Jens Peder Jensen

Top-down control in freshwater lakes: the role of nutrient state, submerged macrophytes and water depth

Based on data from 233 Danish lakes, enclosure experiments, full-scaleexperiments and published empirical models we present evidence that top-downcontrol is more important in shallow lakes than in deep lakes, excepting lakeswith a high abundance of submerged macrophytes. The evidence in support is: (1)That at a given epilimnion total phosphorus concentration (TP) the biomass offish per m2 is independent of depth, which means that biomassper m3is markedly higher in shallow lakes. (2) That the biomass of benthic invertebratesis higher in shallow lakes, which means that the benthi-planktivorous fish areless dependent on zooplankton prey than in deep lakes. By their ability to shiftto zooplankton predation their density can remain high even in periods whenzooplankton is scarce and they can thereby maintain a potentially high predationpressure on zooplankton. (3) That the possibilities of cladocerans to escapepredation by vertical migration are less. (4) That the zooplankton:phytoplanktonmass ratio per m2 is lower and presumably then also thegrazing pressure onphytoplankton. (5) That nutrient constraints appear to be weaker, as evidenced bythe fact that at a given annual mean TP, summer TP is considerably higher inshallow lakes, especially in eutrophic lakes lacking submerged macrophytes. (6)That negative feedback on cladocerans by cyanobacteria is lower as cyanobacterialdominance is less frequent in shallow lakes and more easily broken (at least inNorthern temperate lakes), and (7) That top-down control by benthi-planktivorousfish is markedly reduced in lakes rich in submerged macrophytes because theplants serve as a refuge for pelagic cladocerans and encouragepredatory fish at the expense of prey fish. We conclude that manipulation of fishand submerged macrophytes may have substantial impact on lake ecosystems, inparticular in shallow eutrophic lakes. On the contrary, if the conditions formore permanent changes in plant abundance or fish community structure are lackingthe feed-back mechanisms that endeavour a return to the original turbid state willbe particularly strong in shallow lakes.

Fish manipulation as a lake restoration tool in shallow, eutrophic, temperate lakes 2: threshold levels, long-term stability and conclusions

In order to evaluate short-term and long-term effects of fish manipulation in shallow, eutrophic lakes, empirical studies on relationships between lake water concentration of total phosphorus (P) and the occurrence of phytoplankton, submerged macrophytes and fish in Danish lakes are combined with results from three whole-lake fish manipulation experiments. After removal of less than 80 per cent of the planktivorous fish stock a short-term trophic cascade was obtained in the nutrient regimes, where large cyanobacteria were not strongly dominant and persistent. In shallow Danish lakes cyanobacteria were the most often dominating phytoplankton class in the P-range between 200 and 1000 pg P 1−1. Long-term effects are suggested to be closely related to the ability of the lake to establish a permanent and wide distribution of submerged macrophytes and to create self-perpetuating increases in the ratio of piscivorous to planktivorous fish. The maximum depth at which submerged macrophytes occurred, decreased exponentially with increasing P concentration. Submerged macrophytes were absent in lakes > 10 ha and with P levels above 250−300 μg P 1−1, but still abundant in some lakes 10 cm numerically contributed more than 80 per cent of the total planktivorous and piscivorous fish (> 10 cm) in the pelagical of lakes with concentrations above 100 μg P 1−1. Below this threshold level the proportion of planktivores decreased markedly to ca. 50 per cent at 22 μg P 1−1. The extent of the shift in depth colonization of submerged macrophytes and fish stock composition in the three whole-lake fish manipulations follows closely the predictions from the relationships derived from the empirical study. We conclude that a long-term effect of a reduction in the density of planktivorous fish can be expected only when the external phosphorus loading is reduced to below 0.5−2.0 g m −2 y −1. This loading is equivalent to an in-lake summer concentration below 80−150 μg P 1−1. Furthermore, fish manipulation as a restoration tool seems most efficient in shallow lakes.

Restoration of shallow lakes by nutrient control and biomanipulation—the successful strategy varies with lake size and climate

