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.     

The impact of planktivorous flsh on the structure of a plankton community

SUMMARY. 1. The abundance of pianktivorous juvenile yellow perch, Perca flavescens , was manipulated in three 750 m³ enclosures in a eutrophic lake.
2. There was a significant negative relationship between fish and zoopiankton biomasses. At high fish densities the zooplankton community was dominated by small filter-feeding cladocera. primarily bosmi- nids. At low fish densities the zooplankton community was dominated by large filter-feeding cladocera, primarily daphnids.
3. There was no significant relationship between zooplankton and phytoplankton biomasses when considered over the whole experiment but there was a trend towards lower phytoplankton biomass in the enclosure dominated by daphnids during mid-summer.
4. We conclude that although planktivorous fish have a strong negative impact on zooplankton community biomass and size structure, the relationship at the next lower trophic level, zooplankton and phytoplankton, is much weaker. Therefore, the biomanipulation of planktivorous fish populations as a management technique to control phytoplankton abundance is largely ineffective.     

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.     

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.     

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.     

用鲢鳙直接控制微囊藻水华的围隔试验和湖泊实践

为了控制水体富营养化所形成的水华,国外经典的生物操纵论者提倡放养食鱼性鱼类以控制食浮游生物的鱼类,藉此壮大浮游动物种群,然后借助浮游动物遏制藻类.按照这条思路,以浮游生物为食的鲢和鳙应该是清除的对象.本文作者认为我国湖泊中危害性最大的是微囊藻水华,而浮游动物根本不能摄食这种水华,不如直接利用鲢鳙进行控制,因而1989-1992年间在武汉的东湖进行了三次原位围隔试验,2000年再度重复了一次围隔试验,结果证明,迄今在东湖中已消失18年的微囊藻水华,重新出现在不养鱼的围隔里.在养入一定密度的鲢或/和鳙的围隔中,就没有水华出现.已经出现水华的围隔,放入适当密度的鲢或/和鳙后,在短期内水华又复消失;而放入草鱼的围隔,则水华始终持续,不受影响.说明鲢和鳙能有效地遏制微囊藻水华.鲢、鳙遏制水华的有效放养密度(亦即有效生物量)为46-50g·m^-3.东湖湖泊中鲢、鳙的生物量,直到1985年才达到这个水平,所以1985年湖里反常地没有出现水华.往后的年份,东湖的鱼产量越来越高,也就是湖里鲢、鳙的生物量越来越大,所以迄今没有微囊藻卷土重来的机会.东湖的水质和其他非生物条件仍适于蓝藻水华的发生与发展,如果东湖鲢、鳙的产量下降到阈值以下,亦即降到1985年以前1000t水平,那么微囊藻及其它蓝藻水华的重来将是不可避免的.经典的生物操纵论学说和措施不能解决微囊藻水华猖獗的问题.相反,用滤食浮游生物的鲢和鳙直接遏制微囊藻却是有实验根据并已被渔业生产实践证明行之有效的.     

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。     

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.     

Response of plankton communities to whole-lake Ca(OH)2 and CaCO3 additions in eutrophic hardwater lakes

1. The impact of whole-lake lime (slaked lime, Ca(OH)₂, and/or calcite, CaCO₃) addition on plankton communities was evaluated in eutrophic hardwater lakes on the North American Boreal Plain.
2. Two lakes received a single treatment of lime (Ca(OH)₂ at 74 or 107 mg L^–1), two lakes received multiple treatments with Ca(OH)₂ and/or CaCO₃ (5–78 mg L^–1), and four lakes were untreated and served as reference systems.
3. Over the long-term (> 1 year), phytoplankton biomass was reduced in multiple-dose lakes, but not in single-dose lakes. Cyanobacteria typically dominated the algal community in the years before, during and after lime treatment in both single- and multiple-dose lakes.
4. In the single-dose lakes, randomized intervention analysis showed no significant change in the biomass of zooplankton after lime addition.     

The role of light for fish–zooplankton–phytoplankton interactions during winter in shallow lakes – a climate change perspective

1. Variations in the light regime can affect the availability and quality of food for zooplankton grazers as well as their exposure to fish predation. In northern lakes light is particularly low in winter and, with increasing warming, the northern limit of some present-day plankton communities may move further north and the plankton will thus receive less winter light.
2. We followed the changes in the biomass and community structure of zooplankton and phytoplankton in a clear and a turbid shallow lake during winter (November–March) in enclosures both with and without fish and with four different light treatments (100%, 55%, 7% and <1% of incoming light).
3. In both lakes total zooplankton biomass and chlorophyll- a were influenced by light availability and the presence of fish. Presence of fish irrespective of the light level led to low crustacean biomass, high rotifer biomass and changes in the life history of copepods. The strength of the fish effect on zooplankton biomass diminished with declining light and the effect of light was strongest in the presence of fish.
4. When fish were present, reduced light led to a shift from rotifers to calanoid copepods in the clear lake and from rotifers to cyclopoid copepods in the turbid lake. Light affected the phytoplankton biomass and, to a lesser extent, the phytoplankton community composition and size. However, the fish effect on phytoplankton was overall weak.
5. Our results from typical Danish shallow eutrophic lakes suggest that major changes in winter light conditions are needed in order to have a significant effect on the plankton community. The change in light occurring when such plankton communities move northwards in response to global warming will mostly be of modest importance for this lake type, at least for the rest of this century in an IPCC A2 scenario, while stronger effects may be observed in deep lakes.     

