Vertical distribution of benthic community responses to fish predators, and effects on algae and suspended material

The vertical positioning of benthic invertebrates should be a trade-off between the risky, but productive, sediment surface and the safer, but physiologically harsher, conditions deeper down in the sediment. This is because the foraging efficiency of benthic fish decreases with sediment depth, whereas the sediment surface is generally better oxygenated and has a higher resource quality than lower layers. We studied how two benthic fish predators, bream (Abramis brama) and ruffe (Gymnocephalus cernuus), affected the community composition and vertical distribution of benthos, and their indirect effects on algae and suspended material, in field enclosures. Whereas bream had significant effects on the density, composition and distribution of the benthos, ruffe had no such effects. The total benthos biomass in bream treatments was an-order of magnitude lower in the upper sediment layer (0–1 cm) and three times lower in the middle layer (1–3 cm) than in the controls, whereas there were no significant effects in the deepest layer (3–10 cm). Bivalves persisted in the deepest layer although their density was reduced in shallow sediment, whereas gastropods faced the risk of local extinction in the presence of bream. As indirect effects, small-bodied cladocerans, phytoplankton, periphyton and both organic and inorganic suspended material were higher in the bream treatments. We␣conclude that the impact of bream diminished substantially with increasing sediment depth, enabling invertebrates to survive in the sediment and to persist in the presence of bream. However, there were␣no␣indications of any group adjusting their vertical position behaviourally as a response to predation threat.     

Effects of nutrients and grazers on periphyton phosphorus in lake enclosures

SUMMARY. 1. Periphyton. measured as particulate phosphorus (PP) and expressed as periphyton PP, growing on vertically oriented substrata (polyvinyl impregnated nylon) under different nutrient loadings, light intensities (exposures), and grazer communities was examined in eight large enclosures (750 m3) where nutrients (N and P) and planktivorous fish (1+yellow perch) were added in a 2x2 factorial design.
2. During the first 3 weeks of the experiment (25 June to 15 July), there was a significantly higher accumulation of phosphorus into periphyton (periphyton PP) with fertilization, but fish addition had no effect. During the fourth to seventh weeks (16 July to 12 August), addition of fish was associated with lower abundance of amphipods and chironomids and higher concentration of periphyton PP. In the enclosures without fish, these invertebrates were over 25 times more abundant, and periphyton PP decreased substantially compared to the June-July period. Fertilization increased periphyton PP only at high exposures in the enclosures with fish.
3. Exposure had a significant effect on periphyton PP. In the enclosures with fish, high abundance of nanoplankton reduced water transparency, and periphyton PP was lower in the deeper waters which may have been due to limitation by low light. Lower periphyton PP was also observed at the surface on sunny sides of enclosures without fish, and therefore with high water transparency. This pattern may have been due to inhibitory effects of high light intensity.
4. Periphyton communities in the enclosures with fish had higher uptake rates for planktonic phosphorus, and lower rates of phosphorus release, suggesting that periphyton with high phosphorus demand may have high internal cycling of assimilated phosphorus.     

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.     

鲤对浅水湖泊水质影响的围隔实验研究

鲤是我国浅水湖泊常见鱼类,鲤可以通过扰动底泥、牧食浮游动物、排泄营养盐等途径影响湖泊水质。为了研究鲤对浅水湖泊水质的影响,论文于2009年6月23日至8月18日在暨南大学明湖进行原位围隔实验。实验分为三个处理组(三组围隔):(1)在围隔中放养鲤(放鱼组);(2)在围隔底部放置塑料隔网,在隔网上部空间放养鲤(放鱼放网组);(3)围隔中不放隔网也不放鲤(空白对照组)。实验结果表明:鲤可以显著增加水体悬浮物、总氮、总磷和叶绿素a含量,显著降低了水体透明度。而鲤是否扰动底泥对水体悬浮物、总氮、总磷和叶绿素a含量等指标的影响不显著。 因此,该研究中鲤对浅水湖泊的影响可能主要是通过排泄等途径实现的,鲤扰动底泥对水质的影响不大。     

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.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Context-dependent effects of freshwater mussels on stream benthic communities

