ALBERT

Notes: 1. The effect of total nitrogen (TN) and phosphorus (TP) loading on trophic structure and water clarity was studied during summer in 24 field enclosures fixed in, and kept open to, the sediment in a shallow lake. The experiment involved a control treatment and five treatments to which nutrients were added: (i) high phosphorus, (ii) moderate nitrogen, (iii) high nitrogen, (iv) high phosphorus and moderate nitrogen and (v) high phosphorus and high nitrogen. To reduce zooplankton grazers, 1+ fish (Perca fluviatilis L.) were stocked in all enclosures at a density of 3.7 individuals m−2.2. With the addition of phosphorus, chlorophyll a and the total biovolume of phytoplankton rose significantly at moderate and high nitrogen. Cyanobacteria or chlorophytes dominated in all enclosures to which we added phosphorus as well as in the high nitrogen treatment, while cryptophytes dominated in the moderate nitrogen enclosures and the controls.3. At the end of the experiment, the biomass of the submerged macrophytes Elodea canadensis and Potamogeton sp. was significantly lower in the dual treatments (TN, TP) than in single nutrient treatments and controls and the water clarity declined. The shift to a turbid state with low plant coverage occurred at TN 〉2 mg N L−1 and TP 〉0.13–0.2 mg P L−1. These results concur with a survey of Danish shallow lakes, showing that high macrophyte coverage occurred only when summer mean TN was below 2 mg N L−1, irrespective of the concentration of TP, which ranged between 0.03 and 1.2 mg P L−1.4. Zooplankton biomass and the zooplankton : phytoplankton biomass ratio, and probably also the grazing pressure on phytoplankton, remained overall low in all treatments, reflecting the high fish abundance chosen for the experiment. We saw no response to nutrition addition in total zooplankton biomass, indicating that the loss of plants and a shift to the turbid state did not result from changes in zooplankton grazing. Shading by phytoplankton and periphyton was probably the key factor.5. Nitrogen may play a far more important role than previously appreciated in the loss of submerged macrophytes at increased nutrient loading and for the delay in the re-establishment of the nutrient loading reduction. We cannot yet specify, however, a threshold value for N that would cause a shift to a turbid state as it may vary with fish density and climatic conditions. However, the focus should be widened to use control of both N and P in the restoration of eutrophic shallow lakes.

Notes: 1. The variable ecological response of lakes to reduced nutrient loading (oligotrophication) at sites in Europe and North America was discussed at a workshop held in Silkeborg (Denmark) in January 2003. Studies of lake oligotrophication were presented based on both long-term monitoring and data generated by palaeolimnological methods.2. This introduction to the special issue provides short summaries of a series of the papers presented and their limnological context. Results show that the majority of lakes had approached a new equilibrium of phosphorus (P) and nitrogen (N) concentrations 10–15 years (P) and 0–5 years (N) after a major reduction in loading, irrespective of hydraulic retention time. Phytoplankton biomass decreased and a shift towards meso-oligotrophic species dominance occurred. The fish responded surprisingly fast to the loading reduction in most lakes. As a result, the percentage of piscivores increased and total fish biomass declined markedly, which may explain an increase in the body size of cladocerans and an increase in the zooplankton to phytoplankton biomass ratio seen in many of the lakes.3. Monitoring has in general been initiated after the effects of eutrophication became apparent. In this context palaeolimnological techniques become very useful because they allow limnologists to extend time scales of coverage and to define restoration targets and baseline conditions. Moreover, lake sediments pre-dating anthropogenic disturbance can be used to examine ecological response to, for instance, climate variability, allowing problems associated with multiple stressors to be addressed.4. It is concluded that there is a great need for a synthetic, holistic approach to studying lake oligotrophication, combining multiple techniques of palaeolimnological sediment analysis with detailed but temporally limited long-term monitoring of chemical and biological variables. This is important, not least to assess future responses to nutrient loading reductions, as global warming will interact with a range of external stressors and ultimately affect lake management strategies to deal with the resultant feedbacks.

