Freshwater Biology (2002) 47, 2453–2465

Biomanipulation of lake ecosystems: successful
applications and expanding complexity in the
underlying science
¨ R G E N B E N N D O R F , † P E T E R K A S P R Z A K * and R A I N E R K O S C H E L *
*Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Berlin, Germany
†Dresden University of Technology, Institute of Hydrobiology, Dresden, Germany

1. To illustrate advances made in biomanipulation research during the last decade, seven
main topics that emerged after the first biomanipulation conference in 1989 are discussed
in relation to the papers included in this special issue and the general literature.
2. The substantially higher success rates of biomanipulations in shallow as opposed to
stratified lakes can be attributed to several positive feedback mechanisms relating mainly
to the recovery of submerged macrophytes.
3. The role of both nutrient loading and in-lake concentrations in predicting the success of
biomanipulations is emphasised and supported by empirically defined threshold values.
Nutrient recycling by aquatic organisms (such as fish) can contribute to the bottom-up
effects on lake food webs, although the degree can vary greatly among lakes.
4. Ontogenetic niche shifts and size-structured interactions particularly of fish populations
add to the complexity of lake food webs and make scientifically sound predictions of
biomanipulation success more difficult than was previously envisaged.
5. Consideration of appropriate temporal and spatial scales in biomanipulation research is
crucial to understanding food web effects induced by changes in fish communities. This
topic needs to be further developed.
6. An appropriate balance between piscivorous, planktivorous and benthivorous fishes is
required for long-lasting success of biomanipulations. Recommended proportions and
absolute densities of piscivorous fish are currently based on data from only a few
biomanipulation experiments and need to be corroborated by additional and quantitative
assessments of energy flow through lake food webs.
7. Biomanipulation effects in stratified lakes can be sustained in the long term only by
continued interventions. Alternate stable states of food web composition probably exist
only in shallow lakes, but even here repeated interventions may be needed as long as
nutrient inputs remain high.
8. Biomanipulation is increasingly used as a lake restoration technique by considering the
needs of all lake users (sustainability approach). The combination of water quality
management and fisheries management for piscivores with positive effects for both
appears to be particularly promising.
9. Biomanipulation research has contributed substantially to progress in understanding
complex lake food webs, which should in turn promote a higher success rate of future
whole-lake biomanipulations.

Correspondence: Thomas Mehner, Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Department of Biology and Ecology
of Fishes, POB 850 119, D-12561 Berlin, Germany. E-mail: 
2002 Blackwell Science Ltd


1990. 1998. 1986). Drenner & Hambright. The distinction between shallow and stratified lakes There is widespread consensus that biomanipulation probably has a much higher success rate in shallow than in stratified deep lakes (Gulati et al. niche shifts. 1990) serve as a starting point to illustrate the main topics in biomanipulation research about 12–15 years ago (Table 1). 1988). Scheffer. 1998.. Kitchell & Hodgson. In addition. and findings from a range of other studies that have been published mostly since 1990. 1998) when the proceedings of the first international meeting on the topic appeared (Gulati et al.. the top-down : bottom-up theory (McQueen. we elucidate the recent developments in the new topics in some detail by combining results from papers presented at the symposium in Rheinsberg and included in this special issue. Post & Mills. 1985).. We will point out the progress made during the last decade by relating our conclusions on the current state of biomanipulation research to the synthesis of the 1989 conference (Lammens et al. The focus of the early reviews was on enclosure and laboratory experiments. Drenner & Hambright. and the holistic foodweb model (Persson et al.. Finally. 1990. nutrients Introduction More than 10 detailed reviews on biomanipulation have been published since 1990 (e. 1998. and the limitations of simple food-chain models to explain some of the responses observed in lakes were highlighted. This remarkable activity demonstrates the continuing immense interest in this issue from both a scientific and practical viewpoint. The results of the first international conference on biomanipulation held in Amsterdam in 1989 (Gulati et al. Smith & Slobodkin. the trophic cascade model (Carpenter. More recently. 1999). Some detailed attention was given to species and functional groups that are indirectly affected by biomanipulation. 