Studies on the Effects of Silver Carp (Hypophthalmichthys molitrix) on the Phytoplankton in a Shallow Hypereutrophic Lake through an Enclosure Experiment

The responses of nutrients, water transparency, zooplankton and phytoplankton to a gradient of silver carp biomass were assessed using enclosure methods. The gradient of four silver carp biomass levels was set as follows: 0, 116, 176 and 316 g m^—2. Nutrients did not show any statistically significant differences among the treatments. An outburst of Daphnia only occurred in fishless enclosures where phytoplankton biomass was the lowest and water clarity significantly increased. While among fish enclosures, the small‐sized Moina micrura dominated throughout the experiment and both zooplankton and phytoplankton biomasses decreased with increased fish biomass. No large colonial cyanobacterial blooms occurred in the fishless enclosures as predicted. This might be due to low water temperature, short experiment time and the occurrence of large bodied Daphnia in our experiment. Cryptophyta was the most dominant group in most of the enclosures and the lake water throughout the experiment. The fishless enclosure had much lower proportion of Cyanophyta but higher proportion of Trachelomonas sp.     

An experimental study of the effects of fish zooplanktivory on the phytoplankton of a Brazilian reservoir

Two short-term (4–5 week) sets of enclosure experiments were conducted during winter periods (1982, 1983) in Americana Reservoir near Sao Paulo to test effects of fish predation reducing grazing pressure by large cladoceran zooplankton (mainlyDaphnia gessneri) on phytoplankton density, cell size distribution, biomass, species composition and richness. Two enclosures were stocked with the zooplanktivorous characin fishAstyanax whereas two others remained fishless during each set of experiments. Within two weeks or less, phytoplankton cell density and biomass were significantly reduced in the fishless enclosures, particularly in the small cell size (< 15 µm) fraction. Changes also occurred in species composition and richness. These shifts generally were maintained or intensified during the course of the experiments.     

Effects of a filter-feeding fish [silver carp, Hypophthalmichthys molitrix (Val.)] on phyto- and zooplankton in a mesotrophic reservoir: results from an enclosure experiment

SUMMARY 1. Silver carp, Hypophthalmichthys molitrix (Val.), feeds on both phyto- and zooplankton and has been used in lake biomanipulation studies to suppress algal biomass. Because reports on the effects of silver carp on lake food webs have been contradictory, we conducted an enclosure experiment to test how a moderate biomass of the fish (10 g wet weight m⁻³) affects phytoplankton and crustacean zooplankton in a mesotrophic temperate reservoir.
2. Phytoplankton biomass <30 μm and particulate organic carbon (POC) <30 μm were significantly higher in enclosures with silver carp than in enclosures without fish, whereas Secchi depth was lower. Total copepod biomass declined strongly in both treatments during the experiment, but it was significantly higher in fish-free enclosures. Daphnid biomass was also consistently higher in enclosures without fish, although this effect was not significant. However, the presence of fish led to a fast and significant decrease in the size at maturity of Daphnia galeata Sars. Thus, the moderate biomass of silver carp had a stronger negative effect on cladoceran zooplankton than on phytoplankton.
3. Based on these results and those of previous studies, we conclude that silver carp should be used for biomanipulation only if the primary aim is to reduce nuisance blooms of large phytoplankton species (e.g. cyanobacteria) that cannot be effectively controlled by large herbivorous zooplankton. Therefore, stocking of silver carp appears to be most appropriate in tropical lakes that are highly productive and naturally lack large cladoceran zooplankton.     

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.     