Major efforts have been made world-wide to improve the ecological quality of shallow lakes by reducing external nutrient loading. These have often resulted in lower in-lake total phosphorus (TP) and decreased chlorophyll a levels in surface water, reduced phytoplankton biomass and higher Secchi depth. Internal loading delays recovery, but in north temperate lakes a new equilibrium with respect to TP often is reached after <10–15 years. In comparison, the response time to reduced nitrogen (N) loading is typically <5 years. Also increased top-down control may be important. Fish biomass often declines, and the percentage of piscivores, the zooplankton:phytoplankton biomass ratio, the contribution of Daphnia to zooplankton biomass and the cladoceran size all tend to increase. This holds for both small and relatively large lakes, for example, the largest lake in Denmark (40 km2), shallow Lake Arreso, has responded relatively rapidly to a ca. 76% loading reduction arising from nutrient reduction and top-down control. Some lakes, however, have proven resistant to loading reductions. To accelerate recovery several physico-chemical and biological restoration methods have been developed for north temperate lakes and used with varying degrees of success. Biological measures, such as selective removal of planktivorous fish, stocking of piscivorous fish and implantation or protection of submerged plants, often are cheap versus traditional physico-chemical methods and are therefore attractive. However, their long-term effectiveness is uncertain. It is argued that additional measures beyond loading reduction are less cost-efficient and often not needed in very large lakes. Although fewer data are available on tropical lakes these seem to respond to external loading reductions, an example being Lake Paranoa, Brazil (38 km2). However, differences in biological interactions between cold temperate versus warm temperate-subtropical-tropical lakes make transfer of existing biological restoration methods to warm lakes difficult. Warm lakes often have prolonged growth seasons with a higher risk of long-lasting algal blooms and dense floating plant communities, smaller fish, higher aggregation of fish in vegetation (leading to loss of zooplankton refuge), more annual fish cohorts, more omnivorous feeding by fish and less specialist piscivory. The trophic structures of warm lakes vary markedly, depending on precipitation, continental or coastal regions locations, lake age and temperature. Unfortunately, little is known about trophic dynamics and the role of fish in warm lakes. Since many warm lakes suffer from eutrophication, new insights are needed into trophic interactions and potential lake restoration methods, especially since eutrophication is expected to increase in the future owing to economic development and global warming.

The impact of nutrient state and lake depth on top-down control in the pelagic zone of lakes: A study of 466 lakes from the temperate zone to the Arctic

Using empirical data from 466 temperate to arctic lakes covering a total phosphorus (TP) gradient of 2–1036 gL –1 , we describe how the relative contributions of resource supply, and predator control change along a nutrient gradient. We argue that (a) predator control on large-bodied zooplankton is unimodally related to TP and is highest in the most nutrient-rich and nutrient-poor lakes and generally higher in shallow than deep lakes, (b) the cascading effect of changes in predator control on phytoplankton decreases with increasing TP, and (c) these general patterns occur with significant variations—that is, the predation pressure can be low or high at all nutrient levels. A quantile regression revealed that the median share of the predatorsensitive Daphnia to the total cladoceran biomass was significantly related unimodally to TP, while the 10% and 90% percentiles approached 0 and 100%, respectively, at all TP levels. Moreover, deep lakes (more than 6 m) had a higher percentage of

Shallow lake restoration by nutrient loading reduction—some recent findings and challenges ahead

Shallow lakes respond to nutrient loading reductions. Major findings in a recent multi-lake comparison of data from lakes with long time series revealed: that a new state of equilibrium was typically reached for phosphorus (P) after 10–15 years and for nitrogen (N) after <5–10 years; that the in-lake Total N:Total P and inorganic N:P ratios increased; that the phytoplankton and fish biomass often decreased; that the percentage of piscivores often increased as did the zooplankton:phytoplankton biomass ratio, the contribution of Daphnia to Zooplankton biomass, and cladoceran size. This indicates that enhanced resource and predator control often interact during recovery from eutrophication. So far, focus has been directed at reducing external loading of P. However, one experimental study and cross-system analyses of data from many lakes in north temperate lakes indicate that nitrogen may play a more significant role for abundance and species richness of submerged plants than usually anticipated when total phos-phorus is moderate high. According to the alter-native states hypothesis we should expect ecological resistance to nutrient loading reduction and P hysteresis. We present results suggesting that the two alternative states are less stable than originally anticipated. How global warming af-fects the water clarity of shallow lakes is debat-able. We suggest that water clarity often will decrease due to either enhanced growth of phytoplankton or, if submerged macrophytes are stimulated, by reduced capacity of these plants to maintain clear-water conditions. The latter is supported by a cross-system comparison of lakes in Florida and Denmark. The proportion of small fish might increase and we might see higher aggregation of fish within the vegetation (leading to loss of Zooplankton refuges), more annual fish cohorts, more omnivorous feeding by fish and less specialist piscivory. Moreover, lakes may have prolonged growth seasons with a higher risk of long-lasting algal blooms and at places dense floating plant communities. The effects of global warming need to be taken into consideration by lake managers when setting future targets for critical loading, as these may well have to be adjusted in the future. Finally, we highlight some of the future challenges we see in lake restoration research.