1. We asked whether unionid mussels influence the distribution and abundance of co‐occurring benthic algae and invertebrates. In a yearlong field enclosure experiment in a south‐central U.S. river, we examined the effects of living mussels versus sham mussels (shells filled with sand) on periphyton and invertebrates in both the surrounding sediment and on mussel shells. We also examined differences between two common unionid species, Actinonaias ligamentina (Lamarck 1819) and Amblema plicata (Say 1817). 2. Organic matter concentrations and invertebrate densities in the sediment surrounding mussels were significantly higher in treatments with live mussels than treatments with sham mussels or sediment alone. Organic matter was significantly higher in the sediment surrounding Actinonaias than that surrounding Amblema. Actinonaias was more active than Amblema and may have increased benthic organic matter through bioturbation. 3. Living mussels increased the abundance of periphyton on shells and the abundance and richness of invertebrates on shells, whereas effects of sham mussels were similar to sediment alone. Differences in the amount of periphyton growing on the shells of the two mussel species reflected differences in mussel activity and shell morphology. 4. Differences between living and sham mussel treatments indicate that biological activities of mussels provide ecosystem services to the benthic community beyond the physical habitat provided by shells alone. In treatments containing live mussels we found significant correlations between organic matter and chlorophyll a concentrations in the sediment, organic matter concentrations and invertebrate abundance in the sediment and the amount of chlorophyll a on the sediment and invertebrate abundance. There were no significant correlations among these response variables in control treatments. Thus, in addition to providing biogenic structure as habitat, mussels likely facilitate benthic invertebrates by altering the availability of resources (algae and organic matter) through nutrient excretion and biodeposition. 5. Effects of mussels on sediment and shell periphyton concentrations, organic matter concentrations and invertebrate abundance, varied seasonally, and were strongest in late summer during periods of low water volume, low flow, and high water temperature. 6. Our study demonstrates that freshwater mussels can strongly influence the co‐occurring benthic community, but that effects of mussels are context‐dependent and may vary among species.     

Top-down and bottom-up impacts of juvenile fish in a littoral reed stand

1. Research has often focused on pelagic food chains and processes of lakes; less is known about the contribution of benthic energy flows to whole‐lake ecosystem energetics. This stems from the fact that the shoreline and littoral habitats, which provide a key linkage between sediment and water column, have only recently become a significant focus for study. 2. This study aimed to quantify the feeding and phosphorus allocation of a juvenile fish community in a littoral zone of a shallow lake in response to the biomass succession of the invertebrate prey community. Habitats comprising reed and adjacent open water were sampled over two consecutive years during day and night. 3. Although there were substantial year‐to‐year differences in the biomass of invertebrates, the fish community composition, diet consumption rates and phosphorus allocations were very similar in both study years. Biomasses and predation impacts by juvenile fish on prey groups were substantially higher within the reeds than in the adjacent open water habitat. This may be explained by the refuge‐seeking behaviour of the fish. 4. In general, invertebrates were negligibly influenced by fish feeding, with the exception for a strong top‐down control of large cladocerans. In response to the resulting low Daphnia biomass, fish were forced to switch to a higher degree of benthivory. Consequently, juvenile fish in littoral reed stands may shift benthic‐derived energy and phosphorus via the excretion of soluble reactive phosphorus into the open water.     

Fish induced macrophyte loss in shallow lakes: top–down and bottom–up processes in mesocosm experiments

SUMMARY 1. Macrophyte loss from Sites of Special Scientific Interest in England has become widespread over the last 20 years. One reason for this may be changing trends in angling, a multimillion pound industry that has an enormous impact on aquatic ecosystems. Stocking with cyprinid fish is a common angling management practice but the particular fish species and distribution of their biomass may be crucial to the ecosystem. 2. Carp (Cyprinus carpio), roach (Rutilus rutilus), bream (Abramis brama) and tench (Tinca tinca) at biomasses ranging from 0 to 800 kg ha^?1 and at various sizes were placed into experimental mesocosms in Little Mere, a shallow, fertile lake in Cheshire, U.K. The effects these treatments had on the aquatic ecosystem were studied over two summers. Specifically the effects of the treatments on macrophyte growth, benthic and macrophytic macro‐invertebrate populations, water chemistry, epiphyton production and plankton survival were investigated. 3. Carp had a greater detrimental effect on the macrophytes than bream, tench and in particular roach. A biomass of fish > 200 kg ha^?1 adversely affected the extent of macrophyte growth. 4. The decline in macrophyte growth was most likely as a result of increased epiphyton growth that probably reduced the amount of light and carbon dioxide available to the plant. There were no observed direct fish impacts on macrophytes. 5. The chemical data suggested that inorganic nitrogen levels were low and it is possible that release of nitrogen, from fish excreta, followed by immediate uptake, could have been a major factor stimulating epiphyton growth and subsequently macrophyte loss. Phosphorus concentrations increased even in the controls and substantial amounts were available. Phosphorus stimulation can therefore be discounted. Macrophyte‐associated macro‐invertebrates were positively correlated with epiphyton load but had no impact on the extent of epiphytic growth. Shading from disturbed sediment or phytoplankton was also unimportant.     