Predation by underyearling perch (Perca fluviatilis) on a Daphnia galeata population in a short-term enclosure experiment

1. The predation impact of underyearling perch (20 mm total length) on the dynamics of Daphnia galeata was studied in three 6.5-m³ enclosures during a 17-day experiment in June 1995. These data were compared with zooplankton succession in three fish-free control enclosures and in the pelagic zone of Bautzen reservoir, Germany. 2. Due to individual growth during the experiment, fish biomass in the enclosures increased from 210 mg wet body mass (w.b.m.) m^–3 to 830 mg w.b.m. m^–3, equivalent to an increase from 20 kg ha^–1 to 75 kg ha^–1. 3. In the enclosures with fish, biomass of daphnids decreased steadily to values below 1 mg wet weight (w.w.) l^–1 within 17 days, whereas in the fish-free controls and in Bautzen reservoir the Daphnia biomass fluctuated around 8 mg w.w. l^–1. Other zooplankton species exhibited little or no change. Approximately 60 kg ha^–1 was calculated as the critical underyearling perch biomass which may induce a drastic decline of the Daphnia galeata population in Bautzen reservoir. Comparison with values from other lakes is difficult due to differences in water depths and Daphnia biomasses. 4. Mean individual biomass of daphnids, egg ratio and proportion of adult daphnids were significantly lower in the enclosures with fish compared with the control enclosures at the end of the experiment. This may be explained partly by preferential predation of the large, egg-carrying daphnids by fish. However, no difference was found in clutch size and size at first reproduction, possibly due to the short duration of the experiment. 5. It must be assumed that the ‘summer depression’ of daphnids observed in many waters is not the exclusive effect of direct reduction of daphnids by fish predation. Even the high biomass of underyearling perch stocked in the enclosures did not completely account for Daphnia mortality. Instead, the selective loss of large size classes, combined with low food resources and reduced reproduction rates, may induce the marked declines in daphnids.     

Effects of leaping grey mullet Liza saliens (Osteichthyes, Mugilidae) in the macrophyte beds of oligohaline Mediterranean coastal lagoons

Some mugilid fish are known to enhance small phytoplankton in freshwater macrophyte-free environments due to zooplankton depletion. This suggests that they may have negative effects on natural macrophyte beds of freshwater and oligohaline lagoons due to phytoplantkon enhancement. To test this hypothesis, we compared the ecosystems of control enclosures that contained no fish with those of enclosures stocked with Liza saliens at two different densities. The occurrence of L. saliens at a density of 321±92.42 kg ha^–1 reduced cladoceran density, depleted epiphytic chironomid larvae, enhanced mayfly nymphs and cyclopoid copepods and reduced the organic matter content of sediment, all in comparison with control enclosures. At a density of 673±42.04 kg ha^–1, L. saliens reduced total zooplankton density, depleted epiphytic and sediment dwelling chironomid larvae and enhanced mayfly nymphs. The organic matter contents of sediment was not affected. These results showed that L. saliens was very effective in reducing zooplankton density even when macrophyte biomass was high. However, these effects do not affect phytoplankton density, probably because zooplankton was dominated by species with low filter-feeding rates and macrophytes depleted nutrients.     

Effects of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community of a deep,tropical reservoir: an enclosure experiment

1. An in situ enclosure experiment was conducted in a deep reservoir of southern China to examine (i) the effects of a low biomass (4 g wet weight m^?3) of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community and (ii) the response of Daphnia to eutrophication. 2. In the absence of fish, Daphnia galeata dominated the zooplankton community, whereas calanoids were dominant in the fish treatments, followed by D. galeata. Silver carp stocking significantly reduced total zooplankton biomass, and that of D. galeata and Leptodorarichardi, but markedly increased the biomass of smaller cladocerans, copepod nauplii and rotifers. In contrast, nutrient enrichment had no significant effect on the plankton community except for cyclopoids. 3. Chlorophyta, Cryptophyta and Bacillariophyta were dominant phytoplankton groups during the experiment. Chlorophyta with high growth rates (mainly Chlorella vulgaris in the fish enclosures and Ankyra sp. in the fishless enclosures) eventually dominated the phytoplankton community. Total phytoplankton biomass and the biomass of edible phytoplankton [greatest axial linear dimension (GALD) < 30 μm], Chlorophyta, Cryptophyta, Bacillariophyta and Cyanobacteria showed positive responses to fish stocking, while inedible phytoplankton (GALD ≥ 30 μm) was significantly reduced in the fish enclosures. However, there was no significant effect on the plankton community from the interaction of fish and nutrients. 4. Overall, the impact of fish on the plankton community was much greater than that of nutrients. High total phosphorus concentrations in the control treatment and relatively low temperatures may reduce the importance of nutrient enrichment. These results suggest it is not appropriate to use a low biomass of silver carp to control phytoplankton biomass in warmer, eutrophic fresh waters containing large herbivorous cladocerans.     

