Changes in phosphorus and nitrogen cycling following food web manipulations in a shallow Dutch lake

1. The flow of phosphorus and nitrogen through the food web of the shallow, eutrophic lake Wolderwijd was analysed for 2 different years before and for 1 year after food web manipulation.
2. After fish removal the water became clear and the growth of macrophytes began. Fish removal resulted in a significant reduction of the total nutrient pool in the water, but differences between the nutrient cycles before and after the experiment were mainly caused by a gradual change driven by a reduced phosphorus input.
3. The zooplankton biomass before and after food web manipulation did not change significantly. Unfavourable food conditions and predation by young fish limited zooplankton biomass after the food web manipulation.
4. After fish removal benthic algae, fish, zoobenthos and macrophytes form the largest pools of nutrients apart from the sediment top layer. However, they contribute only little to nutrient cycles in the water column.     

Modelling nutrient cycles in relation to food web structure in a biomanipulated shallow lake

The modelPCLAKE describes the phosphorus and nitrogen cycles within a shallow lake ecosystem, including the sediment and a simplified biological food web. All components are modelled in a generalized way rather than a very detailed one. This model has been applied to Lake Zwemlust, a small biomanipulated lake in The Netherlands. Formerly, this highly eutrophic lake was dominated by cyanobacteria and devoid of macrophytes. Biomanipulation was carried out in 1987 by pumping-out of the water, removal of all fish, and refilling of the lake with seepage water. The lake was restocked with some rudd, pike, zooplankton and seedlings of macrophytes, and then monitored up to 1992. Macrophytes developed rather quickly and reached their maximum biomass during the six-years period in 1989. Despite the continuously high nutrient (N and P) loading, algal biomass remained low due to nitrogen limitation, caused by competition with the macrophytes. From 1990 onwards, the macrophytes declined again and a species shift occurred, following an increase of herbivorous birds on the lake and the development of herbivorous fishes.Model simulations grossly reproduced the observed developments in Lake Zwemlust before and after the biomanipulation measures. The existence of multiple steady states at the same trophic state and the possible shift between them could be simulated well. This study also demonstrates the interrelation between system structure and the distribution and cycling of nutrients. It is concluded, that within general boundary conditions set by the trophic state of the system, the food web structure determines the actual nutrient flows and the occurrence of nutrient limitations of the primary producers. It is shown that both aspects can be integrated in one mathematical model. The long-term stability of the macrophyte dominance in the lake is discussed.     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Changes in phosphorus cycling in a shallow lake due to food web manipulations

SUMMARY. 1. Food web manipulation, by removal of planktivorous or benthivorous fish, is a promising method for reducing phytoplankton concentrations in shallow lakes. The part that nutrients may play in the success of such a measure is not well documented.
2. In this study, we analysed the flow of phosphorus through the food web of the shallow, eutrophic Lake Wolderwijd/Nuldernauw. Our studies occurred in the years 1981 (when a bloom of cyanobacteria occurred) and 1987 (no bloom); a hypothetical situation was also examined in which most of the bream are assumed to be removed.
3. The analysis shows that the success of biomanipulation is probably due not only to an increased grazing pressure on the phytoplankton, but also to a decreased availability of phosphorus. The reason for this is the removal of detrital phosphorus by increased sedimentation as a result of a predicted increase in growth of macrophytes after biomanipulation.     

Top-down or Bottom-up Effects by Fish: Issues of Concern in Biomanipulation of Lakes

A large-scale biomanipulation trial was carried out on Lake Vesijärvi in Finland during 1989–1993. Following the mass removal of coarse fish the biomass of cyanobacteria collapsed from 1.4 g/m^?3 to below 0.4 g/m^?3, while total phosphorus concentration declined from 45 μ g/L to 30 μ g/L. No relevant changes in zooplankton communities were observed. The results suggest that the success of food web manipulation as a tool for lake restoration is not necessarily dependent on the grazing rate of zooplankton. The effects of reduced fish-mediated internal loading and recycling of nutrients are in many cases stronger than those of reduced planktivory. Alternative stable states of water quality may also exist in lakes not covered by macrophytes, owing to the changes in the behavior of fish stocks. Year-to-year variation in the littoral zone may cause large oscillations in lake ecosystems—for example, through the recruitment of fish. In addition, the nutrients translocated by fish from the littoral zone may affect the nutrient dynamics of the pelagial plankton community. In terms of phytoplankton species composition and the ratio of phosphorus to chlorophyll a, the water quality in Lake Vesijärvi has improved in a stepwise fashion within the last 10 years. This is probably due to the fact that the five-year mass removal of fish in Enonselkä fulfilled the requirement of sustained management of fish stocks in order to maintain nonequilibrial conditions between alternate stable states. The prediction of the water quality development is obscured, however, by spatial and temporal within-lake variation, which sets high requirements for sampling programs.     

OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

Phytoplankton food quality control of planktonic food web processes

We developed a mechanistic model of nutrient, phytoplankton, zooplankton and fish interactions to test the effects of phytoplankton food quality for herbivorous zooplankton on planktonic food web processes. When phytoplankton food quality is high strong trophic cascades suppress phytoplankton biomass, the zooplankton can withstand intense zooplanktivory, and energy is efficiently transferred through the food web sustaining higher trophic level production. Low food quality results in trophic decoupling at the plant-animal interface, with phytoplankton biomass determined primarily by nutrient availability, zooplankton easily eliminated by fish predation, and poor energy transfer through the food web. At a given nutrient availability, food quality and zooplanktivory interact to determine zooplankton biomass which in turn determines algal biomass. High food quality resulted in intense zooplankton grazing which favored fast-growing phytoplankton taxa, whereas fish predation favored slow-growing phytoplankton. These results suggest algal food quality for herbivorous zooplankton can strongly influence the nature of aquatic food web dynamics, and can have profound effects on water quality and fisheries production. Handling editor: D. Hamilton     

Pathways of Increased Water Clarity After Fish Removal from Ventura Marsh; a Shallow, Eutrophic Wetland

We investigated the pathways by which water clarity increases following fish removal by evaluating the effects of a benthivorous fish reduction in a large, shallow, eutrophic, wetland in a predominately agricultural watershed in Iowa, U.S.A. Phytoplankton was phosphorus limited prior to manipulation. After a substantial fish removal was obtained, water clarity increased as a result of decreased suspended sediment and phytoplankton biomass. Trophic cascading, mitigated by release from fish predation and decreased physical interference from suspended sediments, appears to determine water clarity. Inorganic suspended solids declined immediately after fish were removed but the biomass of Daphnia and Ceriodaphnia did not increase until a few weeks after fish removal. High grazing by zooplankton likely reduced phytoplankton biomass during the height of the clear-water phase. Phytoplankton appeared to be limited by zooplankton grazing for approximately two months before reverting to bottom-up control. An increase in suspended sediment and/or increased predation pressure on zooplankton, due to the return of juvenile carp, appears to account for the decline of larger-bodied zooplankters and the switch back to bottom-up control. Macrophyte diversity and density increased substantially after the initiation of the clear-water phase.     

Stable isotope analysis of archived roach (Rutilus rutilus) scales for retrospective study of shallow lake responses to nutrient reduction

1. There is increasing interest in the use of stable isotope analysis of archived materials to study the long-term impacts of lake perturbations, including nutrient manipulation or species invasion. We tested the utility of this approach in a shallow productive lake using the zooplanktivorous early life stages of roach ( Rutilus rutilus ), a fish species that is widespread throughout Eurasian lakes.
2. Barton Broad is a shallow lake with a well-documented history of earlier eutrophication followed by nutrient reduction, including sediment removal from 1997 to 2000. Using scale samples collected pre- and post-sediment removal, we demonstrated a strong, positive relationship between roach scale δ ¹³C and total phosphorus. We argue that this reflects a decrease in the phytoplankton production which had dominated dissolved inorganic carbon dynamics, and a relative increase in the contribution of respired carbon in the food web.
3. We also derived a scale : muscle isotope relationship for roach which allowed us to model changes in fish muscle against putative prey. Concomitant isotopic shifts in preserved zooplankton samples indicated that the phosphorus reduction measures had an ecosystem-wide impact and that changes in roach scale isotope values were not a result of fish switching diet.
4. Roach scale δ ¹⁵N increased after sediment removal. Since this was not due to a switch in fish diet, we suggest that it probably reflects the loss of nitrogen-fixing, heterocystous cyanobacteria from the plankton.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

Facilitation of clear-water conditions in shallow lakes by macrophytes: differences between charophyte and angiosperm dominance