Karyotype variability in successive generations after hybridization between the great sturgeon, Huso huso (L.), and the sterlet, Acipenser ruthenus L.

Effects of particle size, fish size and temperature on the filtration rate of silver carp were determined. When feeding at 20°C on zooplankton and spherical particles (yeast, micronic beads and pollen), 32-g silver carp filter particles larger than 70 urn at a maximum rate of 18.251 h⁻¹. For particles smaller than 70 μm, filtration rates decrease with decreasing particle size until there is no measured filtration for particles smaller than 10 μm. Filtering rates ( FR ) for particles between 10 and 50 μm are described by the equation, FR =−20.8 + 21.7 × log particle diameter. Filtration rates rise as fish size, particle size and temperature increase. Filtration rates per unit biomass, however, fall as fish size increases: FR = 1.54 W^0.713, where FR is the maximum filtration rate in ¹ h ¹ fish ¹ and W is weight of fish in grammes. The results of these trials are consistent with the hypothesis that particle selection by silver carp is a mechanical, passive function of gill raker morphology.     

Comparative studies on the feeding selectivity of silver carp Hypophthalmichthys molitrix and bighead carp Aristichthys nobilis

Silver carp and bighead carp exhibited size-selection for food particles in aquarium experiments, but did not select their preferred species of plankton actively when they were distributed evenly in the water. They also possessed the capacity of selection for feeding area. The removal rates (% g⁻¹ fish weight) of silver carp for smaller plankton (phytoplankton) were higher than those of bighead carp. The removal rates by the latter for bigger plankton (zooplankton) were higher than those of silver carp, but for plankton about 70 μm dia. the rates by the two species were almost equal.     

Can a community of small-bodied grazers control phytoplankton in rivers?

1. Phytoplankton, zooplankton and grazing were monitored throughout the growing season for three years (1994–96) in the Belgian section of the River Meuse.
2. A size structure analysis of the algal community shows that there was a summer shift toward larger algal units, following a decline in phytoplankton biomass. These changes occurred after an increase in zooplankton biomass and diversity.
3. Daily filtration rates of grazers ranged from 1 to 113% day^–1 and maxima were observed during the summer period. Higher rates tended to correspond with peaks of rotifer biomass. A decline in total phytoplankton biomass within two weeks followed the increase in zooplankton biomass and filtration rate. A rapid biomass recovery was then observed, along with a shift of the algal community toward larger units. When grazing activity was not sustained, due to zooplankton fluctuations, the change in phytoplankton size structure was less marked.
4. We suggest that the composition of the phytoplankton community of large rivers may at times be controlled by grazers. However, such biotic interactions can take place only when physical constraints are reduced, i.e. when discharge is low, and when increased transfer time, high temperature and availability of grazeable algae allow high zooplankton biomass.     

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,     

Covariation of stream community structure and biomass of algae, invertebrates and fish with forest cover at multiple spatial scales

1. To evaluate the spatial extent of the effects of forest cover on stream ecosystems, we measured algae, invertebrate, and fish biomass and invertebrate and fish community structure in 38 small first- to third-order streams in the National Capital Region of Canada along with forest cover at different spatial scales.
2. We considered 55 spatial scales of forest cover including several buffer widths (doubling 10–320 m) and lengths (doubling 10–1280 m, entire riparian distance upstream from sampling area) and entire catchments to determine which spatial scale maximized the correlation with biomass and metrics of community structure.
3. The proportion of variability in biomass and structural metrics explained by forest cover generally increased with increasing scale, suggesting that catchment-wide disturbances are the most influential determinants of benthic and fish communities.
4. Catchment forest cover explained more variation in algal (adjusted r ²   =   0.54), invertebrate (adjusted r ²   =   0.51) and fish (adjusted r ²   =   0.33) biomass than structural metrics of invertebrates and fish (adjusted r ²   =   0.08–0.27).
5. Analyses of the partial effects of forest cover at three scales (reach, riparian and the entire catchment) on biomass and community structure metrics identified catchment and reach scales as being most influential and never detected a significant partial effect of forest cover at the riparian scale.
6. These results suggest that maintenance or protection of reach and riparian buffers alone will not sufficiently protect stream function and structure from catchment-wide impacts.     

