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.     

Evaluation of the potential fish productivity of lake oiskoe (Ergaky Mountain Range,West Sayan) basing on food supply

As a result of integrated studies carried out at the end of August–September 2008–2010, the icthyofauna and food reserve for fish in Oiskoe lake (Ergaky range) were investigated. The trophic status of the lake was estimated based on the biomass of phyto- and zooplankton and zoobenthos. The potential icthyomass, fish production and possible fishing amount were estimated based on calculations of the average biomass and production per season for zooplankton and zoobenthos.     

Experimental study of the impacts of silver carp on plankton communities of eutrophic Villerest reservoir (France)

We examined the impact of five silver carp biomass levels (0, 8, 16, 20, and 32 g m⁻³) on plankton communities and water quality of Villerest eutrophic reservoir (France). We realized the experiments using outdoor mesocosms. The presence of silver carp led to changes in zooplankton and phytoplankton assemblages. High fish biomass strongly reduced cladoceran abundance (through predation). Silver carp inefficiently grazed down particles < 20 μm. More importantly, however, the suppression of herbivorous cladocerans resulted in the increase of small size algae which were relieved from grazing and benefit from high nutrient concentrations. In contrast, in mesocosms without fish, the dominance of cladocerans (mainly Daphnia) controlled small size algae and probably also larger size algae (colonial chlorophytes, cyanobacteria). Thus, the Secchi disc transparency increased markedly. Through cascade effects, the modification of grazers communities led to changes in the utilization patterns of the added nutrients by phytoplankton communities. In high fish biomass treatments, nutrients were more efficiently accumulated into particulate fractions compared with no-fish and low-fish biomass treatments that were characterized by higher dissolved nutrients concentrations. Zooplankton was an essential source of food for silver carp. The productivity of zooplankton sustained a moderate silver carp biomass (up to 16 g m⁻³). In the presence of the highest fish biomass, the productivity of zooplankton was not large enough and silver carps fed on additional phytoplankton. Although mesocosms with high fish biomass were characterized by a slight cyanobacteria development compared with other fish mesocosms, silver carp was not effective in reducing cyanobacteria dominance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Impact of moderate silver carp biomass gradient on zooplankton communities in a eutrophic reservoir. Consequences for the use of silver carp in biomanipulation.

We examined the impacts of moderate gradient silver carp biomass (five levels from 0 to 36 g.m-3, i.e. about 0-792 kg.ha-1) on zooplankton communities of the eutrophic Villerest reservoir (France). During our mesocosm experiment changes in zooplankton assemblages were dependent on silver carp biomass. In the fishless and low fish biomass treatments, zooplankton abundance increased through time, owing to a peak in cladoceran density, but decreased (mainly cladocerans) at highest fish biomass. Copepods and rotifers were less affected at the highest fish biomass and dominated zooplankton communities. We highlighted that the presence of high silver carp biomass could lead to changes in phytoplankton assemblage via the impact on herbivorous zooplankton. Since silver carp efficiently graze on particles > 20 microns, the suppression of herbivorous cladocerans could result in an increase in small size algae (< 20 microns) abundance since these species would be released from grazers as well as competitors (large algae grazed by silver carp) and nutrients levels would be enhanced by fish internal loading. Our results showed that the use of low silver carp biomass (< 200 kg.ha-1) would allow us to minimize these negative effects.     

Autotroph:herbivore biomass ratios; carbon deficits judged from plankton data

A survey on phytoplankton:zooplankton biomass ratios was performed in 342 Norwegian lakes, covering a wide range in lake size and productivity (total phosphorus: 3–246 g l^–1), but with most localities being oligo- to mesotrophic. Mean phytoplankton biomass was 88 g C l^–1, yet with the majority below 50 g C l^–1and a median of 25 g C l^–1. Total zooplankton biomass displayed a mean and median of 37 and 26 g C l^–1, respectively. Cladocerans were by far the dominant group, making up a median of almost 60% of total zooplankton biomass. Total zooplankton biomass as well as that of major aggregated metazoan taxa (cladocerans, calanoid copepods, cyclopoid copepods and rotifers) all showed a positive, but weak correlation with total phytoplankton biomass. These weak correlations suggest that algal biomass per se is a poor predictor of zooplankton biomass. An average phyto-:zooplankton biomass ratio (C:C) of 2.8 (SD±4.7) was found. 30% of the lakes had a phyto-:zooplankton biomass ratio below unity. While there was no correlation between the phyto-:zooplankton biomass ratio with increasing productivity in terms of P concentration, there was a higher biomass ratio in lakes with high fish predation pressure. The low ratio of phyto-:zooplankton biomass suggest major requirements from non-algal sources of C in the zooplankton diet. The need for dietary subsidizing is also supported by the fact that more than 75% of the lakes had algal biomass less than the estimated threshold for net positive growth of zooplankton, although it should be kept in mind that a high share of picoplankton would imply an underestimation of autotroph biomass in these lakes. Since the C-deficiency apparently is most pronounced in oligotrophic systems, it contradicts the view that the detritus pathways plays a predominant role in highly productive systems only, but while the source of detritus probably is mostly of autochthonous origin in eutrophic lakes, allochthonous detritus will be more important in oligotrophic systems.     

