Effect of larval fish and nutrient enrichment on plankton dynamics in experimental ponds

The hypotheses that larval fish density may potentially affect phytoplankton abundance through regulating zooplankton community structure, and that fish effect may also depend on nutrient levels were tested experimentally in ponds with three densities of larval walleye, Stizostedion vitreum (0, 25, and 50 fish m^–3), and two fertilizer types (inorganic vs organic fertilizer). A significant negative relationship between larval fish density and large zooplankton abundance was observed despite fertilizer types. Larval walleye significantly reduced the abundances of Daphnia, Bosmina, and Diaptomus but enhanced the abundance of various rotifer species (Brachionus, Polyarthra, and Keratella). When fish predation was excluded, Daphnia became dominant, but Daphnia grazing did not significantly suppress blue-green algae. Clearly, larval fish can be an important regulator for zooplankton community. Algal composition and abundance were affected more by fertilizer type than by fish density. Inorganic fertilizer with a high N:P ratio (20:1) enhanced blue-green algal blooms, while organic fertilizer with a lower N:P ratio (10:1) suppressed the abundance of blue-green algae. This result may be attributed to the high density of blue-green algae at the beginning of the experiment and the fertilizer type. Our data suggest that continuous release of nutrients from suspended organic fertilizer at a low rate may discourage the development of blue-green algae. Nutrient inputs at a low N:P ratio do not necessarily result in the dominance of blue-green algae.     

Trophic changes,without changes in the external nutrient loading

The impact of the fish population on trophic properties of lake water, was experimentally studied in an oligotrophic Swedish forest lake. Biotic changes following fish removal resulted in a development in oligotrophic direction as shown by the drop in limnetic primary production, pH, total phosphorus, total nitrogen and the increased transparency.     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.     

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.     

Effects of different fish species and biomass on plankton interactions in a shallow lake

Thirty-six enclosures, surface area 4 m², were placed in Little Mere, a shallow fertile lake in Cheshire, U.K. The effects of different fish species (common carp, common bream, tench and roach) of zooplanktivorous size, and their biomass (0, 200 and 700 kg ha^–1) on water chemistry, zooplankton and phytoplankton communities were investigated. Fish biomass had a strong effect on mean zooplankton size and abundance. When fish biomass rose, larger zooplankters were replaced by more numerous smaller zooplankters. Consequently phytoplankton abundance rose in the presence of higher densities of zooplanktivorous fish, as zooplankton grazing was reduced. Fish species were also significant in determining zooplankton community size structure. In enclosures with bream there were significantly greater densities of small zooplankters than in enclosures stocked with either carp, tench and, in part, roach. When carp or roach were present, the phytoplankton had a greater abundance of Cyanophyta than when bream or tench were present. Whilst top-down effects of fish predation controlled the size partitioning of the zooplankton community, this, in turn apparently controlled the bottom-up regeneration of nutrients for the phytoplankton community. At the zooplankton–phytoplankton interface, both top-down and bottom-up processes were entwined in a reciprocal feedback mechanism with the extent and direction of that relationship altered by changes in fish species. This has consequences for the way that top-down and bottom-up processes are generalised.     

Predictability and possible mechanisms of plankton response to reduction of planktivorous fish

The predictability of plankton response to reductions of planktivorous fish was investigated by comparing the plankton community in three biomanipulated lakes and ten unmanipulated lakes differing in intensity of fish predation. Data collected on total phosphorus, phytoplankton and zooplankton biomass and share of cyanobacteria and large grazers, as well as specific growth rate of phytoplankton, were further used to test some of the proposed underlying response-mechanisms. In the biomanipulated lakes the algal biomass and share of cyanobacteria decreased, specific growth rate of phytoplankton increased, and zooplankton biomass and share of large grazers increased or remained unchanged. This pattern was largely reflected in the differences in food-chain structure between the unmanipulated lakes with highversus those with low fish predation. The qualitative response to planktivorous fish reduction thus seems largely predictable. The biomanipulated lakes differed, however, in magnitude of response: the smallest hypertrophic, rotenone-treated lake (Helgetjern) showed the most dramatic response, whereas the large, deep mesotrophic lake (Gjersjøen), which was stocked with piscivorous fish, showed more moderate response, probably approaching a new steady state. These differences in response magnitude may be related to different perturbation intensity (rotenone-treatmentversus stocking with piscivores), food-chain complexity and trophic state. Both decreased phosphorus concentration and increased zooplankton grazing are probably important mechanisms underlying plankton response to biomanipulation in many lakes. The results provide tentative support to the hypothesis that under conditions of phosphorus limitation, increased zooplankton grazing can decrease algal biomassvia two separate mechanisms: reduction of the phosphorus pool in the phytoplankton, and reduction of the internal C:P-ratio in the phytoplankton cells.