Indirect effects of fish on macrophytes in Bays Mountain Lake: evidence for a littoral trophic cascade

Summary We began this experiment to test specific hypotheses regarding direct and indirect effects of fish predation on the littoral macroinvertebrate community of Bays Mountain Lake, Tennessee. We used 24 m² enclosures in which we manipulated the presence and absence of large redear sunfish (Lepomis microlophus>150 mm SL), and small sunfish (L. macrochirus and L. microlophus <50 mm SL) over a 16-mo period. Here we report on effects of fish predation on gastropod grazers that appear to cascade to periphyton and macrophytes.Both large redear sunfish and small sunfish maintained low snail biomass, but snails in fish-free controls increased significantly during the first 2-mo of the experiment. By late summer of the first year of the experiment, the difference in biomass between enclosures with and without fish had increased dramatically (>10×). Midway through the second summer of the experiment, we noted apparent differences in the abundance of periphyton between enclosures containing fish and those that did not. We also noted differences in the macrophyte distribution among enclosures. To document these responses, we estimated periphyton cover, biovolume and cell size frequencies as well as macrophyte distributions among enclosures at the end of the experiment. When fish were absent, periphyton percent cover was significantly reduced compared to when fish were present. Periphyton cell-size distributions in enclosures without fish were skewed toward small cells (only 12% were greater than 200 m³), which is consistent with intense snail grazing. The macrophyte Najas flexilis had more than 60 x higher biomass in the fish-free enclosures than in enclosures containing fish; Potamogeton diversifolius was found only in fish-free enclosures. These results suggest a chain of strong interactions (i.e. from fish to snails to periphyton to macrophytes) that may be important in lake littoral systems. This contrasts sharply with earlier predictions based on cascading trophic interactions that propose that fish predation on snails would enhance macrophyte biomass.     

Interactions between phytoplankton,zooplankton and fish in a shallow,hypertrophic lake: a study of phytoplankton collapses in Lake Søbygård,Denmark

Since 1983 severe phytoplankton collapses have occurred 1–4 times every summer in the shallow and hypertrophic Lake Søbygård, which is recovering after a ten-fold decrease of the external phosphorus loading in 1982. In July 1985, for example, chlorophyll a changed from 650 µg l^–1 to about 12 µg 1^–1 within 3–5 days. Simultaneously, oxygen concentration dropped from 20–25 mg O₂l^–1 to less than 1 mg O₂l^–1, and pH decreased from 10.7 to 8.9. Less than 10 days later the phytoplankton biomass had fully recovered. During all phytoplankton collapses the density of filter-feeding zooplankton increased markedly, and a clear-water period followed. Due to marked changes in age structure of the fish stock, different zooplankton species were responsible for the density increase in different years, and consequently different collapse patterns and frequencies were observed.The sudden increase in density of filter-feeding zooplankton from a generally low summer level to extremely high levels during algae collapses, which occurred three times from July 1984 to June 1986, could neither be explained by changes in regulation from below (food) nor from above (predation). The density increase was found after a period with high N/P ratios in phytoplankton or nitrate depletion in the lake. During that period phytoplankton biomass, primary production and thus pH decreased, the latter from 10.8–11.0 to 10.5. We hypothesize that direct or indirect effects of high pH are important in controlling the filter-feeding zooplankton in this hypertrophic lake. Secondarily, this situation affects the trophic interactions in the lake water and the net internal loading of nutrients. Consequently, not only a high content of planktivorous fish but also a high pH may promote uncoupling of the grazing food-web in highly eutrophic shallow lakes, and thereby enhance eutrophication.A tentative model is presented for the occurrence of collapses, and their pattern in hypertrophic lakes with various fish densities.     

