Existence of a macrophyte-dominated clear water state over a very wide range of nutrient concentrations in a small shallow lake

Little Mere, a small shallow lake, has been monitored for four years, since its main source of nutrients (sewage effluent) was diverted. The lake has provided strong evidence for the persistence of a clear water state over a wide range of nutrient concentrations. It had clear water at extremely high nutrient concentrations prior to effluent diversion, associated with high densities of the large body-sized grazer, Daphnia magna, associated with low fish densities and fish predation. Following sewage effluent diversion in 1991, the nutrient concentrations significantly declined, the oxygen concentrations rose, and fish predation increased. The dominance of large body-sized grazers shifted to one of relatively smaller body-sized animals but the clear water state has been maintained. This is probably due to provision of refuges for grazers by large nymphaeid stands (also found prior to diversion). There has been a continued decrease in nutrient concentrations and expansion of the total macrophyte coverage, largely by submerged plants, following effluent diversion. The grazer community of Little Mere has also responded to this latter change with a decline in daphnids and increase in densities of weed-associated grazers. The presence of large densities of such open water grazers was the apparent main buffer mechanisms of the clear water state until 1994. The lake has, so far, maintained its clear water in the absence of such grazers. Thus, new buffer mechanisms appear to operate to stabilize the ecosystem. Little Mere appears to have shifted from previous top-down controlled clear water state to a bottom-up controlled clear water state.     

Consequences of reduced nutrient loading on a lake system in a lowland catchment: deviations from the norm?

1. Lake restoration from eutrophication often rests on a simple paradigm that restriction of phosphorus sources will result in recovery of former relatively clear‐water states. This view has apparently arisen from early successful restorations of deep lakes in catchments of poorly weathered rocks. Lakes in the lowlands, however, particularly shallow ones, have proved less tractable to restoration. This study of three lowland lakes provides insights that illuminate a more complex picture. 2. The lakes lie in a sequence along a single stream in a mixed urban and rural landscape. Severely deoxygenating effluent from an overloaded sewage treatment works was diverted from the catchment in 1991. Effects on two lakes, Little Mere (z_max <2 m) and Rostherne Mere (z_max 31 m) were followed until 2002. Mere Mere (z_max = 8 m), upstream of the former works, acted as a comparison for changes in water chemistry. Mere Mere showed no change in total phosphorus (TP), total inorganic nitrogen, or planktonic chlorophyll a concentrations. Increased winter rainfall was associated with higher winter soluble reactive phosphorus (SRP) and ammonium concentrations in its water. 3. Little Mere changed from a deoxygenated, highly enriched, fishless system, with large populations of Daphnia magna Straus, clear water and about 40% aquatic plant cover, to a slightly less clear system following diversion. Daphnia magna was replaced by D. hyalina Leydig as fish recolonised. Spring peaks of chlorophyll a declined but summer concentrations increased significantly. Annual mean chlorophyll a concentrations thus showed no change. Submerged plants became more abundant (up to 100% cover), with fluctuating community composition from year to year. Summer release of SRP from the sediment was substantial and has not decreased since 1993. The summer phytoplankton was apparently controlled by nitrogen availability perhaps with some influence of zooplankton grazing. SRP was always very abundant. The lake appeared to have reached a quasi‐stable state by 2002. 5. Rostherne Mere showed a steady decline in TP and SRP concentrations following effluent diversion apparently as a result of steady dilution by water with lower phosphorus concentration. Decline in phosphorus concentrations was much less rapid than expected because of internal remobilisation from the hypolimnion and sediments. There have been no changes in chlorophyll a concentration or of nitrogen availability and by 2002 the phytoplankton probably remained limited by a combination of mixing, grazing and nitrogen. 6. A seeming paradox is, thus, that immense changes in phosphorus budgets have shown no consequences for phytoplankton chlorophyll concentrations in either of the lakes, although the seasonal distribution has altered in Little Mere. Although these case studies deviate from others, for both shallow and deep lakes, they represent distinctive situations rather than undermining conventional models.     

