to numerous environmental stresses, including year-round severe nutrient deficiency, low temperatures, extreme shade, hypersalinity, and 24-hour darkness during the winter ¹. These extreme environmental conditions limit the biota in Lake Bonney almost exclusively to microorganisms ².Single-celled microbial eukaryotes (called "protists") are important players in global biogeochemical cycling ³ and play important ecological roles in the cycling of carbon in the dry valley lakes, occupying both primary and tertiary roles in the aquatic food web. In the dry valley aquatic food web, protists that fix inorganic carbon (autotrophy) are the major producers of organic carbon for organotrophic organisms ^{4, 2}. Phagotrophic or heterotrophic protists capable of ingesting bacteria and smaller protists act as the top predators in the food web ⁵. Last, an unknown proportion of the protist population is capable of combined mixotrophic metabolism ^{6, 7}. Mixotrophy in protists involves the ability to combine photosynthetic capability with phagotrophic ingestion of prey microorganisms. This form of mixotrophy differs from mixotrophic metabolism in bacterial species, which generally involves uptake dissolved carbon molecules. There are currently very few protist isolates from permanently ice-capped polar lakes, and studies of protist diversity and ecology in this extreme environment have been limited ^{8, 4, 9, 10, 5}. A better understanding of protist metabolic versatility in the simple dry valley lake food web will aid in the development of models for the role of protists in the global carbon cycle.We employed an enrichment culture approach to isolate potentially phototrophic and mixotrophic protists from Lake Bonney. Sampling depths in the water column were chosen based on the location of primary production maxima and protist phylogenetic diversity ^{4, 11}, as well as variability in major abiotic factors affecting protist trophic modes: shallow sampling depths are limited for major nutrients, while deeper sampling depths are limited by light availability. In addition, lake water samples were supplemented with multiple types of growth media to promote the growth of a variety of phototrophic organisms.RubisCO catalyzes the rate limiting step in the Calvin Benson Bassham (CBB) cycle, the major pathway by which autotrophic organisms fix inorganic carbon and provide organic carbon for higher trophic levels in aquatic and terrestrial food webs ¹². In this study, we applied a radioisotope assay modified for filtered samples ¹³ to monitor maximum carboxylase activity as a proxy for carbon fixation potential and metabolic versatility in the Lake Bonney enrichment cultures.     

Nutrient transfer from soil nematodes to plants: a direct pathway provided by the mycorrhizal mycelial network

A pathway for the transfer of nutrients from dead nematodes to mycorrhizal plants is described for the first time. Plants of Betula pendula were grown in transparent microcosms in the mycorrhizal (M) or non‐mycorrhizal (NM) condition, either with or without nematode necromass of known nitrogen (N) and phosphorus (P) contents as the major potential source of these elements. Plants colonized by the mycorrhizal fungus Paxillus involutus produced greater yields and had larger N and P contents in the presence of nematodes than did their NM counterparts. The symbiotic systems were shown to exploit the N and P originally contained in necromass more effectively, and to transfer the nutrients to the plants in quantities approximately double those seen in NM systems. Even so, NM plants obtained sufficient N and P from dead nematodes to enable some enhancement of growth. Our observations confirm that mycorrhizal fungi provide the potential for the recycling of nutrients contained in this quantitatively important component of the soil mesofauna and demonstrate that the symbiotic pathway is considerably more effective than that provided by saprotrophs alone. The consequences of this nutrient transfer pathway for nutrient recycling in temperate forest ecosystems are considered.     

Effects of aluminum,iron, oxygen and nitrate additions on phosphorus release from the sediment of a Danish softwater lake

The effects of oxygen, aluminum, iron and nitrate additions on phosphate release from the sediment were evaluated in the softwater Lake Vedsted, Denmark, by a 34-day experiment with undisturbed sediment cores. Six treatments were applied: (1) Control - O₂ (0–20% saturation), (2) O₂ (100% saturation) (3) Al³⁺ – O₂, (4) Fe³⁺ + O₂, (5) Fe³⁺ – O₂, and (6) NO₃ ^– – O₂. Al₂(SO₄)₃*18 H₂O and FeCl₃*4H₂O were added in amounts that theoretically should immobilize the exchangeable P-pool in the top 5 cm of the sediment, while sodium nitrate concentrations were increased to 5 mg N l^–1. The four treatments with metals or NO₃ ^– reduced the P efflux from the sediment significantly as compared to the suboxic control treatment. Mean accumulated P-release rates for suboxic treatments with Al³⁺, Fe³⁺, and NO₃ ^– were: –0.27 mmol m^–2 (st. dev = 0.02 mmol m^–2, N = 5), 0.58 mmol m^–2 (st. dev = 0.30 mmol m^–2, N = 5) and 1.40 mmol m^–2 (st. dev = 0.14 mmol m^–2, N = 5), respectively. The oxic treatment with Fe³⁺ had a P efflux of 0.36 mmol m^–2 (st. dev = 0.08 mmol m^–2, N = 5). The two highest P-release rates were observed in the control treatment and the treatment with O₂ (14.50 mmol m^–2 (st. dev = 3.90 mmol m^–2, N = 5) and 2.31 mmol m^–2 (st. dev = 0.80 mmol m^–2, N = 5), respectively). In order to identify changes in the P and Fe binding sites in the sediment as caused by the treatments, a sequential P extraction procedure was applied on the sediment before and after the efflux experiment. Addition of O₂, Fe³⁺ and NO₃ ^– to the sediment increased the amounts of oxidized Fe³⁺ and P_BD. Al³⁺ addition resulted in a lower fraction of P_BD but a correspondingly higher fraction of Al-bound P. Addition of Al³⁺ decreased the Fe-efflux from the suboxic sediment as well as the amount of oxidized Fe³⁺ in the sediment. This questions the use of Al compounds that contain sulfate because of the possible formation of FeS, which will restrict upward migration of Fe²⁺ and the formation of new Fe-oxides in the surface sediment. Instead, we suggest the use of AlCl₃ for lake restoration purposes.     