鲢放养和施肥对盐碱池塘围隔生态系统浮游生物群落的影响

报道了单养链（Hypophthalmichthys molitrix)和施肥对盐碱池塘围隔生态系统浮游生物群落的影响,链的放养使浮游植物丰度,叶绿素a 含量和和初组生产力增大,浮游植物小型化,生物量以小型硅藻和绿藻占优势,裸藻和金藻占有相当比重;浮游动物生物量减少,特别是枝角类的生物量无鱼围隔大于有鱼围隔,且多是较大型的种类,施肥特别是施无机肥能显著地提高浮游植物丰度和初级生产力,浮游动物生物量也增大,施有机肥围隔浮游植物和浮游动物生物量虽高于有鱼对照围隔,但其浮游植物初级生产力,叶绿素a含量,浮游生物多样性指数,P／R系数均较低,链鱼的生长最差,文中讨论了滤食性鱼类和施肥对浮游生物的影响。     

A Short-Term Study of Vertical and Horizontal Distribution of Zooplankton During Thermal Stratification in Lake Kinneret,Israel

This study documents for the first time both vertical and horizontal distribution patterns of the zooplankton community in Lake Kinneret during the period of thermal stratification. The zooplankton distribution patterns were explored in relation to abiotic (temperature, oxygen) and biotic (picocyanobacteria, ciliates, flagellates, phytoplankton, fish) environmental gradients. Sampling was carried out on 6–7 July 1992 at five stations and six depths from nearshore to offshore. Zooplankton abundance and biomass varied from 5 to 267 ind. l^–1(mean: 95 ind. l^–1), and from 0.1 to 65 d.w. mg m^–3(mean: 24 d.w. mg m^–3). Zooplankton taxonomic groups (Rotifera, Cladocera, Cyclopoida, Calanoida) and size classes (micro-, meso- and macrozooplankton) showed peaks of maximal density and biomass in the epilimnetic and metalimnetic strata (5 and 14 m). Depth, accounting for 31–39% of total spatial variation, reflected the vertical distribution of zooplankton in relation to temperature and oxygen declines, and the higher concentration of food resources (protists and phytoplankton) in the epilimnion and metalimnion. Onshore–offshore distance, accounting for 17–22% of the total spatial variance, reflected different distribution patterns shown among zooplankton groups and size classes. The macrozooplankton (Copepoda, Cladocera) was more abundant offshore, whereas microzooplankton (Rotifera and nauplii) predominated nearshore. These horizontal distribution patterns were related to small increases in temperature and phytoplankton biomass, and higher concentrations of fish in the littoral zone. Although limited to a short temporal scale, our study indicated that zooplankton spatial distribution in Lake Kinneret during the period of thermal stratification was related to physicochemical, food and predation factors, manifested differently along the vertical and nearshore–offshore gradients.     

Ecological restoration in eutrophic Lake Wuli: A large enclosure experiment

A large-scale enclosure experiment for lake restoration was carried out in Lake Wuli, a northern bay of shallow and eutrophic Lake Taihu in China. The large enclosure with an area of 10 ha was set up in the littoral zone and was bordered by waterproof fabric which did not cover the sediments. Multiple approaches were used and included fish removal, piscivorous fish stocking, shoreline reconstruction, aquatic macrophyte planting, benthic macro-animal stocking, and silver carp cultivation in pens for reduction of cyanobacteria. The results showed that the coverage of aquatic macrophytes increased from 0% to 45.7%. Mean concentrations of TN and TP inside the enclosure from May 2004 to May 2008 were 22.2% and 26.0% of those outside, respectively. Secchi depth was 0.40 m outside the enclosures and 0.75 m inside. However, responses of phytoplankton to the restoration project lagged behind improvement of water quality and reestablishment of aquatic plants. The phytoplankton biomass gradually decreased after the third year of the restoration. Stocking piscivorous fish and planting submerged macrophytes could not increase zooplankton biomass and enhance graze pressure on phytoplankton, most likely due to high omnivorous fish density and lower nutrition inside the enclosure. Higher grazing pressure of zooplankton on phytoplankton was observed in May and October every year. Zooplankton to phytoplankton biomass ratios were significantly negatively correlated with phytoplankton biomass outside (r = −0.440, p < 0.01) and inside the enclosure (r = −0.336, p < 0.05) from February 2004 to March 2007. Therefore, phytoplankton biomass inside and outside the enclosure was lower in May and October. Higher grazing pressure of zooplankton on phytoplankton in spring may result in occurrence of the clear-water phase that facilitated growth of submerged macrophytes in the littoral in Lake Wuli, and a clear-water state and improved water quality would likely be sustained throughout the year after reestablishment of submerged macrophytes.     