Changes in a deep lake following sewage diversion – a challenge to the orthodoxy of external phosphorus control as a restoration strategy?

1. The restoration of deep lakes has traditionally focused on reducing the external phosphorus loading. 2. Following the diversion of sewage effluent, that led to marked reductions in nutrient concentrations in its main inflow, Rostherne Mere has shown no reduction in phosphorus or chlorophyll a concentrations. A shallow lake upstream (Little Mere), however, has shown a marked response to effluent diversion. 3. Nutrient budgets for Rostherne Mere reveal that sewage effluent was by far the most significant external source of total phosphorus and that diffuse drainage from the catchment was the most significant external source of dissolved inorganic nitrogen. Phosphorus loads from groundwater and a bird roost were insignificant. Internal sources of phosphorus were, however, considerable and were largely responsible for the observed delay in recovery. 4. Phosphorus limitation of phytoplankton biomass may never be attainable because of substantial internal and diffuse sources of phosphorus, combined with a long retention time. Nitrogen is likely to be more important in limiting phytoplankton biomass. Control of diffuse nitrogen sources may therefore be more effective in the restoration of the deeper lakes of this region.     

Two mesocosm experiments investigating the control of summer phytoplankton growth in a small shallow lake

1. Mesocosm experiments were carried out to examine the relative importance of top down (fish predation) and bottom up (nutrient addition) controls on phytoplankton abundance in a small shallow lake, Little Mere, U.K., in 1998 and 1999. These experiments were part of a series at six sites across Europe. 2. In the 1998 experiment, top‐down processes (through grazing of large Cladocera) were important in determining phytoplankton biomass. The lack of plant refugia for zooplankton was probably important in causing an increasing chlorophyll a concentration even at intermediate fish density. Little Mere normally has abundant macrophytes but they failed to develop substantially during both years. Bottom‐up control was not important in 1998, most probably because of high background nutrient concentrations, as a result of nutrient release from the sediments. 3. In 1999 neither top‐down nor bottom‐up processes were significant in determining phytoplankton biomass. Large cladoceran grazers were absent even in the fish‐free enclosures, probably because dominance of cyanobacteria and high phytoplankton biomass made feeding conditions unsuitable. As in 1998, bottom‐up control of phytoplankton was not important, owing to background nutrient concentrations that were even higher in 1999 than in 1998, perhaps because of the warmer, sunnier weather. 4. The differing outcomes of the two experiments in the same lake with similar experimental designs highlight the importance of starting conditions. These conditions in turn depended on overall weather conditions prior to the experiments.     

The potential of artificial refugia for maintaining a community of large-bodied cladocera against fish predation in a shallow eutrophic lake

The Norfolk Broads are a series of shallow lakes which are highly eutrophic and typified by dense populations of phytoplankton and an absence of submerged aquatic plants. The zooplankton community is subject to intense predation pressure by young fish and is dominated by small-bodied organisms which have a low potential for reducing phytoplankton populations through grazing. Various designs and densities of artificial refugia for zooplankton against fish predation were established in Hoveton Great Broad in order to enhance populations of large-bodied Cladocera. Initially some of the refuges contained higher densities and larger individuals ofDaphnia andCeriodaphnia than the surrounding open water. However, towards the end of the first season and throughout the subsequent two years, population densities and size-structure were similar both within and outside the refuges, although there was still evidence of enhanced body-size ofDaphnia within the refuges compared with the open water. The provision of habitat structures designed as refugia from fish predation did not enhance large-bodied cladoceran populations enough to promote this restoration strategy as feasible for eutrophic and shallow lakes.     

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.     