A number of mechanisms result in a feedback between water clarity and macrophytes and, consequently, the occurrence of alternative stable states in shallow lakes. We hypothesize that bottom-up mechanisms and interactions within the benthic food web are more important in a charophyte-dominated clear-water state, while top-down mechanism and interactions in the planktonic food web prevail at angiosperm dominance. Charophytes, which dominate at lower nutrient concentrations and develop higher densities than most angiosperms, can have a higher influence on sedimentation, resuspension, and water column nutrients. During dominance of dense submerged vegetation like charophytes, zooplankton can be hampered by low food quality and quantity and by high predation pressure from juvenile fish, which in turn are favoured by the high refuge potential of this vegetation. Grazing pressure from zooplankton on phytoplankton can therefore be low in charophytes, but the main feedback in angiosperm-dominated ecosystems. Charophytes offer a higher surface than most angiosperms to periphyton, which favors benthic invertebrates. These support macrophytes by grazing periphyton and constitute a central link in a trophic cascade from fish to periphyton and macrophytes. To test these hypotheses, more experiments and field measurements comparing the effect of charophytes and angiosperms on water clarity are needed.     

Restoration by biomanipulation of lake Bleiswijkse Zoom (The Netherlands): First results

In 1987, the Bleiswijkse Zoom, a small, shallow lake in The Netherlands, was divided into two compartments to investigate the possible use of biomanipulation as a tool for restoring the water quality of hypertrophic lakes. The density of the fish stock before restoration was about 650 kg.ha^–1, composed mainly of bream, white bream and carp. Pikeperch was the main fish predator in the lake. In April, 1987, in one compartment (Galgje) all planktivorous bream and white bream and about 85% of the benthivorous bream and carp were removed. Advanced pikeperch fry were introduced as predator during the transient period. The other compartment (Zeeltje) was used as a reference. Removal of the fish in Galgje resulted in low concentrations of chlorophyll-a, total phosphorus, nitrogen and suspended solids. The absence of bottom-stirring activity by benthivorous fish and the low chlorophyll-a concentrations led to an increase in the Secchi disk transparency from 20 to 110 cm. Within two months after removal of the fish, macrophytes, mainly Characeae, became abundant. Until July the high density of large zooplankton species caused low algal biomass. From June onwards, the zooplankton densities decreased, but the algal concentrations remained low. This is probably because of nutrient limitation or depression of algal growth by macrophytes or both. Compared with the non-treated compartment the number of fish species in the treated compartment was higher. Perch, rudd and roach, i.e. the species associated with aquatic vegetation, were found in the samples. The survival of the O⁺ pikeperch was poor. The pikeperch could not prevent the growth of young cyprinids. Within two months after the removal of the fish a habitat for northern pike was created.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Trophic structure, species richness and biodiversity in Danish lakes: changes along a phosphorus gradient

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.     

Response of a shallow Mediterranean lake to nutrient diversion: does it follow similar patterns as in northern shallow lakes?

1. In view of the paucity of data on the response of warm shallow lakes to reductions in nutrient loading, this paper presents a long‐term limnological data set to document changes in the food‐web of a shallow Mediterranean lake (Lake Albufera, Valencia, Spain) that has experienced reductions in phosphorus (P) (77%) and nitrogen (N) (24%) loading following sewage diversion. 2. Nine years after sewage diversion, P concentration in the lake was reduced by 30% but remained high (TP = 0.34 mg L^?1), although the mean water retention time in the lake was only 0.1 years. Nitrate concentrations did not significantly change, probably because the lake continued to receive untreated effluents from ricefields. 3. Chlorophyll a concentration was reduced by half (annual mean of 180 μg L^?1). Cyanobacteria abundance remained high but its composition changed towards smaller species, both filamentous and chroococcal forms. 4. Cladocera abundance increased and reached peaks twice a year (December to March and July to September). After nutrient reduction, short‐term clear‐water phases (up to 5 weeks) occurred during February to March in several years, concomitant with annual flushing of the lake and lower fish densities. The abundance of Cladocera in winter contrasted with the spring peaks observed in northern restored shallow lakes. The zooplankton to phytoplankton biomass ratio remained lower than in northern temperate shallow lakes, probably because of fish predation on zooplankton. 5. Improvement of the water quality of Lake Albufera remained insufficient to counteract littoral reed regression or improve underwater light allowing submerged plants re‐colonise the lake. 6. Sewage diversion from Lake Albufera impacted the food web through the plankton, but higher trophic levels, such as fish and waterfowl, were affected to a lesser degree. Although the fish species present in the lake are mainly omnivorous, long‐term data on commercial fish captures indicated that fish communities changed in response to nutrient level and trophic structure as has been observed in restored shallow lakes at northern latitudes. 7. Phosphorus concentrations produced similar phytoplankton biomass in Lake Albufera as in more northern shallow lakes with abundant planktivorous fish and small zooplankton. However, in Lake Albufera, high average concentrations were maintained throughout the year. Overall, results suggest that nutrient control may be a greater priority in eutrophicated warm shallow lakes than in similar lakes at higher latitudes.     