1975). This suggests that predictions of bioma- nipulation success may require more detailed understanding of interactions within aquatic food webs. 25 of 41 case studies. 1998. Seven partially different themes were identified at the second symposium on biomanipulation held in 2000 in Rheinsberg near Berlin. the potential of biomanipulation for improving lake water quality was assessed based on the available empirical and theoretical evidence (Table 1). management. Hansson et al..g. Hansson et al. which replaced the formerly dominating small-scale two-level experiments.2454 T.. Germany (Table 1). Biomanipulation refers here to the deliberate reduction of planktivory. 1990). 1990). McQueen. The desired reduction of planktivory may be achieved either by removing zooplanktivorous fish manually or by promoting an abundant piscivorous fish community by stocking and protection measures to increase predation pressure on the planktivorous fish. more synthetic view of biomanipulation at the whole-lake scale.. Freshwater Biology. In this review and synthesis paper. This shift was accompanied by and resulted from a large number of whole-lake biomanipulation experiments. 2453–2465 . None of the reviews leaves a doubt that biomanipulation can be an effective and powerful tool for water quality improvement. which is followed by an increase in the abundance and size of zooplankton (predominantly large Daphnia species) and results in increased grazing pressure on phytoplankton and ultimately clearer water of lakes. (1998) noted that most of the successful applications were founded essentially on simple food chain theory (Hairston. and their links with the trophic levels adjacent to them. lakes. Lamarra & Lynch. Benndorf. The main advantage  2002 Blackwell Science Ltd. Keywords: heterogeneity. 1998). maintenance. Hansson et al.. Many studies focused on separate trophic levels. 1960) and its derivatives such as the biomanipulation concept (Shapiro. the ability of shallow lakes to switch between two alternate stable states was clearly expressed. This shift in emphasis is best characterised as an effort towards a broader. interest has shifted to elucidating responses observed after whole-lake manipulations (Hansson et al. The expectation that this simplistic approach works in all situations is in contrast with the abundance of publications highlighting the complexity of aquatic food webs. 47. such as phytoplankton and zooplankton. Mehner et al. The average success rate of food-web manipulations is about 60% (10 of 17 case studies. McQueen. 1999) and only 15% of the whole-lake biomanipulation experiments (6 of 41 studies) reanalyzed by Drenner & Hambright (1999) were considered a definite failure.

Germany. 1990) and the second symposium in Rheinsberg. 47. as summarised in this special issue Amsterdam 1989 Rheinsberg 2000 Edibility of phytoplankton Quality and quantity of phytoplankton Effects of nutrient concentration on phytoplankton community structure – Ecosystem stability Deepwater areas and macrophytes as refugia Alternate stable states of shallow lakes Distinction between shallow and stratified lakes Role of macrophytes Alternate stable states of shallow lakes Indirect effects Invertebrate predators Rotifers Benthivorous fish – phytoplankton link – Biomanipulation and ecosystem research Resource-quality response not predicted by food-chain models Size–structured interactions Stronger role of benthivorous fish Nutrient supply and recycling Biomanipulation efficiency threshold of P-loading Threshold of in-lake P-concentration Trophic state of the lake Nutrient recycling by fish and zooplankton Benthivorous fish The effects of nutrient supply and recycling on the success of biomanipulation Ontogenetic niche shifts and size–structured trophic interactions Ontogenetic niche shifts in piscivores Role of young-of-the-year fish Size refuges of prey against gape-limited predators The importance of ontogenetic niche shifts and size-structured interactions Temporal and spatial heterogeneity in food webs Diel migrations of fish Littoral–pelagic coupling Benthic–pelagic coupling ‘Timing’ of predator–prey interactions The role of temporal variability and spatial heterogeneity in food webs Zooplankton as a key factor Top-down effects of zooplankton related to trophic state Overestimated role of planktivorous fish Greater attention directed to pelagic than benthic zone – – Proportion of piscivores to planktivores ⁄ benthivores Role of predatory invertebrates Balance between piscivores and planktivores and benthivores Importance of omnivory Situation in tropical water bodies  2002 Blackwell Science Ltd. Freshwater Biology..Biomanipulation today – summary and review 2455 Table 1 Summary of the main topics presented at the first biomanipulation conference in 1989 in Amsterdam (synthesis by Lammens et al. 2453–2465 Topic The distinction between shallow and stratified lakes The relative proportion of piscivores and planktivores ⁄ benthivores in the fish community .