Response of a eutrophic, shallow subtropical lake to reduced nutrient loading

1. Lake Apopka (FL, U.S.A.) was subjected to decades of high nutrient loading from farms developed in the 1940s on converted riparian wetlands. Consequences included perennially high densities of cyanobacteria, low water transparency, elimination of submerged vegetation, modified fish community, and deposition of nutrient‐rich, flocculent sediments. 2. Initial steps were taken to reduce phosphorus (P) loading. Through strengthened regulation and purchase of farms for restoration, external P loading was reduced on average from 0.56 to 0.25 g P m^?2 year^?1 (55%) starting in 1993. The P loading target for the lake is 0.13 g P m^?2 year^?1. 3. For the first 6 years of P loading reduction the annual sedimentation coefficient (σ) averaged 13% less than the prior long‐term value (0.97 versus 1.11 year^?1). The sedimentation coefficient, σ, was lower in the last 3 years of the study, but this period included extreme low‐water conditions and may not be representative. Annual σ was negative (net P flux to the water column) only 1 year. 4. Wind velocity explained 43% of the variation in σ during the period before reductions in total phosphorus (TP) concentration of lake water, but this proportion dropped to 6% after TP reductions. 5. Annual mean TP concentrations differed considerably from values predicted from external loading and hydraulic retention time using the Vollenweider–Organization for Economic Co‐operation and Development relationship. Reductions in lake water TP concentration fit model predictions better when multiyear (3‐year) mean values were used. 6. Evidence available to date indicates that this shallow, eutrophic lake responded to the decrease in external P loading. Neither recycling of sediment P nor wind‐driven resuspension of sediments prevented improvements in water quality. Reductions in TP concentration were evident about two TP‐resident times (2 × 0.9 year) after programmes began to reduce P loading. Improvements in concentrations of chlorophyll a and total suspended solids as well as in Secchi transparency lagged changes in lake‐water TP concentration but reached similar magnitudes during the study.     

Coupling between benthic biomass of Microcystis and phosphorus release from the sediments of a highly eutrophic lake

Variations in microbial biomass and activity in the sediments of hypereutrophic Lake Vallentunasjön were followed during a period of 5 years. The data were compared to the calculated release of phosphorus from the sediments during the same period. A strong co-variation was found between biomass of Microcystis, heterotrophic bacterial activity in the sediments and internal phosphorus loading. These parameters exhibited mainly a declining trend during the investigation period. A pronounced stability of the sediment chemistry, including the fractional composition of the sediment phosphorus, during the studied period indicates that microbial activity affected the phosphorus release from the sediments. Calculations of the percentage of sediment bacteria that was associated to the mucilage of Microcystis colonies imply, together with the specific bacterial production, that Microcystis in the sediment stimulates bacterial production. In the highly phosphorus-saturated sediments of Lake Vallentunasjön this would ultimately lead to an increased release of phosphorus from the sediment. Lake Vallentunasjön does not follow the common pattern of recovery after reduction of external phosphorus loading. The large biomasses and long survival of Microcystis in the sediment are probably important reasons for the delayed recovery of the lake.     

Cascading Trophic Interactions from Fish to Bacteria and Nutrients after Reduced Sewage Loading: An 18-Year Study of a Shallow Hypertrophic Lake

The effects of major reductions in organic matter, total phosphorus (TP), and total nitrogen (TN) loading on the chemical environment, trophic structure, and dynamics of the hypertrophic, shallow Lake Søbygård were followed for 18 years. After the reduction in organic matter loading in 1976, the lake initially shifted from a summer clear-water state, most likely reflecting high grazing pressure by large Daphniaspecies, to a turbid state with extremely high summer mean chlorophyll a (up to 1400 μg L^{? 1}), high pH (up to 10.2), and low zooplankton grazing. Subsequently, a more variable state with periodically high grazing rates on phytoplankton and bacteria was established. Changes in zooplankton abundance and grazing could be attributed to variations in cyprinid abundance due to a fish kill (probably as a consequence of oxygen depletion) and pH-induced variations in fish recruitment and fry survival. The results suggest strong cascading effects of fish on the abundance and size of zooplankton and phytoplankton and on phytoplankton production. A comparatively weak cascading effect on ciliates and bacterioplankton is suggested. Due to high internal loading, only minor changes were observed in lake-water TP after a reduction in external TP loading of approximately 80% in 1982; net retention of TP was still negative 13 years after the loading reduction, despite a short hydraulic retention time of a few weeks. TN, however, decreased proportionally to the TN-loading reduction in 1987, suggesting a fast N equilibration. Only minor improvement in the environmental state of the lake has been observed. We suggest that another decade will be required before the lake is in equilibrium with present external P loading.