The relative importance of top-down and bottom-up control of phytoplankton in a shallow macrophyte-dominated lake

1. Mechanisms stabilizing the plant-dominated clear-water state were investigated in Little Mere, U.K. Replicated, factorial, mesocosm experiments, carried out in 1995 and 1996, were designed to investigate the relative importance of top-down (zooplankton grazing) and bottom-up (nitrogen-limitation) control in limiting algal growth, and the role of macrophytes in these processes. Treatments included increased salinity (1995) and sticklebacks (1996) to reduce zooplankton numbers, weekly nitrate additions and removal of macrophytes. 2. Contrary to the results of other studies, submerged plants did not reduce nitrate concentrations. Owing to the high stickleback density in the enclosures with fish, macrophytes did not provide a refuge for zooplankton during the experiment. In Little Mere, however, where fish densities are lower, macrophytes probably play a key role in maintaining clear water by providing refuge for pelagic zooplankton and habitat for attached Cladocera. 3. Phytoplankton in Little Mere was not nitrogen- (N) limited during the growing season. Although nitrogen availability sets a maximum potential phytoplankton biomass it was not realized owing to control by zooplankton grazing.     

Applicability of planktonic biomanipulation for managing eutrophication in the subtropics

Although large-bodied cladocerans such asDaphnia can reduce algal biomass significantly in temperate lakes if freed from fish predation, the applicability of such biomanipulation techniques for eutrophication management in the subtropics and tropics has been examined only recently. Subtropical cladoceran assemblages differ from those of temperate lakes by their low species richness, early summer gameogenesis, and greatly reduced body size. Eutrophic Florida lakes are dominated by pump-filter feeding fish rather than by size selective planktivores as a temperate lakes. Cladocerans in Florida lakes can increase in abundance significantly if freed from fish but fail to have an impact on algal biomass or composition. The greatest potential for using biomanipulation to manage phytoplankton-dominated lakes in the subtropics and tropics lies with phytophagous fish. Future research should concentrate on defining the role of individual fish taxa on phytoplankton composition and community structure, nutrient cycling, and planktonic productivity before embarking on whole lake manipulation projects.     

Controlling Harmful Cyanobacteria: Taxa-Specific Responses of Cyanobacteria to Grazing by Large-Bodied Daphnia in a Biomanipulation Scenario

Lake restoration practices based on reducing fish predation and promoting the dominance of large-bodied Daphnia grazers (i.e., biomanipulation) have been the focus of much debate due to inconsistent success in suppressing harmful cyanobacterial blooms. While most studies have explored effects of large-bodied Daphnia on cyanobacterial growth at the community level and/or on few dominant species, predictions of such restoration practices demand further understanding on taxa-specific responses in diverse cyanobacterial communities. In order to address these questions, we conducted three grazing experiments during summer in a eutrophic lake where the natural phytoplankton community was exposed to an increasing gradient in biomass of the large-bodied Daphnia magna. This allowed evaluating taxa-specific responses of cyanobacteria to Daphnia grazing throughout the growing season in a desired biomanipulation scenario with limited fish predation. Total cyanobacterial and phytoplankton biomasses responded negatively to Daphnia grazing both in early and late summer, regardless of different cyanobacterial densities. Large-bodied Daphnia were capable of suppressing the abundance of Aphanizomenon, Dolichospermum, Microcystis and Planktothrix bloom-forming cyanobacteria. However, the growth of the filamentous Dolichospermum crassum was positively affected by grazing during a period when this cyanobacterium dominated the community. The eutrophic lake was subjected to biomanipulation since 2005 and nineteen years of lake monitoring data (1996–2014) revealed that reducing fish predation increased the mean abundance (50%) and body-size (20%) of Daphnia, as well as suppressed the total amount of nutrients and the growth of the dominant cyanobacterial taxa, Microcystis and Planktothrix. Altogether our results suggest that lake restoration practices solely based on grazer control by large-bodied Daphnia can be effective, but may not be sufficient to control the overgrowth of all cyanobacterial diversity. Although controlling harmful cyanobacterial blooms should preferably include other measures, such as nutrient reductions, our experimental assessment of taxa-specific cyanobacterial responses to large-bodied Daphnia and long-term monitoring data highlights the potential of such biomanipulations to enhance the ecological and societal value of eutrophic water bodies.     