The loss of submerged plants with eutrophication II. Relationships between fish and zooplankton in a set of experimental ponds, and conclusions

SUMMARY. 1. In 1982 and 1983 sets of experimental ponds were left with their submerged plant communities intact (plant ponds) or were cleared manually of them (cleared ponds). The ponds were all fertilized with ammonium nitrate and with variable amounts of phosphate. In 1982 fish were removed from the ponds. Zooplankton communities were dominated by large Cladocera with Daphnia prominent in the cleared ponds and Simocephalus in the plant ponds. There was no detectable effect of differential phosphorus additions on zooplankton communities or populations.
2. In 1983 zooplanktivorous fish (mainly roach) were stocked in the ponds. In the plant ponds the fish did not survive, probably through severe deoxygenation and the zooplankton community again included large-bodied Simocephalus. Fish survival was variable in the cleared ponds. Where fish stocks were absent or low (0.5–1 g m⁻²) a Daphnia- dominated community persisted; at intermediate fish stocks (18.1 g m⁻²) Eudiaptomus gracilis was predominant and where fish stock was high (22.8–29.1 g m⁻²) Bosmina longirostris , and cyclopoid copepods dominated the communities. Mean biomass of the zooplankton community declined with increase in fish stock to between 5.1 and 18.1 g m⁻² then increased.
3. On the basis of results from the experimental ponds and elsewhere, a new hypothesis is put forward to account for the switch from aquatic plant to phytoplankton dominance in eutrophicated shallow lakes. It envisages dominance by either group to be possible as alternative states over a wide range of high nutrient loadings. It suggests that each state is buffered against increased loading by mechanisms involving plant and algal physiology and zooplankton grazer populations. The nature of the buffers and the reasons by which one state may be switched to the other are, discussed.     

Seasonal succession of zooplankton in the north basin of Lake Biwa

To examine the seasonal succession of the entire zooplankton community in Lake Biwa, zooplankton biomass (on an areal basis) and its distribution patterns among crustaceans, rotifers and ciliates were studied in the north basin from April 1997 to June 1998. Seasonal changes in phytoplankton and population dynamics of Daphnia galeata were also examined to assess food condition and predation pressure by fish. From March to November, crustaceans dominated zooplankton biomass, but rotifers and ciliates were dominant from December to February. Among crustaceans, Eodiaptomus japonicus was the most abundant species, followed by D. galeata. Zooplankton biomass increased from January to a peak in early April, just before the spring bloom of phytoplankton, then decreased in mid-April when mortality rate of D. galeata increased. From mid-June, zooplankton increased and maintained a high level until the beginning of November. During this period, both birth and mortality rates of D. galeata were relatively high and a number of rotifer and crustacean species were observed. However, their abundances were very limited except for E. japonicus which likely preys on ciliates and rotifers. In Lake Biwa, food sources other than phytoplankton, such as resuspended organic matter from the sediments, seems to play a crucial role in zooplankton succession from winter to early spring, while zooplankton community seems to be regulated mainly by fish predation from summer to fall.     

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.     

Zooplankton biomass and community structure in a Danube River floodplain system: effects of hydrology

1. Zooplankton density and biomass was examined in a Danube River floodplain section with highly variable hydrological dynamics. Temporal patterns were analysed to assess the effects of hydrological conditions on zooplankton community structure and the differential response of the two major zooplankton taxa, rotifers and crustaceans.
2. Calculated floodplain water age was used as an integrated parameter describing hydrological conditions and connectivity.
3. Total zooplankton biomass, crustacean biomass and crustacean species number were significantly positively related to water age. Rotifer biomass followed a hump-shaped relationship with water age, and rotifer species number decreased with increasing water age.
4. Rotifers dominated the community in periods of low to medium water ages. In periods of higher water ages the community was dominated by crustaceans.
5. We propose that the hydrological regime of floodplains is crucial for zooplankton biomass patterns and succession, through the alternation of washing-out effects, taxon-specific potential of reproduction and biological interactions. Flood events and high water levels reset the community to an early successional phase.     

围隔中鲢和菹草控藻效果及其生态学意义

在滇池沿岸的池塘中 ,构建了面积为 10 0m2 ,水深为 85cm ,底质均为淤泥的四个围隔 ,其中在三个围隔内分别放养密度为 10 0g/m3 、75g/m3 、5 0g/m3 的鲢。它们均在一定程度上控制浮游植物的生物量并改善水体透明度 ,其中初始放养鲢 75g/m3 的围隔控制效果最明显。放养三种密度的鲢在有效控制微囊藻水华的同时 ,原先水体的优势种类发生质的变化 ,水体中浮游植物的多样性也明显增加。放鱼 2个月后 ,在生物操纵的围隔中同时种入菹草 ,在75g/m3 的鲢控藻的围隔中 ,菹草长势良好 ,浮游动物大量出现 ,透明度进一步增加