Influence of benthic and interstitial processes on nutrient changes along a regulated reach of a large river (Rhône River,France)

An enclosure study was conducted in Ranger Lake in south-central Ontario, Canada from 4 July to 5 August 1997 to determine predation effects of the larvae of the phantom midge fly Chaoboruson the zooplankton community. Zooplankton assemblages were established in 12 enclosures (2 m in diameter, 7.5 m deep). Three densities of fourth-instar Chaoborus trivittatus (0 l^–1, 0.1 l^–1 and 0.5 l^–1) were introduced as predator treatments to the enclosures. Temperature, dissolved oxygen and zooplankton community composition were monitored for six weeks. To determine if the zooplankton community composition changed, a repeated measures multivariate analysis was performed on percent biomass of Bosmina and calanoid copepods. There were no significant differences in mean taxon percent biomass among predator treatments. There were significant differences in mean taxon percent biomass between water layers (epilimnion and metalimnion). There were also significant differences in lengths of Bosmina and calanoid copepods among predator treatments at the end of the experiment. Crop content analysis of C. trivittatusshowed that Bosmina constituted 88–98% of the prey items found in the crops. These results demonstrate that the use of deep enclosures, a Chaoborus species which vertically migrates, and lower natural densities of Chaoborus may provide prey with an important natural refuge from predation and so allow a more accurate determination of the predation impact of Chaoborus trivittatusin temperate lakes where fish control Chaoborus densities.     

Scaling food chains in aquatic mesocosms: do the effects of depth override the effects of planktivory?

To assess the effects of physical dimension and planktivorous fish on phytoplankton standing crop, we repeated an experiment at different scales in plastic enclosures during summer 1995 in Lake Créteil, France. Enclosures were scaled for a constant surface (1.5 × 1.5 m) as depth was increased from 2.5 to 4.5 m. Even-link (zooplankton and phytoplankton) and odd-link (planktivorous fish, zooplankton and phytoplankton) food webs were established in both shallow and deep enclosures. Fish densities in the deep enclosures were scaled to allow comparisons with shallow ones for both in individuals m⁻² or individuals m⁻³. We explicitly designed this experiment to examine the scale-dependent behavior of the top-down mechanism of algal biomass control in lakes, and in particular to test the hypothesis of stronger cascading effects of fish on lower trophic levels at reduced depth. Both fish and enclosure size had highly significant effects on phytoplankton biomass over the duration of the experiment. No depth × fish interaction effects were observed. The presence of planktivorous fish enhanced phytoplankton biomass in both shallow and deep enclosures, although the reduction in depth generally produced a stronger effect. The mean concentration of chlorophyll a in the deep odd-link systems (ca 5 mg m⁻³) was lower than in the shallow even-link systems (ca 17 mg m⁻³). Statistical interpretation did not change when data were expressed as phytoplankton biomass per unit of surface area. Light limitation and zooplankton grazing are the most probable mechanisms explaining our results in these nutrient-enriched systems. Moreover, we found that the strength of the cascading effect of fish on plankton was not a function of depth. We believe that further studies on scaling effects should be conducted in order to improve our understanding of ecological patterns and to extrapolate results from micro/mesocosms to natural ecosystems. Received: 18 January 1999 / Accepted: 7 June 1999     

Effects of fish density on planktonic communities and water quality in a manipulated forest pond