Mesocosm experiments on the interaction of sediment influence, fish predation and aquatic plants with the structure of phytoplankton and zooplankton communities

1. Little Mere, U.K., received large quantities of sewage effluent until 1991, when the effluent was diverted. Experiments, carried out in mesocosms in 1992 and 1993, were designed to predict the effects of: (i) reduced external nutrient loading; (ii) reduced internal loading from the sediment; and (iii) recolonization by fish of the better aerated water. Treatments included isolation of the water from the underlying sediment or exposure to the sediment (which lacked plants in 1992, but was covered by Potamogeton berchtoldii in 1993) crossed with different population densities of Rutilus rutilus in 1992 and of Perca fluviatilis in 1993.
2. Exposure to sediment (as opposed to isolation from it) resulted in no net change in the biovolumes of most major algal groups, but this masked major complementary effects on individual species. The experiments showed a decreasing influence of the sediment, between 1992 and 1993, in determining water chemistry, and an increasing pH between years but no increase in cyanophyte dominance. This had been anticipated because a lake upstream provides abundant inocula, and conditions in Little Mere after diversion of effluent were expected to favour cyanophytes.
3. Roach and perch additions to the mesocosms resulted in reductions in Daphnia populations but increases in numbers of small Cladocera and copepods. Plant-associated Cladocera were unaffected by fish. The presence of submerged plants to some extent reduced fish predation effects on Daphnia hyalina .
4. The experimental results in general accurately predicted subsequent events in the open lake.     

Mesocosm experiments on the interaction of sediment influence, fish predation and aquatic plants with the structure of phytoplankton and zooplankton communities

1. Little Mere, U.K., received large quantities of sewage effluent until 1991, when the effluent was diverted. Experiments, carried out in mesocosms in 1992 and 1993, were designed to predict the effects of: (i) reduced external nutrient loading; (ii) reduced internal loading from the sediment; and (iii) recolonization by fish of the better aerated water. Treatments included isolation of the water from the underlying sediment or exposure to the sediment (which lacked plants in 1992, but was covered by Potamogeton berchtoldii in 1993) crossed with different population densities of Rutilus rutilus in 1992 and of Perca fluviatilis in 1993.
2. Exposure to sediment (as opposed to isolation from it) resulted in no net change in the biovolumes of most major algal groups, but this masked major complementary effects on individual species. The experiments showed a decreasing influence of the sediment, between 1992 and 1993, in determining water chemistry, and an increasing pH between years but no increase in cyanophyte dominance. This had been anticipated because a lake upstream provides abundant inocula, and conditions in Little Mere after diversion of effluent were expected to favour cyanophytes.
3. Roach and perch additions to the mesocosms resulted in reductions in Daphnia populations but increases in numbers of small Cladocera and copepods. Plant-associated Cladocera were unaffected by fish. The presence of submerged plants to some extent reduced fish predation effects on Daphnia hyalina .
4. The experimental results in general accurately predicted subsequent events in the open lake.     

Gape Limitation, Prey Size Refuges and the Top–down Impacts of Piscivorous Largemouth Bass in Shallow Pond Ecosystems