perch and pike stocked over a 5-year period induced a decrease in planktivorous and benthivorous fish density. 2002). 1997). 2002). thus contributing to a decreased availability of nitrogen for phytoplankton growth (Van Donk et al. with drastic fish removals and heavy external nutrient loading working antagonistically. McQueen et al. and that an annual loading threshold below 0. a stratified eutrophic hardwater lake. rarely occur when dense macrophyte beds are present suggest that macrophytes compete with phytoplankton successfully for nutrients (Van Donk et al. 2002). 2002) indicate the switching of shallow lakes between the turbid and clear-water states. In Lake Udbyover (Denmark). Similarly. van Nes & Mooij.) and pike (Esox lucius L. (2) the feeding efficiency of predatory fish such as perch (Perca fluviatilis L. Recognition of this positive feedback mechanism has led to the theory of alternate stable states in shallow lakes.. 1981).8 g of total P m)2 of lake surface area must be reached before lake water quality can be improved by biomanipulation (Benndorf et al. 1986.6–0. which in turn expands the water depth and bottom area where macrophytes can grow. Mehner et al. suggest that the improved water quality observed in that lake over the past 10 years can only partly be attributed to biomanipulation. The effects of nutrient supply and recycling on the success of biomanipulation It has long been proposed that bottom-up effects of nutrients on the structure of pelagic food webs remain effective even in strongly top-down manipulated lakes (e. 1990). and (5) resuspension of bottom material is generally lower in macrophyte beds (Barko & Smart. 2453–2465 . Freshwater Biology. Jeppesen et al. (3) experimental and observational results showing that high densities of phytoplankton. 1990) and may excrete allelopathic substances against cyanobacteria (Declerck et al. plankton-dominated state and a clear..g. with rapid shifts occurring between a turbid... (1991) determined an in-lake P-concentration of about 100 lg L)1 as the critical level below which long-term effects of biomanipulation can be expected in shallow lakes.. which eventually led to the re-appearance of submerged macrophytes (Skov et al.. There is also a convincing example that stocking of piscivores without manual removal of planktivores may shift shallow lakes into the clear state. because the improve 2002 Blackwell Science Ltd. Benndorf (1987) suggested therefore that the reduction of nutrient runoff from the catchment may be an important prerequisite for successful biomanipulation. Similarly. 47. Van de Bund & Van Donk. which promote the clear-water state of shallow lakes through a number of mechanisms: (1) Macrophyte beds can act as a refuge for zooplankton from fish predation (Stansfield et al. Grimm & Backx. Shallow lakes in the Netherlands are sensitive to changes in bream biomass (Lammens. 1986). especially cyanobacteria. (4) Conditions inside macrophyte beds may increase denitrification.) in macrophyte beds is higher than that of planktivorous or benthivorous cyprinids such as roach [Rutilus rutilus (L. long-term data from Lake Zwemlust (the Netherlands. Long-term data from Feldberger Haussee..)] or bream [Abramis brama (L. 2000). 1993). macrophyte-dominated state (Scheffer.)] (Winfield. Table 1 (Continued ) Amsterdam 1989 Rheinsberg 2000 Topic Management Assessment of the application potential of biomanipulation Role of nutrient supply and concentration Long-term maintenance of biomanipulation Repeated stocking measures necessary Drastic manual fish removal required Press perturbations necessary Alternative stable states of food-web configuration The requirement of long-term maintenance of biomanipulation measures Management and sustainability Co-operation of all lake users Water quality and sustainable fisheries management The management and sustainability aspects of biomanipulation of food-web manipulations in shallow lakes is the potential for (re)colonisation of large bottom areas by macrophytes.2456 T. All these mechanisms work together towards stabilising or even enhancing water clarity. Germany. 1998).