     

Early responses of plankton and turbidity to biomanipulation in a shallow prairie lake

We evaluated the effect of a fish removal from a shallow, turbid, eutrophic lake. By late May (following an October fish removal), the cladoceran community shifted from small-bodiedBosmina andChydorus (less than 100 l⁻¹) to largerDaphnia (over 100 l⁻¹). During the periods of peak daphnid abundance (late May–June) chlorophyll-a concentrations and edible diatoms were reduced and water transparency improved dramatically. Total phosphorus was not significantly lowered during this period. Although this clear-water phase was short-lived (May, June and early July), it corresponded to the critical period of plant growth and allowed dramatic increases in submergent macrophytes.     

Long-term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Müggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Müggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N₂‐fixing species Aphanizomenon flos‐aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re‐colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Müggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.     

Effects of fertiliser and crayfish on plankton and nutrient dynamics in hardwater ponds

Although phosphorus fertilisation can improve productivity in most freshwater ponds, phosphate may become limiting in extremely hard water due to rapid precipitation with calcium. Hence we studied the characteristics of plankton and nutrient dynamics in water containing >400 mg CaCO₃ l^–1in pond and microcosm systems. The field experiment was conducted in eight earthen ponds involving two nutrient ratios (N:P = 1:1 and 20:1) with or without crayfish. Fertilisation significantly increased concentrations of NO₂–N and NO₃–N, but soluble reactive phosphorus was depleted to the level prior to fertilisation within 24 h. The laboratory test showed that after 6 h of fertilisation, 45% phosphorus was precipitated by calcium, 30% phosphorus was assimilated by phytoplankton and only 25% phosphorus remained in water column. The phytoplankton abundance in hardwater ponds was regulated by the abundance of zooplankton population rather than by either crayfish or fertilisation. The presence of crayfish only increased the concentration of total phosphorus. This study suggests that when phytoplankton production is required in crayfish ponds the maintenance of phytoplankton abundance will depend on the effective control of zooplankton rather than fertilisation. Due to the rapid precipitation of phosphorus by calcium in hard water ponds, more frequent phosphorus fertilisation is needed to enhance primary productivity.     

Responses to fish predation and nutrients by plankton at different levels of taxonomic resolution

1. To improve mechanistic understanding of plankton responses to eutrophication, a mesocosm experiment was performed in the shallow littoral zone of a south Swedish lake, in which nutrient and fish gradients were crossed in a fully factorial design. 2. Food chain theory accurately predicted total biomass development of both phyto‐ and zooplankton. However, separating zooplankton and algae into finer taxonomic groups revealed a variety of responses to both nutrient and fish gradients. 3. That both nutrients and fish are important for phytoplankton dynamics was seen more clearly when viewing each algal group separately, than drawing conclusions only from broad system variables such as chlorophyll a concentration or total phytoplankton biovolume. 4. In some taxa, physiological constraints (e.g. sensitivity to high pH and low concentrations of free CO₂) and differences in competitive ability may be more important for the biomass development than fish predation, grazing by herbivorous zooplankton, and nutrient availability. 5. We conclude that food chain theory accurately predicted responses in system variables, such as total zooplankton or algal biomass, which are shaped by the dynamics of certain strong interactors (‘keystone species’), such as large cladocerans, cyanobacteria and edible algae (<50 μm), whereas responses at finer taxonomic levels cannot be predicted from current theory.     

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 submerged macrophytes in Danish lakes to nutrient loading reductions and biomanipulation

During the last two decades the nutrient loading to Danish lakes has been reduced with the aim to improve water quality. However, because of internal P-loading and biological resistance, the expected improvement has been delayed. Therefore, to reduce the duration of the recovery period and to accelerate recolonisation of submerged macrophytes, several lakes have been biomanipulated with the purpose of improving the top-down control by zooplankton. To elucidate the effects of these measures, we undertook an analysis of data on submerged macrophytes monitored annually in 17 lakes for 8 years. The results obtained show that the macrophyte coverage in non-biomanipulated lakes remained relatively stable following the external nutrient reduction. However, a small increase in macrophyte coverage occurred in a few lakes. In two of the four biomanipulated lakes, in contrast, macrophyte coverage increased from 0 up to 80% within 2–4 years following manipulation. In the other two lakes macrophyte colonisation failed. However, in lakes with a successful recolonisation, large inter-annual variations in macrophyte coverage, varying between 2 and 80% among growth seasons, occurred. We conclude that the potential of macrophyte recolonisation after nutrient loading reduction on the short term is higher in biomanipulated lakes than in lakes subjected to loading reduction only, although biomanipulation does not provide a guarantee for macrophyte recolonisation or a stable macrophyte community when colonisation occurs.

     

Phytoplankton biomass reduction after planktivorous fish reduction in a shallow,eutrophic lake: a combined effect of reduced internal P-loading and increased zooplankton grazing

Hydrobiologia - No recovery was recorded in the shallow and eutrophic Lake Væng, Denmark, after a sewage diversion in 1981, due to an internal phosphorus loading and a dominance of...     