Top–down control of phytoplankton biomass through piscivorous fish manipulation has been explored in numerous ecological and biomanipulation experiments. Piscivores are gape-limited predators and it is hypothesized that the distribution of gape sizes relative to distribution of body depths of prey fish may restrict piscivore effects cascading to plankton. We examined the top–down effects of piscivorous largemouth bass on nutrients, turbidity, phytoplankton, zooplankton and fish in ponds containing fish assemblages with species representing a range of body sizes and feeding habits (western mosquitofish, bluegill, channel catfish, gizzard shad and common carp). The experimental design consisted of three replicated treatments: fishless ponds (NF), fish community without largemouth bass (FC), and fish community with largemouth bass (FCB). Turbidity, chlorophyll a, cyclopoid copepodid and copepod nauplii densities were significantly greater in FC and FCB ponds than in NF ponds. However, these response variables were not significantly different in FC and FCB ponds. The biomass and density of shallow-bodied western mosquitofish were reduced and bluegill body depths shifted toward larger size classes in the presence of largemouth bass, but the biomass and density of all other fish species and of the total fish community were unaffected by the presence of largemouth bass. Our results show that top–down impacts of largemouth bass in ecosystems containing small- and deep-bodied fish species may be most intense at the top of the food web and alter the size distribution and species composition of the fish community. However, these top–down effects may not cascade to the level of the plankton when large-bodied benthivorous fish species are abundant.     

The effects of tench (Tinca tinca (L.)) and sticklebacks (Gasterosteus aculeatus L.) on planktonic and benthic communities in mesocosms in a shallow lake

The effects of introducing a zooplanktivorous fish, three-spined stickleback, (Gasterosteus aculeatus) and a benthivorous fish, tench (Tinca tinca) separately and in combination to replicated experimental enclosures with two density levels of white water lily (Nymphaea alba) were studied in Little Mere, UK. Numbers of Daphnia hyalina were high and only slightly diminished at reduced lily densities, probably due to stickleback predation, but there was no consequential effect on phytoplanktonic chlorophyll a concentrations. Tench reduced the numbers of gastropods but not of other macroinvertebrates, and in turn increased the biomass of periphyton growing on artificial substrata within the enclosures. The higher lily density reduced oxygen concentrations and pH values and increased total phosphorus and soluble reactive phosphorus concentrations but otherwise had little effect on water chemistry. There was little interactive effect of the fish species. The results are integrated with those of six other such enclosure experiments carried out in Little Mere since 1992.     

Ecological and socio‐economic impacts of invasive water hyacinth (Eichhornia crassipes): a review

1. Water hyacinth (Eichhornia crassipes) is one of the world’s most invasive aquatic plants and is known to cause significant ecological and socio‐economic effects. 2. Water hyacinth can alter water clarity and decrease phytoplankton production, dissolved oxygen, nitrogen, phosphorous, heavy metals and concentrations of other contaminants. 3. The effects of water hyacinth on ecological communities appear to be largely nonlinear. Abundance and diversity of aquatic invertebrates generally increase in response to increased habitat heterogeneity and structural complexity provided by water hyacinth but decrease due to decreased phytoplankton (food) availability. 4. Effects of water hyacinth on fish are largely dependent on original community composition and food‐web structure. A more diverse and abundant epiphytic invertebrate community may increase fish abundance and diversity, but a decrease in phytoplankton may decrease dissolved oxygen concentrations and planktivorous fish abundance, subsequently affecting higher trophic levels. 5. Little is known about the effects of water hyacinth on waterbird communities; however, increases in macroinvertebrate and fish abundance and diversity suggest a potentially positive interaction with waterbirds when water hyacinth is at moderate density. 6. The socio‐economic effects of water hyacinth are dependent on the extent of the invasion, the uses of the impacted waterbody, control methods and the response to control efforts. Ecosystem‐level research programmes that simultaneously monitor the effects of water hyacinth on multiple trophic‐levels are needed to further our understanding of invasive species.     