We examined the effects of fish on lower trophic levels in a small pond in eastern Finland. The pond was divided into four sections with plastic curtains and stocked with crucian carp (Carassius carassius); two sections had low (4.4–5.5 g m^–3) and two high (10.4–13.7 g m^–3) densities of fish. In the summer of 1987, the pond was sampled weekly for phyto- and zooplankton until the fish were removed by rotenone in September after a three month experiment. Fish density as well as the extent of macrophyte cover had a considerable impact on planktonic communities and water quality. Mean zooplankton biomass was significantly lower and phytoplankton biomass higher, at high fish density. Water transparency was correlated negatively with chlorophyll-a at low fish density but turbidity appeared to reduce transparency at high fish density. The composition and dynamics of the plankton also differed at different fish densities. The mechanisms behind these effects, and the influences of habitat and fish behaviour on the results, are discussed.     

The impact of zooplankton status on the management of Lake Kinneret (Israel)

Monthly averages of standing stock wet biomass of zooplankton in Lake Kinneret (Israel) varied between 11 and 76 g m^–2 during 1969–1981, with the exception of two months. Averaged contributions of different groups were: Cladocera 58%, Copepoda 35% and Rotifera 7%. Total standing crop wet biomass is highest during January–June, averages varied between 35 and 50 g m^–2, and decreases during summer–fall (23–36 g m^–2). The winter biomass of Cladocera fluctuated between 22 and 35 g m^–2 and dropped to a range of 9–23 g m^–2 in summer, whereas copepod biomass varied very little around an average of 18 g (ww) m^–2 with the exception of low values from April to June. The stock biomass of Rotifera is relatively high during winter floods season (December-March) whilst in summer it is very low.Young stages of fish in Lake Kinneret feed mostly on zooplankton and zoobenthic forms. The most abundant fish in the Kinneret ecosystem, Mirogrex terraesanctae terraesanctae, also feed on zooplankton at the adult stage throughout the year, and herbivorous fish consume zooplankton during the summer when lake plankton resources are limited.The summer ecosystem of Lake Kinneret is characterised as a steady state type, in which the impact of the zooplankton-chain is of great importance. Increase of predation pressure on zooplankton by fish can disequilibrate the balanced trophic relations existing between nannoplankton production and zooplankton grazing capacity. Such a situation can lead to organics accumulation as nannoplankton blooms, resulting in water quality deterioration. Management options aimed at preventing collapse of zooplankton populations are discussed.     

Does high nitrogen loading prevent clear‐water conditions in shallow lakes at moderately high phosphorus concentrations?

1. The effect of total nitrogen (TN) and phosphorus (TP) loading on trophic structure and water clarity was studied during summer in 24 field enclosures fixed in, and kept open to, the sediment in a shallow lake. The experiment involved a control treatment and five treatments to which nutrients were added: (i) high phosphorus, (ii) moderate nitrogen, (iii) high nitrogen, (iv) high phosphorus and moderate nitrogen and (v) high phosphorus and high nitrogen. To reduce zooplankton grazers, 1⁺ fish (Perca fluviatilis L.) were stocked in all enclosures at a density of 3.7 individuals m^?2. 2. With the addition of phosphorus, chlorophyll a and the total biovolume of phytoplankton rose significantly at moderate and high nitrogen. Cyanobacteria or chlorophytes dominated in all enclosures to which we added phosphorus as well as in the high nitrogen treatment, while cryptophytes dominated in the moderate nitrogen enclosures and the controls. 3. At the end of the experiment, the biomass of the submerged macrophytes Elodea canadensis and Potamogeton sp. was significantly lower in the dual treatments (TN, TP) than in single nutrient treatments and controls and the water clarity declined. The shift to a turbid state with low plant coverage occurred at TN >2 mg N L^?1 and TP >0.13–0.2 mg P L^?1. These results concur with a survey of Danish shallow lakes, showing that high macrophyte coverage occurred only when summer mean TN was below 2 mg N L^?1, irrespective of the concentration of TP, which ranged between 0.03 and 1.2 mg P L^?1. 4. Zooplankton biomass and the zooplankton : phytoplankton biomass ratio, and probably also the grazing pressure on phytoplankton, remained overall low in all treatments, reflecting the high fish abundance chosen for the experiment. We saw no response to nutrition addition in total zooplankton biomass, indicating that the loss of plants and a shift to the turbid state did not result from changes in zooplankton grazing. Shading by phytoplankton and periphyton was probably the key factor. 5. Nitrogen may play a far more important role than previously appreciated in the loss of submerged macrophytes at increased nutrient loading and for the delay in the re‐establishment of the nutrient loading reduction. We cannot yet specify, however, a threshold value for N that would cause a shift to a turbid state as it may vary with fish density and climatic conditions. However, the focus should be widened to use control of both N and P in the restoration of eutrophic shallow lakes.     