Biomanipulation today – summary and review ment concurred with a decline in the nutrient loading (Mehner et al. appear not to support the idea that the P input rate must fall below a certain threshold for herbivory to control phytoplankton biomass (Carpenter et al. whereas mesotrophic lakes have four levels with abundant populations of predatory fish. as nutrients were supplied experimentally only during short dosage periods. including experiments involving nutrient additions to lakes. 1990. 2453–2465 2457 A re-discovered effect of biomanipulation is the enhancement of nutrient recycling by aquatic animals.. which exhibit high metabolic rates. the experiments may not have mimicked external loading well.. Faafeng & Nilssen.. when growth rates and P demand of roach and the often very large internal P loading from the sediment (Søndergaard.) stir up the lake bottom and thus increase sediment re-suspension. Tarvainen. Benthivorous fish species such as bream and common carp (Cyprinus carpio L. The removal of cyprinids from Lake Ringsjo¨n. Increasing calcite precipitation and phosphorus coprecipitation may have further contributed to the decline in nutrient concentrations and. but see Attayde & Hansson. A similar argument for stronger top-down control in mesotrophic lakes was derived from experimental work on zooplankton–phytoplankton interactions: Carney (1990) and Elser & Goldman (1991) proposed the ‘mesotrophic maximum hypothesis’ stating that the top-down impact on phytoplankton is highest under mesotrophic conditions. P-recycling by fish may have a low impact on total P availability in most eutrophic lakes. However. recent studies in North America. In contrast. mainly perch. and it has been suggested that the removal of benthivorous fish determines the outcome of biomanipulation in shallow lakes more strongly than the removal of planktivorous fish (Lammens et al. Sweden. (1992). In contrast to the findings above.  2002 Blackwell Science Ltd. the phosphorus accumulation rate of the sediment was probably higher than under equilibrium conditions. In contrast. indicating that the lakes most heavily impacted by nutrient inputs would show the greatest response to a reduction in planktivory. indirectly. (1992) argued that both oligotrophic and eutrophic lakes in Europe possess three-level food chains without piscivores (phytoplankton. in part. Zooplankton biomass did not increase in this lake following biomanipulation (Horppila et al. In addition.. implying that benthivorous fish may exert bottom-up effects on water quality (Tatrai & Istvanovics. resulted in a rapid change in the diversity of benthic invertebrates. 2002). Drenner & Hambright. zooplankton. 1997). suggesting that biomanipulation success was mainly triggered by bottom-up forces induced by intensive fish removal. The trophic state of lakes has an impact on foodchain length. Sarvala & Helminen (2002) calculated that P-release by a roach population could not directly account for the late summer increase in P concentration observed in the biomanipulated Lake Ko¨ylio¨nja¨rvi in Finland. indicating that these fish did indeed disturb .. According to Persson et al. 47. Many effects of fish on phytoplankton that were originally attributed to the feeding of fish on zooplankton can be better explained as indirect effects of nutrient excretion and. 2001) are balanced. 2001). high light attenuation resulting from high concentrations of coloured DOC (up to 9. to the reduction of phytoplankton biomass in that lake (Koschel. Similarly. with a strong top-down influence on planktivorous roach populations. the reduced Precycling after a massive fish kill of the omnivorous Tilapia and Oreochromis in a shallow tropical reservoir contributed to a short-term improvement in water quality (Starling et al. Reduced rates of nutrient recycling resulting from a reduced biomass of planktivorous fish were apparently responsible for the success of biomanipulation in the Finnish Lake Vesija¨rvi. planktivorous fish). Thus. the feeding activity of these fishes may directly destroy or uproot macrophytes. although young fish. Jensen & Jeppesen. Freshwater Biology. water turbidity and internal nutrient loading (Breukelaar et al. 2001).. 1994). 1993. However. Therefore. 1986). dominated the population. the hypotheses above predict that the effectiveness of biomanipulation measures may differ systematically with the trophic state of lakes. 2001). 1990. which leads to an indirect bottom-up influence on pelagic food-web structure. 1998). the absence of piscivorous perch in eutrophic lakes is because of the fact that perch is competitively inferior to roach under the unfavourable food conditions for perch in eutrophic lakes. Schindler et al. egestion by fish (Brabrand.. Sarnelle (1992) found that Daphnia grazing on phytoplankton increased with phosphorus concentration. including fish. 1999).5 units m)1) inhibited the development of inedible phytoplankton in those experiments. In addition. thus delineating the potential for biomanipulation by changes in the upper trophic levels. Persson et al.