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.     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Impact of fish predation on cladoceran body weight distribution and zooplankton grazing in lakes during winter

1. It is well accepted that fish, if abundant, can have a major impact on the zooplankton community structure during summer, which, particularly in eutrophic lakes, may cascade to phytoplankton and ultimately influence water clarity. Fish predation affects mean size of cladocerans and the zooplankton grazing pressure on phytoplankton. Little is, however, known about the role of fish during winter. 2. We analysed data from 34 lakes studied for 8–9 years divided into three seasons: summer, autumn/spring and winter, and four lake classes: all lakes, shallow lakes without submerged plants, shallow lakes with submerged plants and deep lakes. We recorded how body weight of Daphnia and then cladocerans varied among the three seasons. For all lake types there was a significant positive correlation in the mean body weight of Daphnia and all cladocerans between the different seasons, and only in lakes with macrophytes did the slope differ significantly from one (winter versus summer for Daphnia). 3. These results suggest that the fish predation pressure during autumn/spring and winter is as high as during summer, and maybe even higher during winter in macrophyte‐rich lakes. It could be argued that the winter zooplankton community structure resembles that of the summer community because of low specimen turnover during winter mediated by low fecundity, which, in turn, reflects food shortage, low temperatures and low winter hatching from resting eggs. However, we found frequent major changes in mean body weight of Daphnia and cladocerans in three fish‐biomanipulated lakes during the winter season. 4. The seasonal pattern of zooplankton : phytoplankton biomass ratio showed no correlation between summer and winter for shallow lakes with abundant vegetation or for deep lakes. For the shallow lakes, the ratio was substantially higher during summer than in winter and autumn/spring, suggesting a higher zooplankton grazing potential during summer, while the ratio was often higher in winter in deep lakes. Direct and indirect effects of macrophytes, and internal P loading and mixing, all varying over the season, might weaken the fish signal on this ratio. 5. Overall, our data indicate that release of fish predation may have strong cascading effects on zooplankton grazing on phytoplankton and water clarity in temperate, coastal situated eutrophic lakes, not only during summer but also during winter.     

Changes in the fish and zooplankton communities of Ringsjön,a Swedish lake undergoing man-made eutrophication

During the 20th century Lake Ringsjön has developed from a mesotrophic to a eutrophic lake, and the phytoplankton community has changed from a rather diverse community to a monoculture of blue-green algae. The eutrophication process has accelerated during the last decade. The most important external nutrient loading of today comes from agriculture.Although phosphorus has been shown to be the primary nutrient leading to excessive algal growth in fresh water, several biotic factors — such as interactions between nutrients, phytoplankton, zooplankton and planktivorous fish — may play a decisive role in the occurrence and maintenance of large algal blooms.The aim of this investigation was to study the changes in the fish community of Lake Ringsjön, especially the most dominant planktivores, and the state of the zooplankton community during the seventies. The fish fauna is dominated by cyprinids, especially roach, and there are relatively few predatory fish. During the seventies the mean size of roach decreased, and measurements of the zooplankton community indicated that the predation pressure on zooplankton had increased. The mean sizes of cladocerans such as Daphnia and Bosmina, which were selected for by the planktivorous fish, decreased; the size of the calanoid Diaptomus, which was not preyed upon by the dominating fish species, did not change. The growth of zooplankton-feeding stages of several fish species was retarded, which meant that the growth of young perch decreased, while older roach were mainly affected. In the prevailing situation, planktivorous roach can maintain a numerous population of small individuals, whereas the predatory perch is at a disadvantage, and predation on zooplankton is intense.     