Combined stable isotope and gut contents analysis of food webs in plant-dominated, shallow lakes

Summary 1. To determine feeding links between primary producers, invertebrates and fish, stable isotope analyses and gut content analyses of fish were conducted on the components of four shallow, eutrophic to hypertrophic, plant-dominated lakes.
2. Although separation of basal resources was possible, the diets of both fish and invertebrates were broad, comprising food from different compartments (planktonic, epiphytic/benthic), as well as from different trophic levels.
3. Mixing models were used to determine the extent to which periphyton production supported higher trophic levels. Only one species of invertebrate relied upon periphyton production exclusively.
4. Fish density affected the diets of invertebrates. The response was different for planktonic and epiphytic/benthic invertebrates. The proportion of periphyton production in the diets of zooplankton appeared to increase with fish density, whilst it decreased for other invertebrates.
5. As all zooplankton samples were collected in the open water at dusk, these results are further evidence for the diurnal horizontal migration of zooplankton. Although not conclusive, they are consistent with a behavioural response by invertebrates and zooplankton in the presence of fish.     

Relations between planktivorous fish abundance,zooplankton and phytoplankton in three lakes of differing productivity

Hydrobiologia - Water chemistry, phytoplankton, zooplankton and fish populations werestudied over several years in three shallow, non-stratified lakes withdiffering nutrient loadings and fish...     

Ecological effects of reduced nutrient loading (oligotrophication) on lakes: an introduction

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

Effect of young-of-the-year walleye (Percidae: Stizostedion vitreum) on plankton dynamics and water quality in ponds

To understand the impact of young-of-the-year (YOY) fish on food web dynamics and water quality, we stocked larval walleye (9 mm TL) (Stizostedion vitreum) in six experimental ponds using two fish densities (10 and 50 fish m^–3) with three replicates. At high fish density, the average abundances of cladocerans and copepods and the Secchi depth were lower whereas abundances of rotifers and algae, gross primary productivity (GPP), pH and total phosphorus concentration were higher than at low fish density. Fish impact on bacterial abundance, dissolved oxygen, nitrogen and phosphorus concentrations, however, was not significant. The within treatment measurements of all variables except GPP were significantly different over time. Our results indicate that YOY walleye predation at high density can affect plankton community by reducing large zooplankton biomass and water clarity, and increasing phytoplankton abundance. The impact of YOY piscivorous fish on plankton should be considered when biomanipulation is applied for improvement of water quality.     

The influence of mesozooplankton on phytoplankton nutrient limitation: a mesocosm study with northeast Atlantic plankton

We used marine phytoplankton from mesocosms seeded with different zooplankton densities to study the impact of mesozooplankton on phytoplankton nutrient limitation. After 7 d of grazing (copepod mesocosms) or 9 d (appendicularian mesocosms) phytoplankton nutrient limitation was studied by enrichment bioassays. After removal of mesozooplankton, bioassay bottles received either no nutrients, phosphorus or nitrogen alone, or a combination of nitrogen and phosphorus and were incubated for 2 d. Phytoplankton reproductive rates in the bottles without nutrient addition were calculated after correction for grazing by ciliates and indicated increasing nitrogen limitation with increasing copepod abundance. No nutrient limitation was found in the appendicularian mesocosms. The increase of nutrient limitation with increasing copepod density seems to be mainly the result of a trophic cascade effect: Copepods released nanoplankton from ciliate grazing pressure, and thereby enhanced nitrogen exhaustion by nanophytoplankton and reduced nitrogen excretion by ciliates. Nitrogen sequestration in copepod biomass, the mechanism predicted by the ecological stoichiometry theory, seems to have been a weaker effect because there was only little copepod growth during the experiment.     

Effects of planktivorous and benthivorous fish on organisms and water chemistry in eutrophic lakes

The effects of planktivorous and benthivorous fish on benthic fauna, zooplankton, phytoplankton and water chemistry were studied experimentally in two eutrophic Swedish lakes using cylindrical enclosures. In enclosures in both lakes, dense fish populations resulted in low numbers of benthic fauna and planktonic cladocerans, high concentration of chlorophyll, blooms of blue-green, algae, high pH and low transparency. In the soft-water Lake Trummen, total phosporus increased in the enclosure with fish, but in the hard-water Lake Bysjön total phosphorus decreased simultaneously with precipitation of calcium carbonate. Enclosures without fish had a higher abundance of benthic fauna and large planktonic cladocerans, lower phytoplankton biomass, lower pH and higher transparency.The changes in enclosures with fish can be described as eutrophication, and those in enclosures without fish as oligotrophication. The possibility of regulation of fish populations as a lake restoration method is discussed.This paper was presented at the XXth SIL Congress in Copenhagen in 1977.     

Size‐based interactions across trophic levels in food webs of shallow Mediterranean lakes

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