The shift from edible to inedible phytoplankton species observed during biomanipulation in response to intense grazing by daphnids is another negative feedback mechanism in foodwebs relating to sizestructure. 1991). they reach a refuge against being fed by predators (Hambright. 1994. 2001. 2002).. The strong predation impact is particularly detrimental for water quality during the summer months (Romare. It has been hypothesised. including predation by (young) fish and predatory invertebrates as well as natural senescence and death (Benndorf et al. Rudstam & Post. Evidence of ontogenetic niches comes from fish communities in both North America (Werner. To overcome this problem. 1998). it is likely that the midsummer decline of daphnids is caused by many interacting mechanisms. Beeck et al. 2000). which are capable of feeding on phytoplankton. 1999). Another mechanism relating to the size-structure of fish communities is the interaction between piscivores and their prey. 1999). young-of-the-year (YOY) fish often increase greatly after several years of biomanipulation (Mehner et al. Bergman & Andersson. Bergman & Hansson. planktivorous species such as common bream or roach can be preyed upon only at younger stages. In addition.2458 T. The importance of ontogenetic niche shifts and size-structured interactions Body size of organisms is important for determining the shape and strength of trophic interactions. 1984). As a result. in particular. Freshwater Biology.. If prey fish can grow to a sufficiently large size. Persson & Eklo¨v. In addition. preventing the top-down control of large planktivores by piscivores (Lammens. 2000). The role of temporal variability and spatial heterogeneity in food webs The differing temporal or spatial scales on which most trophic interactions in lakes occur have received relatively little attention. 1999). 1999).). as these fish also suppress daphnids. and planktivorous YOY fish influenced daph 2002 Blackwell Science Ltd. their stocking is recommended only in geographical regions where daphnids are naturally absent (Radke & Kahl. 1996). Of particular interest for biomanipulation is the diet shift of fish that start their life as planktivores but shift to piscivory when reaching a certain length (e. different developmental stages must be treated separately when such interactions are analysed. Sander lucioperca (L. As these small fish have higher biomass-specific food requirements and occur in high densities. 1999)..)]. The potential growth rate of piscivores depends on the spatial distribution of their prey fish (Mason & Brandt. Mehner et al. perch and pikeperch. Persson. 1996). 2002).g. 1994). although quantitative evidence for this mechanism is limited (Luecke et al. The intensity of fish– zooplankton interactions may vary along spatial gradients in lakes (George & Winfield. 1999). which is often accompanied by reduced water transparency resulting from high phytoplankton biomass. 1990. Establishing a top trophic level of piscivores with these species therefore entails enhanced planktivory by the young (Mehner et al. egg-bearing daphnids (Mehner. has been suggested (Starling et al. stocking lakes with fish such as silver carp [Hypophthalmichthys molitrix (Val. Mehner et al. However. 1998). as the abundance of adult planktivorous fishes decreases. As a result. 47. the lake bottom. their predation impact on zooplankton is higher than that of the same biomass of adult planktivorous fish (Romare & Bergman.. 1992) and Europe (Persson. Quantitative impacts of YOY fish on zooplankton may have rarely been demonstrated because trophic interactions between YOY fish and zooplankton also depend on size relationships: Fish are gape-limited and thus ingest smaller daphnids after hatching than during later growth stages (Schael. 1995). Wagner & Benndorf. 1995). 1996). However. particularly in aquatic habitats where size-structured fish populations dominate (Werner & Gilliam. strong demographic effects in Daphnia populations would only occur when fish feed preferentially on mature. The predation impact by piscivorous perch and pikeperch on their own YOY descendants varied between littoral and pelagic areas (Do¨rner.. with possible implications for pelagic nutrient concentrations and the food web (Svensson. Because single species may show shifts in diet or habitat use during ontogeny. Bystro¨m & Wahlstro¨m.. competition for food diminishes. Hu¨lsmann & Voigt. 2453–2465 . which favours the recruitment of young stages (Romare & Bergman. 2000. 2002). that feeding by YOY fish may cause the commonly observed phenomenon of midsummer decline in Daphnia abundance.. because strong predation may shift the age structure of prey populations towards larger individuals (Bro¨nmark et al..