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.     

Seasonal variations in stable isotope ratios of two biomanipulation fishes and seston in a large pen culture in hypereutrophic Meiliang Bay, Lake Taihu

This paper reports on seasonal changes in stable carbon and nitrogen isotope ratios of seston and muscle tissue of silver carp and bighead carp during 2004 and 2005, focusing primarily on the carbon sources and trophic relationships among phytoplankton, zooplankton and silver carp and bighead carp in a large fish pen of Meiliang Bay (Lake Taihu, China). δ¹³C showed a minimal value in March 2005 and a maximal value in August 2005 in seston both inside and outside the pen, whereas δ¹⁵N of seston showed the minimum in winter and the maximum during algal blooms. A positive correlation between δ¹³C of silver carp and that of seston suggested that temporal variation of δ¹³C in seston was preserved in fish via the food chain. The differences of δ¹³C among seston, zooplankton and muscle tissue of silver carp and bighead carp ranged only 0.2–1.7%, indicating that plankton production was the primary food source of filter-feeding fishes. According to a mass balance model, we estimated that the contributions of zooplankton to the diets of silver carp and bighead carp were 45.7% and 54.3%, respectively, based on the δ¹⁵N values of zooplankton and planktivorous fishes.     

Food habits, feeding behaviour and impact of triploid bighead carp, Hypophthalmichthys nobilis, in experimental ponds

Food of sterile triploid bighead carp, Hypophthalmichthys nobilis , was examined in ponds receiving water from a hypereutrophic take in Florida. No distinctive seasonal changes in food composition were found. The fish fed selectively on Botryococcus braunii Kuetzing, a large nuisance algal species, which constituted 61% of the volume and 50% of the dry weight of the food. Zooplankton was highly selected, but constituted only 3% of the food volume because of the low zooplankton density in the ponds. The remaining proportion of food consisted mainly of blue-green algae. Final fish biomass was low, ranging from 60 to 97 kg ha⁻¹ in individual ponds. Low biomass was due to slow fish growth and high fish mortality. Despite low biomass, the fish lowered the ratio of blue-green/green algae in the ponds and tended to lower phyto- and zooplankton abundance.     

Changes in the plankton community following introduction of filter-feeding planktivorous fish

1. We conducted enclosure experiments in a shallow eutrophic lake, in which a biomass gradient of the filter-feeding planktivore, silver carp, Hypophthalmichthys molitrix Valenciennes, was created, and subsequent community changes in both zooplankton and phytoplankton were examined.
2. During a summer experiment, a bloom of Anabaena flos-aquae developed (≈ 8000 cells mL⁻¹) solely in an enclosure without silver carp. Concurrent with, or slightly preceding the Anabaena bloom, the number of rotifer species and their abundance increased from seven to twelve species (1700–14 400 organisms L⁻¹) after the bloom in this fish-free enclosure. Protozoans and bacteria were generally insensitive to the gradient of silver carp biomass.
3. During an autumn experiment, on the other hand, large herbivorous crustaceans were more efficient than silver carp in suppressing the algae, partly because the lower water temperature (≈ 24 °C) inhibited active feeding of this warm-water fish and also formation of algal colonies. Heterotrophic nanoflagellate and bacterial densities were also influenced negatively by the crustaceans.
4. Correspondence analysis (CA) was applied to the weekly community data of zooplankton and phytoplankton. A major effect detected in the zooplankton community was the presence/absence of silver carp rather than the biomass of silver carp, whereas that in the phytoplankton community was the fish biomass before the Anabaena bloom, but shifted to the presence/absence of the fish after the bloom.     