Notes: 1. The impact of changes in submerged macrophyte abundance on fish-zooplankton-phytoplankton interactions was studied in eighteen large-scale (100 m2) enclosures in a shallow eutrophic take. The submerged macrophytes comprised Potamategon pectinatus L., P. pusillus L. and Callitriche hermaphroditica L. while the fish fry stock comprised three-spined sticklebacks, Gasterosteus acuteatus L., and roach, Rutilus rutilus L.2. In the absence of macrophytes zooplankton biomass was low and dominated by cyclopoid copepods regardless of fish density, while the phytoplankton biovolume was high (up to 38 mm31) and dominated by small pennate diatoms and chlorococcales. When the lake volume infested by submerged macrophytes (PVI) exceeded 15–20% and the fish density was below a catch per unit effort (CPUE) of 10 (approx. 2 fry m−2), planktonic cladoceran biomass was high and dominated by relatively large-sized specimens, while the phytoplankton biovolume was low and dominated by small fast-growing flagellates. At higher fish densities, zooplankton biomass and average biomass of cladocerans decreased and a shift to cyclopoids occurred, while phytoplankton biovolume increased markedly and became dominated by cyanophytes and dinoflagellates.3. Stepwise multiple linear regressions on log-transformed data revealed that the biomass of Daphnia, Bosmina, Ceriodaphmia and Chydorus were all significantly positively related to PVI and negatively to the abundance of fish or PVI x fish. The average individual biomass of cladocerans was negatively related to fish, but unrelated to PVI. Calculated zooplankton grazing pressure on phytoplankton was positively related to PVI and negatively to PVI x fish. Accordingly the phytoplankton biovolume was negatively related to PVI and to PVI x zooplankton biomass. Cyanophytes and chryptophytes (% of biomass) were positively and Chlorococcales and diatoms negatively related to PVI, while cyanophytes and Chlorococcales were negatively related to PVI x zooplankton biomass. In contrast diatoms and cryptophytes were positively related to the zooplankton biomass or PVI x zooplankton.4. The results suggest that fish predation has less impact on the zooplankton community in the more structured environment of macrophyte beds, particularly when the PVI exceeds 15–20%. They further suggest that the refuge capacity of macrophytes decreases markedly with increasing fish density (in our study above approximately 10 CPUE). Provided that the density of planktivorous fish is not high, even small improvements in submerged macrophyte abundance may have a substantial positive impact on the zooplankton, leading to a lower phytoplankton biovolume and higher water transparency. However, at high fish densities the refuge effect seems low and no major zooplankton mediated effects of enhanced growth of macrophytes are to be expected.

Notes: 1. Concentrations of phosphorus, nitrogen and silica and alkalinity were monitored in eight shallow and four deep Danish lakes for 13 years following a phosphorus loading reduction. The aim was to elucidate the seasonal changes in nutrient concentrations during recovery. Samples were taken biweekly during summer and monthly during winter.2. Overall, the most substantive changes in lake water concentrations were seen in the early phase of recovery. However, phosphorus continued to decline during summer as long as 10 years after the loading reduction, indicating a significant, albeit slow, decline in internal loading.3. Shallow and deep lakes responded differently to reduced loading. In shallow lakes the internal phosphorus release declined significantly in spring, early summer and autumn, and only non-significantly so in July and August. In contrast, in deep lakes the largest reduction occurred from May to August. This difference may reflect the much stronger benthic pelagic-coupling and the lack of stratification in shallow lakes.4. Nitrogen only showed minor changes during the recovery period, while alkalinity increased in late summer, probably conditioned by the reduced primary production, as also indicated by the lower pH. Silica tended to decline in winter and spring during the study period, probably reflecting a reduced release of silica from the sediment because of enhanced uptake by benthic diatoms following the improved water transparency.5. These results clearly indicate that internal loading of phosphorus can delay lake recovery for many years after phosphorus loading reduction, and that lake morphometry (i.e. deep versus shallow basins) influences the patterns of change in nutrient concentrations on both a seasonal and interannual basis.