A 2-year experimental study on nutrient and predator influences on food web constituents in a shallow lake of north-west Spain

1. A 2‐year study was carried out on the roles of nutrients and fish in determining the plankton communities of a shallow lake in north‐west Spain. Outcomes were different each year depending on the initial conditions, especially of macrophyte biomass. In 1998 estimated initial ‘per cent water volume inhabited’ (PVI) by submerged macrophytes was about 35%. Phytoplankton biomass estimated as chlorophyll a was strongly controlled by fish, whereas effects of nutrient enrichment were not significant. In 1999 estimated PVI was 80%, no fish effect was observed on phytoplankton biomass, but nutrients had significant effects. Water temperatures were higher in 1998 than in 1999. 2. In the 1998 experiment, cladoceran populations were controlled by fish and cyanobacteria were the dominant phytoplankton group. There were no differences between effects of low (4 g fresh mass m^?2) and high (20 g fresh mass m^?2) fish density on total zooplankton biomass, but zooplankton biomass was higher in the absence of fish. With the high plant density in 1999, fish failed to control any group of the zooplankton community. 3. Total biovolume of phytoplankton strongly decreased with increased nutrient concentrations in 1998, although chlorophyll a concentrations did not significantly change. At higher nutrient concentrations, flagellate algae became more abundant with likely growth rates that could have overcompensated cladoceran feeding rates. This change in phytoplankton community composition may have been because of increases in the DIN : SRP ratio. Both chlorophyll a concentration and total phytoplankton biovolume increased significantly with nutrients in the 1999 experiment. 4. A strong decline of submerged macrophytes was observed in both years as nutrients increased, resulting in shading by periphyton. This shading effect could account for the plant decline despite lower water turbidity at the very high nutrient levels in 1998.     

Nutrient management and structural shifts in fish assemblages: Lessons learned from an Area of Concern in Lake Ontario

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Community resistance and change to nutrient enrichment and fish manipulation in a vegetated lake littoral

1. High biomass of macrophytes is considered important in the maintenance of a clear‐water state in shallow eutrophic lakes. Therefore, rehabilitation and protection of aquatic vegetation is crucial to the management of shallow lakes. 2. We conducted field mesocosm experiments in 1998 and 1999 to study community responses in the plant‐dominated littoral zone of a lake to nutrient enrichment at different fish densities. We aimed to find the threshold fish biomass for the different nutrient enrichment levels below which large herbivorous zooplankton escapes control by fish. The experiments took place in the littoral of Lake Vesijärvi in southern Finland and were part of a series of parallel studies carried out jointly at six sites across Europe. 3. In 1998, when macrophyte growth was poor, a clear‐water state with low phytoplankton biomass occurred only in unenriched mesocosms without fish or with low fish biomass (4 g fresh mass m^?2). Both nutrient enrichment and high fish biomass (20 g fresh mass m^?2) provoked a turbid water state with high planktonic and periphytic algal biomass. The zooplankton community was dominated by rotifers and failed to control the biomass of algae in nutrient enriched mesocosms. The littoral community thus had low buffer capacity against nutrient enrichment. 4. In 1999, macrophytes, especially free‐floating Lemna trisulca L., grew well and the zooplankton community was dominated by filter‐feeding cladocerans. The buffer capacity of the littoral community against nutrient enrichment was high; a clear‐water state with low phytoplankton biomass prevailed even under the highest nutrient enrichment. High grazing rates by cladocerans, together with reduced light penetration into the water caused by L. trisulca, were apparently the main mechanisms behind the low algal biomass. 5. Effects of fish manipulations were less pronounced than effects of nutrient enrichment. In 1999, clearance rates of cladocerans were similar in fish‐free and low‐fish treatments but decreased in the high‐fish treatment. This suggests that the threshold fish biomass was between the low‐ and high‐fish treatments. In 1998, such a threshold was found only between fish‐free and low‐fish treatments. 6. The pronounced difference in the observed responses to nutrient enrichment and fish additions in two successive years suggests that under similar nutrient conditions and fish feeding pressure either clear or turbid water may result depending on the initial community structure and on weather.