Control mechanisms of arctic lake ecosystems: a limnocorral experiment

To assess the potential impact of human exploitation on arctic lakes and to determine how these eco systems are regulated we initated a limnocorral experiment in Toolik Lake, Alaska, in the summer of 1983. The limnocorrals were 5 m in diameter and from 5–6 m in depth and were open to the sediments. In 1983 four limnocorrals were deployed in an isolated bay of Toolik Lake within a cross-classified treatment regime of high and low inorganic nitrogen and phosphorus additions and high and low free swimming fish additions. The objective of the nutrient addition was to stimulate phytoplankton growth and determine the extent to which increased plant production was passed through pelagic and benthic food chains. The objective of the fish addition was to determine the impact of fish predation on large-bodied zooplankton, especially the zooplanktivorous copepod Heterocope, then to study the effect of altered Heterocope densities on small-bodied zooplankton species population dynamics. In 1984 two more limnocorrals were deployed, one a low fish, 1 × nutrient addition treatment and the other a no fish, no nutrient treatment. The fish manipulation was changed to confining several fish in cages with the cages held in corrals for varying lengths of time. The addition of inorganic nitrogen and phosphorus dramatically increased phytoplankton productivity. This increase in algal biomass and production greatly altered the light environment and water quality in the nutrient treated limnocorrals. The secchi disk depth in the nutrient treated limnocorrals declined each summer reaching as low as 1 m in 1985. Both oxygen content and pH increased in the nutrient treatment corrals. Corrals not receiving nutrient additions remained near lake concentrations for most water quality parameters. While phytoplankton biomass was stimulated in 1983 phytoplankton growth was not sufficient to draw down all the nitrogen and phosphorus added and these nutrients reached high levels in the last half of the summer. In 1984 phosphorus remained above 20 μg in the nutrient-treated corrals but ammonia dropped to reference levels by day 25. In 1985 both nutrient concentrations rapidly declined to reference levels. Most pelagic components responded to the nutrient additions. Microbial production was stimulated in the nutrient treated limnocorrals and bacterial population sizes built up to nearly 8–10 times those of the reference corrals. However, microheterotrophs soon increased in abundance and apparently grazed down bacteria to reference levels. Phytoplankton population density, as estimated by chlorophyll a determinations, increased dramatically with nutrient addition such that each year the phytoplankton densities were higher than before. Primary productivity was also stimulated and appeared not to be light limited even when phytoplankton densities rose to high levels. In the first two years of the experiment zooplankton densities were little altered by the increased phytoplankton densities. However, by 1985 daphnid densities were quite a bit higher in the high nutrient addition limnocorrals. The benthic community and sediment response was much less affected by nutrient addition. Overall sediment respiration increased in the nutrient treated corrals but underlying sediments seemed little affected. Decomposition of Carex litter was likewise little affected by nutrient addition. Benthic invertebrates were also little impacted by the nutrient addition and increased sedimentation of phytoplankton. However, the response of benthic invertebrates is difficult to assess fully in the current experiment because chironomids, a prominent component of the benthic community, failed to recruit into the limnocorrals and the corrals physically shifted during ice-out in the spring of 1984 disturbing the sediment in several corrals. The fish additions in 1983 of free swimming grayling essentially eliminated large bodied zooplankton, especially Heterocope septentrionalis, from all four limnocorrals. In subsequent summers Heterocope were not so dramatically preyed upon but generally were found in higher densities in the low or no fish treatments. However, either when Heterocope were eliminated in 1983 or were in rough inverse proportion to fish density, altered Heterocope abundance had no obvious affect on small-bodied zooplankton abundance. The fish treatment apparently influenced the zooplankton response to high nutrient addition in 1985. In the high nutrient limnocorrals daphnid populations became very abundant, but in the high fish treatment the daphnid responding was the small-bodied D. longiremis while in the low fish treatment the daphnid responding was the large-bodied D. middendorffiana. Thus we have considerable evidence for bottom up control of phytoplankton density and production. This increased production ultimately, but not for two years, stimulated zooplankton density increases. Increased nutrients had little effect on the benthos or sediments. Fish manipulations influenced large-bodied zooplankton but had little effect on small-bodied zooplankton. Because grayling are predominantly plankton feeders in lakes, no fish effect on benthic invertebrates was expected. Limnocorrals thus seem good systems to study nutrient-phytoplankton interactions. They are not as suitable for benthic invertebrate studies and fish manipulations may be difficult. Most other limnocorral studies were of brief duration; however, in the present study the limnocorrals seemed to perform well over a three year period.     

Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.