2001. whereas smaller cladocerans responded positively to the declining fish abundance. 1997). which are active over the whole diurnal cycle (Brabrand & Faafeng. In Lake Mendota. because generation times of different members of pelagic food webs differ widely (Ramcharan et al. food supply. Coupling of the dynamics of organisms differing greatly in regard to generation times may be apparent only over longer time scales. Ho¨lker et al. and population density... Jeppesen et al. 47. A similar effect can be obtained by adding fish kairomones to the water (Gliwicz.A. Planktivorous fish seeking daytime shelter from predation by piscivorous fish or birds either in the deep hypolimnion or in littoral vegetation may lead to a reduction of predatory losses in pelagic zooplankton (Gliwicz & Dawidowicz. acts on state variables such as biomass. Dawidowicz et al. Temporal scales are also important to consider. Such long observations following biomanipulation experiments have not yet been made. (2002) concluded that this dual predation pressure may have been responsible for the extinction of the large Daphnia hyalina Leydig during biomanipulation of the deep Lake Mutek in Poland. 1990.. it is not clear whether mechanisms acting strongly in one particular habitat have a significant impact on the strength of interactions at the whole-lake scale. Consideration of temporal scales was found to be crucial also for understanding the initiation of the midsummer decline of daphnids. new nutrients are supplied to pelagic phytoplankton in a directly usable form.. ageing) determines whether the decline in Daphnia density at the end of spring eventually results in a longer or shorter period in midsummer with extremely low Daphnia abundance (Post & Kitchell. fish may subsidise the pelagic nutrient pool by feeding in nearshore areas and then migrating to the central open-water region (littoral-pelagic coupling) or by feeding at the lake bottom and subsequently moving upwards into the water column (benthicpelagic coupling). In this way.Biomanipulation today – summary and review nids more strongly in littoral than in pelagic areas in the stratified hypertrophic Bautzen Reservoir (Hu¨lsmann et al. 1993. the reproduction rate. and the population growth rate. 2001).. 2453–2465 2459 ited by invertebrate predators such as Chaoborus. Daphnia may be ‘squeezed’ and threatened by predation from both vertebrates in the epilimnion and invertebrates in the hypolimnion. in contrast. However. Time scale considerations also relate to the issue of trophic level control by either predation or resource limitation. Benndorf et al. However. Gliwicz (2002) pointed out that only the bottom-up control is mediated by time-dependent parameters such as the individual growth rate. layers with reduced oxygen concentrations may help daphnids escape fish predation because fish often avoid these zones (Wright & Shapiro. 2002) might induce a behaviourally mediated biomanipulation effect by preventing the daytime feeding of planktivores on daphnids in the open water. In addition. that is at least 50 years in most lakes. the impacts of a massive natural dieoff of a strong cisco (Coregonus artedi Lesueur) year class on plankton community structure and water clarity were detectable for a decade (Lathrop et al.. Hu¨lsmann & Voigt. A structural difference of the trophic cascade within and outside of submerged macrophytes stands has been found in shallow lakes. planktivory can be further suppressed by the presence of pelagic predators such as pikeperch. A similar spatial coupling occurs if fish feed in one area of a lake and excrete in another area. 2002). 1999). Consequently. In both cases.S. stocking with visually oriented pelagic piscivores such as the strongly day-active perch (Jacobsen et al. 2002). U. Consideration of habitat diversity is important also if the physical or chemical characteristics of a water body create spatial refuges that affect the strength of trophic interactions. . Life spans may range from days for phytoplankton to > 20 years for piscivorous fish. 1995). individual body size. 1997. 1992). where the relative timing of all processes involved (predation. Freshwater Biology. if the anoxic hypolimnion is inhab 2002 Blackwell Science Ltd. 2002).... 2002). 1995). Yodzis (1988) suggested that evaluation of the long-term dynamics of whole-lake experiments has to consider time scales twice the sum of the life spans of all members of the trophic chain.. 1998). independent of the rate at which these entities are produced. Top-down control.. For example. resulting in a higher water transparency within macrophyte beds (Schriver et al. Restriction of this fish-mediated nutrient transfer in some cases was found to be more important for the success of biomanipulation than the release of feeding pressure on zooplankton following removal of planktivorous fish (Horppila et al. not over a few months or seasons (Persson et al.