Internal phosphorus loading in shallow Danish lakes

High phosphorus concentrations due to internal loading from the sediment with a strongly negative impact on lake water quality, is often seen in shallow lakes after a reduction of external loading. To analyse the nature of internal loading we studied 1. the seasonal phosphorus concentrations of 265 Danish shallow, mainly eutrophic lakes; 2. seasonal phosphorus mass balances and retention for eight years in 16 eutrophic lakes, and 3. phosphorus mass balances and changing sediment phosphorus profiles for 15 years in one hypertrophic lake. Lake water, inlets and outlets were routinely sampled 10–26 times annually. Total phosphorus (TP) concentrations during summer were two–four times higher than winter values in lakes with a mean summer total phosphorus concentration (TPsum) above 0.2 mg P 1−1. Annual phosphorus retention decreased with increasing TPsum and was lower than predicted from the Vollenweider model, particularly in lakes with TPsum above 0.2 mg P1−1. The seasonal phosphorus retention in lakes with TPsum below 0.1 mg P 1−1 was positive during the whole season, except July and August when mean retention ranged from −10 to −30% of inlet loading. In lakes with TPsum above 0.1 mg P 1−1, the retention was positive during winter, but negative from April to September. The negative retention was most pronounced in lakes with the highest TPsum, particularly in May and July when mean retention ranged from −50 to −68% in lakes with TPsum above 0.2 mg P 1−1. The retention was generally less negative in June, when a clearwater phase typically occurs and close to 0 also in lakes with a high TPsum. Mass balances from the hypertrophic lake have now shown a 15-yr net annual negative retention following reduced external loading. Sediment profiles suggest phosphorus release from depths down to 25 cm and that net internal phosphorus loading may persist for another 15 yrs. It is concluded that internal loading of shallow eutrophic lakes may have a considerable and persistent impact on summer TP after reduced external loading.

Functional ecology and palaeolimnology: using cladoceran remains to reconstruct anthropogenic impact.

The field of lake palaeoecology has undergone significant changes. Powerful quantitative techniques have been developed to investigate anthropogenic impacts on lakes. Inclusion of zooplankton and benthic chydorid cladocerans has provided previously unavailable information on the historical development of planktivorous fish populations, submerged macrophytes and lake production, and has been used to document exotic species introductions, rapid genetic evolution and human disturbance of lakes. In particular, new techniques now allow a more complete evaluation of changes in past and present trophic structure to be made, and provide insights on the rapid evolutionary responses of aquatic invertebrate communities to anthropogenic perturbation of lakes.

Long-term responses to fish-stock reduction in small shallow lakes: interpretation of five-year results of four biomanipulation cases in The Netherlands and Denmark

The effects of fish stock reduction have been studies in 3 Dutch lakes (Lake Zwemlust, Lake Bleiswijkse Zoom and Lake Noorddiep) and 1 Danish lake (Lake Væng) during 4–5 years. A general response id described. The fish stock reduction led in general to a low fish stock, low chlorophyii-a, high transparency and high abunuance of macrophytes. Large Daphnia became abundant, but their density decreased, due to food limitation and predation by fish. The total nitrogen concentration became low due to N-uptake by macrophytes and enhanced denitrification. In Lake Bleiswijkse Zoom the water transparency deteriorated and the clear water state was not stable. The fish stock increased and the production of young fish in summer was high. lear water occurred only in spring. Large daphnids were absent in summer and the macrophytes decreased.In Lake Zwemlust, Lake Væng and Lake Noorddiep the water remained clear during the first five years. In summer of the sixth year (1992) transparency decreased in Lake Zwemlust (with high P-concentration of 1.0 mg P l-1). Also in Lake Væng (with a low nutrient concentration of 0.15 mg P.⋆-1) a short term turbid stage (1.5 month) occurred in summer 1992 after a sudden collapse of the macrophytes. Deterioration of the water quality seems to start in summer and seems related to a collapse in macrophytes. At a low planktivorous fishstock (e.g. Lake Væng)thhe duration of the turbid state is shorter. than in presence of a high planktivorous fish biomass (e.g. Lake Zwemlust, and later years of Lake Bleiswijkse Zoom).

Cascading trophic interactions from fish to bacteria and nutrients after reduced sewage loading: An 18-year study of a shallow hypertrophic lake

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.

Trophic dynamics in turbid and clearwater lakes with special emphasis on the role of zooplankton for water clarity

Within a certain nutrient level shallow lakes may alternate between two states, a clearwater and a turbid state. To obtain more information on the characteristics of these two states, we compared seasonal variations in trophic structure and physico-chemical variables of two clear and two turbid lakes studied during seven or eight years. The clearwater lakes were characterised by a high abundance of submerged macrophytes, high piscivorous:planktivorous fish ratios, high zooplankton:phytoplankton ratios and low chlorophyll a during summer. Submerged macrophytes were almost absent from the turbid lakes, planktivorous fish dominated, the zooplankton:phytoplankton ratio was low and summer chlorophyll a was high. While total phosphorus (TP) was almost constant throughout the season in the clearwater lakes, TP was substantially higher during summer in the turbid lakes reflecting high internal loading. In the clearwater lakes, mean summer chlorophyll a was only 45–51% of winter values, while summer chlorophyll a was 118–259% of winter values in the turbid lakes. Our data suggests that zooplankton, by grazing on phytoplankton, play a major role in maintaining clearwater conditions in eutrophic macrophyte-rich lakes, in particular during summer. In addition, results from a multiple regression on data from 37 lakes and the analyses of the seasonal dynamics in suspended solids provide some evidence that zooplankton grazing diminishes concentrations of detritus and inorganic suspended solids either directly by grazing or more indirectly. Using information also from literature, we argue that the role of zooplankton grazing for water clarity in macrophyte-rich lakes may increase from mesotrophic to eutrophic lakes.