Notes: 1. For 13 years the response of the plankton and fish community to a decline in external phosphorus loading was studied in eight lakes with a mean depth 〈5 m. We conducted chi-square analyses of sign of slope (positive or negative) of bimonthly averages of plankton variables for the eight lakes versus time. For fish, we compared results from two periods, i.e. 1989–1994 versus 1994–2001 as less data were available.2. Fish community structure tended to respond to the lowered concentration of total phosphorus (TP), although not all changes were significant. While catch per unit effort (multi-mesh sized gill nets) of cyprinids (especially bream, Abramis brama and roach, Rutilus rutilus) was highest in the first 5-year period, the quantitative importance particularly of perch (Perca fluviatilis), pike (Esox lucius) and rudd (Scardinius erythropthalmus), a littoral species, increased significantly after 1994.3. No changes occurred in zooplankton biomass, except for an increase in November and December. Biomass of small cladocerans, however, declined during summer and autumn, and the proportion of Daphnia to cladoceran biomass also increased. Average body weight of Daphnia and that of all cladocerans increased. The proportion of calanoids among copepods decreased in summer and the average body weight of cyclopoids and calanoids decreased during summer and autumn/early winter.4. Total biovolume of phytoplankton declined significantly in March to June and tended to decline in November and December as well, while no significant changes were observed during summer and autumn. Non-heterocystous cyanobacteria showed a decreasing trend during summer and autumn, while heterocystous cyanobacteria increased significantly in late summer. An increase in late summer was also evident for cryptophytes and chrysophytes, while diatoms tended to decline during most seasons.5. We conclude that phytoplankton, and probably also fish, responded rapidly to reduced loading, whereas the effect on zooplankton was less pronounced. However, increases in body weight of cladocerans and the zooplankton to phytoplankton biomass ratio during summer indicate reduced top-down control on zooplankton and enhanced grazing on phytoplankton. This conclusion is supported by a tendency for fish biomass to decline and a shift towards greater dominance by piscivores and, thus, an increased likelihood of predator control of zooplanktivorous cyprinids.

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

Notes: 1. It is well accepted that fish, if abundant, can have a major impact on the zooplankton community structure during summer, which, particularly in eutrophic lakes, may cascade to phytoplankton and ultimately influence water clarity. Fish predation affects mean size of cladocerans and the zooplankton grazing pressure on phytoplankton. Little is, however, known about the role of fish during winter.2. We analysed data from 34 lakes studied for 8–9 years divided into three seasons: summer, autumn/spring and winter, and four lake classes: all lakes, shallow lakes without submerged plants, shallow lakes with submerged plants and deep lakes. We recorded how body weight of Daphnia and then cladocerans varied among the three seasons. For all lake types there was a significant positive correlation in the mean body weight of Daphnia and all cladocerans between the different seasons, and only in lakes with macrophytes did the slope differ significantly from one (winter versus summer for Daphnia).3. These results suggest that the fish predation pressure during autumn/spring and winter is as high as during summer, and maybe even higher during winter in macrophyte-rich lakes. It could be argued that the winter zooplankton community structure resembles that of the summer community because of low specimen turnover during winter mediated by low fecundity, which, in turn, reflects food shortage, low temperatures and low winter hatching from resting eggs. However, we found frequent major changes in mean body weight of Daphnia and cladocerans in three fish-biomanipulated lakes during the winter season.4. The seasonal pattern of zooplankton : phytoplankton biomass ratio showed no correlation between summer and winter for shallow lakes with abundant vegetation or for deep lakes. For the shallow lakes, the ratio was substantially higher during summer than in winter and autumn/spring, suggesting a higher zooplankton grazing potential during summer, while the ratio was often higher in winter in deep lakes. Direct and indirect effects of macrophytes, and internal P loading and mixing, all varying over the season, might weaken the fish signal on this ratio.5. Overall, our data indicate that release of fish predation may have strong cascading effects on zooplankton grazing on phytoplankton and water clarity in temperate, coastal situated eutrophic lakes, not only during summer but also during winter.