1999. the impact of pulsed perturbations appears to be too small (Persson et al. and many of the studies documented in literature were initiated because of the practical requirement of lake restoration rather than scientific interest (Carpenter & Kitchell.. 1993. 1988) or optimal (Benndorf. Moreover. Mehner et al. 47.. drastic and fast removal of > 75% of the planktivores within one season (Meijer et al. 1997).. requires control by either piscivores or repeated manual removal (Romare & Bergman. what is the most appropriate balance between planktivores and piscivores? Besides the generally desired high species diversity and length variability of the piscivore stock (Benndorf.2460 T. Benndorf & Kamjunke (1999) recommended a proportion of 30–40% piscivores. allowing other groups that prey on daphnids. unpublished data). 2002). Wysujack & Mehner. 2002). However. In contrast. 1999) has been recommended as a measure to shift shallow lakes to the clear-water state (Hansson et al. 1994. 2001). 1992. particularly if the lakes are connected to other water bodies. 1998). indicating that only repeated press perturbations may hold stratified lakes in the desired food-web structure. 2002). whether such critical fish densities are lake-specific or can be generally defined. 1998). in  2002 Blackwell Science Ltd.. Biomanipulation by piscivore stocking had the lowest success rate among a range of measures evaluated by Drenner & Hambright (1999).. 2002) and up to 50% piscivores (Bautzen Reservoir. Freshwater Biology. If planktivorous fish cannot and should not be completely removed from biomanipulated lakes. Van de Bund & Van Donk. The relative proportion of piscivores and planktivores ⁄ benthivores in the fish community As biomanipulation requires drastic removal of planktivorous fish. Benndorf et al. Detailed energetic balances supporting these empirical values are currently lacking. McQueen. the increased recruitment success of planktivores following a period of planktivorous fish removals. Prevention of immigration may be especially complicated if the connections are used for navigation (Perrow et al. 1999). Perrow et al.. Moreover... It is not yet clear. such as predatory invertebrates (Wissel et al. Benndorf & H. 1993). This recommendation is based on experimental evidence from long-term biomanipulations in German lakes showing that biomanipulation is successful when the fish community consists of >20–25% (Feldberger Haussee. 1992). there is no evidence at present that alternative stable states also occur in stratified lakes (Persson.. however.. 1990) absolute densities of planktivorous fish may be an even better endpoint for fish reductions than the relative removal rates suggested by Meijer et al. The requirement of long-term maintenance of biomanipulation measures Biomanipulation probably does not lead to stable food web configurations and therefore requires continuous efforts to suppress planktivores (Kitchell. For example. in the stratified Feldberger Haussee. to invade and fill the gap. Some critical (Barthelmes. J. 2000). (1999) and others. This conclusion may hold particularly true if alternative stable states of food web configurations exist. The drastic reductions in planktivore fish stocks normally required for successful biomanipulation indicate that press perturbations are required to shift a system into another state. Given that even shallow lakes may show little sign of long-term stability of biomanipulation effects (Perrow et al. 1998). 1990. repeated fish stock reductions have been proposed as a simple and cost-effective management strategy as long as external nutrient supplies have not been markedly reduced (Van De Bund & Van Donk. planktivorous fish may rapidly invade biomanipulated lakes. the question may arise why not create a fish community composed of piscivores only? Intentional changes of both the piscivorous and planktivorous levels in a whole-lake biomanipulation experiment have indicated that the goal of removing the planktivorous level from a lake totally may be impossible to achieve in practice (Wissel et al. as has been proposed for shallow lakes (Scheffer. 2453–2465 .. Therefore. Do¨rner. 1997). the continuous removal of planktivores by low-intensity fishing did not result in a long-term decline of the planktivorous fish stock and did not greatly enhance crustacean biomass (Kasprzak et al. however. total removal of planktivorous fish would leave the planktivore niche unoccupied. Similarly. Even repeated stocking of piscivores may not be sufficient without manual removal of planktivores. Mehner et al. The management and sustainability aspects of biomanipulation Biomanipulation is used to improve the water quality of lakes. 2000).