鲤鱼种和鲢鳙对池塘浮游生物的影响

本文根据１９９０年－９月对９个鱼池的研究,报道鲤鱼种和鲢鳙对浮游生物的影响,结果表明：鲤鱼种使浮游植物和浮游动物生物量增加,浮游植物大型化,浮游动物小型化,并使浮游植物多样性增加,浮游动物多样性下降;鲢鳙密度增加,浮生物小型明显,微型藻类和超微在在浮植物中所占比重显著增加,小型浮游动物（原生动物和轮虫）在浮游动物中比重也明显增;鲢鳙密度对浮游生物多样性和浮游生物量的影响具有阶段性,当鲢鳙密度低时,     

太湖梅梁湾大型控藻围栏对浮游甲壳动物群落结构的影响

2005年对太湖梅梁湾大型鲢、鳙控藻围栏内外浮游甲壳动物群落结构的季节变化进行了监测.结果表明:围栏内外的环境因子、浮游植物生物量、浮游甲壳动物种类组成无显著差异.但鲢、鳙放养对浮游甲壳动物的生物量产生了较大的影响,围栏内浮游甲壳动物的总生物量和枝角类的生物量显著低于围栏外.总体上,枝角类各种类的生物量受鲢、鳙放养的影响程度大于桡足类的种类.太湖梅梁湾浮游甲壳动物的季节演替明显,大部分种类只是季节性出现.冬季和春季以溞(Daphnia sp.)和近邻剑水蚤(Cyclops vicnus)等大型种类为主,夏季和秋季以象鼻溞(Bosmina sp.)、角突网纹溞(Ceriodaphnia cornuta)和中华窄腹剑水蚤(Limnoithona sinensis)等小型种类为主.典范对应分析表明,透明度、温度和浮游植物的生物量是影响浮游甲壳动物季节变化的主要因素.     

Impacts of two biomanipulation fishes stocked in a large pen on the plankton abundance and water quality during a period of phytoplankton seasonal succession

Silver and bighead carp were stocked in a large pen to control the nuisance cyanobacterial blooms in Meiliang Bay of Lake Taihu. Plankton abundance and water quality were investigated about once a week from 9 May to 7 July in 2005. Biomass of both total crustacean zooplankton and cladocerans was significantly suppressed by the predation of pen-cultured fishes. There was a significant negative correlation between the N:P weight ratio and phytoplankton biomass. The size-selective predation by the two carps had no effect on the biomass of green alga Ulothrix sp. It may be attributed to the low fish stocking density (less than 40 g m⁻³) before June. When Microcystis dominated in the water of fish pen, the pen-cultured carps effectively suppressed the biomass of Microcystis, as indicated by the significant decline of chlorophyll a in the >38 μm fractions of the fish pen. Based on the results of our experiment and previous other studies, we conclude that silver and bighead carp are two efficient biomanipulation tools to control cyanobacterial (Microcystis) blooms in the tropical/subtropical eutrophic waters. Moreover, we should maintain an enough stocking density for an effective control of phytoplankton biomass.     

Determination analysis as a tool to study relationships between ecosystem components in water bodies of the Azov, Caspian, and Kara Sea basins

Contingencies of different components of freshwater biocenoses in the Don, Volga, Angara, and Northern Azov basins where studied by determination analysis. A positive contingency between the biomass of total phytoplankton or its divisions and the biomass of zooplankton was demonstrated. The most significant contingencies were revealed for zooplankton and green algae, dinoflagellates, and cyanobacteria. As against zooplankton, the biomass of zoobenthos features much less contingencies with the biomass of microalgae as well as zooplankton, since detritus, higher plant tissues, and small animals are the basis of the diet of benthic organisms. Differences in the contingency pattern between different geographical regions and time periods within the same basin have been revealed using the context procedure in determination analysis.     

Does stocking of filter-feeding fish for production have a cascading effect on zooplankton and ecological state? A study of fourteen (sub)tropical Chinese reservoirs with contrasting nutrient concentrations

Stocking of filter-feeding fish is a common tool used in Chinese reservoirs to increase fish production because of low natural recruitment. Whether such stocking has important negative effects on zooplankton with cascading effects on phytoplankton is debated. We compared the zooplankton communities in fourteen reservoirs with different nutrient concentrations and fish densities. Both chlorophyll a (Chla) and fish catch were positively related with total phosphorus (TP), whereas zooplankton biomass did not show a similar relationship with TP. Zooplankton seemed to be influenced by fish as high fish catches coincided with a low proportion of calanoids of the total copepod biomass, a high proportion of rotifers of the total zooplankton biomass, a low zooplankton:phytoplankton biomass ratio, and the absence of Daphnia irrespective of TP concentration. Both zooplankton biomass and most of the zooplankton:phytoplankton biomass ratios were among the lowest reported in the literature for the nutrient range studied. Furthermore, the Chla:TP ratio was higher than what is typically observed in temperate lakes. We conclude that top-down control of zooplankton is of key importance in reservoirs in South China where frequent stocking of filter-feeding fish seems to contribute to poor water quality in the form of higher algal biomass and reduced clarity.     