Impact of submerged macrophytes on fish-zooplanl phytoplankton interactions: large-scale enclosure experiments in a shallow eutrophic lake

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

Grazer control and nutrient limitation of phytoplankton biomass in two Australian reservoirs

1. Grazer and nutrient controls of phytoplankton biomass were tested on two reservoirs of different productivity to assess the potential for zooplankton grazing to affect chlorophyll/phosphorus regression models under Australian conditions. Experiments with zooplankton and nutrients manipulated in enclosures, laboratory feeding trials, and the analysis of in-lake plankton time series were performed. 2. Enclosures with water from the more productive Lake Hume (chlorophyll a = 3–17.5 μg l^–1), revealed significant zooplankton effects on chlorophyll a in 3/6, phosphorus limitation in 4/6 and nitrogen limitation in 1/6 of experiments conducted throughout the year. Enclosures with water from the less productive Lake Dartmouth (chlorophyll a = 0.8–3.5 μg l^–1), revealed significant zooplankton effects in 5/6, phosphorus limitation in 5/6 and nitrogen limitation in 2/6 of experiments. 3. While Lake Hume enclosure manipulations of the biomass of cladocerans (Daphnia and Diaphanosoma) and large copepods (Boeckella) had negative effects, small copepods (Mesocyclops and Calamoecia) could have positive effects on chlorophyll a. 4. In Lake Hume, total phytoplankton biovolume was negatively correlated with cladoceran biomass, positively with copepod biomass and was uncorrelated with total crustacean biomass. In Lake Dartmouth, total phytoplankton biovolume was negatively correlated with cladoceran biomass, copepod biomass and total crustacean biomass. 5. In both reservoirs, temporal variation in the biomass of Daphnia carinata alone could explain more than 50% of the observed variance in total phytoplankton biovolume. 6. During a period of low phytoplankton biovolume in Lake Hume in spring–summer 1993–94, a conservative estimate of cladoceran community grazing reached a maximum of 0.80 day^–1, suggesting that Cladocera made an important contribution to the development of the observed clear-water phase. 7. Enclosure experiments predicted significant grazing when the Cladocera/Phytoplankton biomass ratio was greater than 0.1; this threshold was consistently exceeded during clear water phase in Lake Hume. 8. Crustacean length had a significant effect on individual grazing rates in bottle experiments, with large Daphnia having highest rates. In both reservoirs, mean crustacean length was negatively correlated with phytoplankton biovolume. The observed upper limit of its variation was nearly twice as high compared to other world lakes.     

Phytoplankton and zooplankton community structure after nutrient additions to the oligotrophic Lake Hecklan,Sweden

Lake Hecklan, in central Sweden, was fertilized with phosphorus and nitrogen during thermal stratification (late May-early Oct) 1984–1987. The nutrient additions were relatively small and raised the total phosphorus concentrations from 6 to 10 µg l^–1. The working hypothesis was that this moderate increase in the phosphorus concentration could increase the phytoplankton biomass without adverse changes in the planktonic community structure. The fertilization increased the phytoplankton biomass from 0.1 to a maximum of 2 mm³ l^–1. Chrysophyceae and Cryptophyceae dominated throughout the experimental period. Thus, the phytoplankton composition remained typical for a Swedish forest lake and provided a potential for increased zooplankton growth. An increased growth of zooplankton was indicated by increased biomass of Cladocera and Copepoda in 1984 and 1985, and by increased fecundity of herbivorous zooplankton.