. 1998.S. boundary conditions such as lake depth. which in turn enabled the anglers to catch larger specimens than before (Lathrop et al. recent studies. underwater light climate.. planktivorous and benthivorous fish and their respective prey  2002 Blackwell Science Ltd. 3. Benndorf & Kamjunke. during recent years.. 2. Lathrop et al. The confounding role of nutrients in determining the potential for successful biomanipulations is now clearer. The consideration of scales is important for understanding trophic interactions along the temporal (diel. However. The scientific data and expertise obtained in biomanipulation experiments in a range of lake types serve as the basis for designing appropriate fish manipulation measures in particular water bodies. 1996. although they can mirror major food-web changes in response to press perturbations. Therefore. quantitative approaches to assess the top-down and bottom-up effects of fish are increasingly used (bioenergetics. 1999. 2001). The scientific background of biomanipulation. In combination with conclusion 1. seasonal. ontogeny) axes. often neglected in earlier studies. 1990. anglers and scientists was part of the planning and manipulation phases (e. 2002). size and bag limits imposed on recreational fishermen were one of the most important restrictions to support biomanipulation. Thus. 2453–2465 2461 groups play a central role in biomanipulation. as compared with the findings until 1989 (Table 1). mixing intensity and trophic state govern the response of complex food webs to changes in predation strength. in addition to generating income for the local fishermen (Starling et al. In addition. Most lake managers currently learn by doing. is now fully acknowledged as these fish are the dominant zooplankton feeders in many lakes.. 2002).Biomanipulation today – summary and review many biomanipulations. increasing interest in catching living bream for stocking of angling waters allows Dutch fishermen to obtain income by combining the desired decline in bream stocks with the effect of enhanced water transparency (Lammens et al. this affirmation is a partial revival of the top-down : bottom-up concept by McQueen et al. Freshwater Biology.. For example. 1992.. using biomanip- . 47. in Lake Mendota. Madgwick & Phillips. Evidence from a reservoir in tropical Brazil suggests that dense populations of omnivorous tilapias increase nutrient concentrations and promote algal blooms as a result of P-recycling and bioturbation. detailed insights into the trophic interactions within pelagic food webs are necessary. (1986). success was often reported in those cases where authorities and lake users supported the biomanipulation (Horppila et al. Kitchell. 1999) and the combination of water quality management and sustainable fisheries management by biomanipulation has received increasing attention during the last years (Lammens. The main progress can be summarised as follows: 1. which predicts the strongest effect of biomanipulation on pelagic food webs when both fish and nutrients are altered. 2002). Guidelines for biomanipulating lakes have been developed (Moss. the co-operation of lake and fishery managers. a professional tilapia cast-net fishery has been established to control the population density of fish and thus limit the occurrence of cyanobacterial blooms and fish kills. for the fine-tuning of ongoing and for predicting success of future biomanipulations. Perrow et al. and more precisely described in. The importance of young-of-the-year fish. and for finetuning ongoing biomanipulations. particularly in North America and parts of Europe where fisheries favour piscivorous species as catch. Besides scales. spatial (habitat coupling.. interannual). lake and catchment relationships). and interorganismic (size structure. Even in tropical countries there is potential to support water-quality goals by stocking with piscivores and controlled fishery.g. 4. has become even more complex than was conceived after the 1989 conference. Wysujack et al. lake food-web ecology. In addition. temperature regime. Conclusion A generally better understanding of the complexity of lake food webs and the frequent application of biomanipulation as a lake restoration technique have emerged from the recent developments in biomanipulation research. simple food-chain models cannot be realistically assumed to reflect the multitude of possible effects of biomanipulations. The thresholds for nutrient supply and concentrations in lakes suggested in earlier studies have been confirmed by. Interactions between piscivorous. U. Benndorf. In contrast to the situation 10–15 years ago.. Biomanipulation can be regarded as an approach to water quality improvement that requires the combination of research and management. 2002).A. individual-based modelling). 1999).

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