Spatial heterogeneity of zooplankton biomass and size structure in southern Quebec lakes: variation among lakes and within lake among epi-, meta- and hypolimnion strata

Environmental control of zooplankton biomass size structure(53–100, 100–202, 202–500 and >500 µm)was investigated in the three limnetic strata of 25 southernQuébec Shield lakes, Canada. Among-lake differences werethe greatest source of variation of zooplankton biomass, whereasthe strong lake–by–stratum interaction observedindicated that the vertical variations of zooplankton biomassand its size fractions were not constant from lake to lake.The analysis of spatial and local factors based on thermal stratais consistent with conceptual models of predation and nutrientcontrol on the biomass and size structure of the zooplankton.Productivity of the aquatic systems, which was driven by lakedepth, flushing rate and total phosphorus concentration, wasthe primary factor influencing total zooplankton biomass andsize structure at among-lake scale in epilimnetic waters. Theeffects of the planktivorous fish on the large zooplankton biomass(>500 µm) was more clearly perceived when the effectof lake depth was removed by partial redundancy analysis. Thisstudy showed that although bottom-up and top-down forces arecomplementary in structuring of zooplankton communities, theycan also act differently on the community attributes (e.g. biomassand size structure). Among-lake zooplankton biomass is predictablefrom lake trophy, but the size structure and vertical distributionof zooplankton communities appear to be controlled by lake stratificationand by inference to interactions with size selective predationby fish. In metalimnetic waters, the 53–100 and 100–202µm zooplankton biomass fractions were primarily dependenton abiotic factors, while the 202–500 and >500 µmfractions were related to planktivory and picophytoplanktonconcentrations. The well-oxygenated and cold hypolimnetic watersof some lakes offered a refuge from surface turbulence and planktivoryto large zooplankton size fractions (202–500 and >500µm).     

Effects of common carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis>) and bream (<Emphasis Type="Italic">Abramis brama</Emphasis>) on the structure of the littoral community in a mesotrophic lake (Mesocosm Experiments)

The effect of common carp and bream on hydrochemical parameters, abundance, and structure of phyto-, zooplankton, and macrozoobenthos has been studied in semienclosed mesocosms installed in the littoral zone of a mesotrophic lake. Significant differences in the biomass of different algal groups in mesocosms with fish in respect to the control were established only for diatoms and were not found for other phytoplankton groups. Common carp had a greater effect on the abundance of large zooplankton species (Diaphanosoma brachyurum) compared to bream. The abundance of the small Bosmina longirostris increased in mesocosms both with bream and common carp. The macrozoobenthos biomass reduced at higher rates in mesocosms with common carp than in those with bream, with the strongest effect of common carp on mayfly larvae. The differences between the consumption of chironomid and oligochaete larvae were not established in mesocosms with common carp. Bream mainly affected the larvae of mayflies and oligochaetes and, to a lesser degree, the chironomid larvae.     

Seasonal dynamics of phyto- and zooplankton and their interactions in the hypereutrophic reservoir

The seasonal dynamics of phyto- and zooplankton, as well as their trophic interactions, were studied for the shallow freshwater hypereutrophic Curonian Lagoon in the Baltic Sea in 2007–2009. The water temperature and high concentrations of nutrients were the limiting factors in the seasonal dynamics of the studied communities. Decreases in the B _z /B _ph ratio and the grazing pressure of phytophagous zooplankton were observed over the last few decades due to the eutrophication process and cyanobacteria blooms. The large zooplankton consumes a significant amount of primary production (Chlorococcales and Bacillariophyta) when the predator pressure of the fish fry is absent. This grazing pressure may precondition cyanobacteria scum.     

The influence of turbidity on growth and survival of fish larvae: a numerical analysis

Growth and survival through the early larval phase probably limit the production potential of many commercially important fish stocks. Attempts to increase the production of these stocks by restocking of juveniles have generally failed. Here, we analyse how enhanced concentrations of phytoplankton and zooplankton affect the survival of fish larvae during their early life stages. The analysis is developed for larvae feeding on copepod eggs and nauplii, with fish and invertebrates as major predators. A model of feeding and growth of fish larvae is applied to assess the benefit of enhanced phytoplankton and zooplankton abundance. The analysis shows that the shading effect of higher phytoplankton concentration may reduce predation rates on fish larvae substantially. This `top-down' effect may be more important for the cohort survivorship than the `bottom-up' mechanism in situations when larval food is sufficiently abundant. However, while increased algal biomass will improve recruitment at high zooplankton concentration, it may also reduce recruitment at low zooplankton concentrations and shallow mixing depths. Both the larvae and their vertebrate predators are dependent on light to detect their prey, and the longer reactive distance of the predators make them more susceptible than the larvae to reduced light levels and increased turbidity. We discuss the implications of reduced predation and increased zooplankton abundance on recruitment and production of fish larvae, and point at environmental conditions where changing algal biomass is likely to affect recruitment success.     

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.