Notes: 1. Using data from 71, mainly shallow (an average mean depth of 3 m), Danish lakes with contrasting total phosphorus concentrations (summer mean 0.02–1.0 mg P L−l), we describe how species richness, biodiversity and trophic structure change along a total phosphorus (TP) gradient divided into five TP classes (class 1–5: 〈0.05, 0.05–0.1, 0.1–0.2, 0.2–0.4,〉 0.4 mg P L−1). 
2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floating-leaved macrophytes, species richness was unimodally related to TP, all peaking at 0.1–0.4 mg P L−1. The Shannon–Wiener and the Hurlbert probability of inter-specific encounter (PIE) diversity indices showed significant unimodal relationships to TP for zooplankton, phytoplankton and fish. Mean depth also contributed positively to the relationship for rotifers, phytoplankton and fish. 
3. At low nutrient concentrations, piscivorous fish (particularly perch, Perca fluviatilis) were abundant and the biomass ratio of piscivores to plankti-benthivorous cyprinids was high and the density of cyprinids low. Concurrently, the zooplankton was dominated by large-bodied forms and the biomass ratio of zooplankton to phytoplankton and the calculated grazing pressure on phytoplankton were high. Phytoplankton biomass was low and submerged macrophyte abundance high. 
4. With increasing TP, a major shift occurred in trophic structure. Catches of cyprinids in multiple mesh size gill nets increased 10-fold from class 1 to class 5 and the weight ratio of piscivores to planktivores decreased from 0.6 in class 1 to 0.10–0.15 in classes 3–5. In addition, the mean body weight of dominant cyprinids (roach, Rutilus rutilus, and bream, Abramis brama) decreased two–threefold. Simultaneously, small cladocerans gradually became more important, and among copepods, a shift occurred from calanoid to cyclopoids. Mean body weight of cladocerans decreased from 5.1 μg in class 1 to 1.5 μg in class 5, and the biomass ratio of zooplankton to phytoplankton from 0.46 in class 1 to 0.08–0.15 in classes 3–5. Conversely, phytoplankton biomass and chlorophyll a increased 15-fold from class 1 to 5 and submerged macrophytes disappeared from most lakes. 
5. The suggestion that fish have a significant structuring role in eutrophic lakes is supported by data from three lakes in which major changes in the abundance of planktivorous fish occurred following fish kill or fish manipulation. In these lakes, studied for 8 years, a reduction in planktivores resulted in a major increase in cladoceran mean size and in the biomass ratio of zooplankton to phytoplankton, while chlorophyll a declined substantially. In comparison, no significant changes were observed in 33 ‘control’ lakes studied during the same period.

Notes: Lake restoration in Denmark has involved the use of several different restoration techniques, all aiming to improve lake water quality and establishing clear-water conditions. The most frequently used method, now used in more than 20 lakes, is the reduction of zooplanktivorous and benthivorous fish (especially roach (Rutilus rutilus) and bream (Abramis brama)) with the objective of improving the growth conditions for piscivores, large-sized zooplankton species, benthic algae and submerged macrophytes. Piscivore stocking (mainly Esox lucius (pike)), aiming especially at reducing the abundance of young-of-the-year fish, has been used in more than 10 lakes and frequently as a supplement to fish removal. Hypolimnetic oxidation, with oxygen and nitrate, has been undertaken in a few stratified lakes and sediment dredging, with the purpose of diminishing the internal phosphorus loading, has been experimented with in one large, shallow lake. Submerged macrophyte implantation has been conducted in some of the biomanipulated lakes to increase macrophyte abundance and distribution. Overall, the results from lake restoration projects, in the mainly shallow Danish lakes, show that external nutrient loading must be reduced to a level below 0.05–0.1 mg P L–1 under equilibrium conditions to gain permanent effects on lake water quality. By using fish removal, at least 80% of the fish stock should be removed over a period of not more than 1–2 years to obtain a substantial effect on lower trophic levels and to avoid regrowth of the remaining fish stock. Stocking of piscivores requires high densities (〉0.1 individuals m–2) if an impact on the plankton level is to be obtained and stocking should be repeated yearly until a stable clear-water state is reached. The experiments with hypolimnetic oxygenation and sediment dredging confirm that internal phosphorus loading can be reduced. Experience from macrophyte implantation experiments indicates that protection against grazing by herbivorous waterfowl may be useful in the early phase of recolonization.

Notes: ABSTRACT 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.