Biomass dependent interactions in pond ecosystems: responses of lower trophic levels to fish manipulations

The effects of mature benthivorous cyprinid fish and theirrecruitment on sediment resuspension, turbidity, phyto- andzooplankton, and benthic macroinvertebrates were studied in fourexperimental ponds. The ponds were stocked with bream (Abramisbrama L.), white bream (Blicca bjoerkna L.), roach (Rutilus rutilus L.) and wild carp (Cyprinus carpio L.) of3+–5+ age classes at standing crop biomass varying from 0 to500 kg ha-1. Cyprinids caused an increase in sedimentresuspension and in turbidity, in proportion to their biomass. Meancrustacean biomass did not significantly affect phytoplanktonbiomass due to intense grazing by fish during spring. Ponds withhigh fish stocks showed reduced midge biomass and vegetation coverand increased biomass of predatory invertebrates.     

Resuspension of algal cells by benthivorous fish boosts phytoplankton biomass and alters community structure in shallow lakes

1. Positive effects of fish on algal biomass have variously been attributed to cascading top‐down effects and to nutrient enrichment by fish excretion. 2. Here, we used a combination of field and laboratory approaches to test an additional hypothesis, namely that the physical resuspension of settled algal cells by fish enhances algal biomass and alters community composition. 3. A multi‐lake survey showed that phytoplankton biomass and the fraction of motile algae increased with the concentration of inorganic suspended solids. This correlation could not be explained by wind‐induced resuspension because of the small size of the lakes. 4. In an enclosure experiment, chlorophyll‐a concentration, phytoplankton abundance and inorganic suspended solids increased significantly in the presence of Cyprinus carpio (common carp), but only if the fish had access to the sediment. No such effects were seen when a net prevented carp reaching the sediment. 5. The effects of enhanced nutrients and reduced zooplankton grazing as a result of fish feeding could not (fully) explain these observations, suggesting that the resuspension by carp of settled algae from a surface film on the sediment was the major factor in the outcome of the experiment. 6. An increase in diatoms and green algae (organisms with a relatively large sedimentation velocity) only in enclosures where carp could reach the sediment supported this view. 7. Several lines of evidence indicate that fish‐induced resuspension of algal cells from the sediment is an important mechanism that affects phytoplankton biomass and community composition in shallow lakes.     

Ecological responses to simulated benthic-derived nutrient subsidies mediated by omnivorous fish

1. Fish can play an important role in coupling benthic and pelagic habitats by consuming benthic prey and providing essential nutrients to algae in dissolved form. However, little is known about the factors affecting the magnitude of this nutrient subsidy. 2. Using laboratory and mesocosm experiments we evaluated how varying ingestion rates of bluegill sunfish (Lepomis macrochirus) affects fish excretion rates of both nitrogen (N) and phosphorus (P). During the 10‐week mesocosm experiment, we also evaluated how varying ingestion rates may affect plankton community dynamics, and nutrient flux between pelagic and benthic habitats. Lastly, bioenergetic/mass balance models were used to examine the nutrient stoichiometry of fish body composition and excretion products. 3. Under laboratory conditions, both N and P excretion rates increased with increased ingestion of benthic prey surrogates (earthworms). This effect was more pronounced for N than P. Furthermore, under the more realistic conditions of the mesocosm experiment ingestion rate had no significant effect on P excretion rate. 4. Increased fish ingestion rate in the mesocosm experiment increased total algal biomass and the flux of nutrients from the water column to sediments. Effects of variable ingestion were much stronger on periphyton biomass and algal sedimentation rates than on phytoplankton or zooplankton biomass or composition. 5. Fish body nutrient composition was greatly affected by ingestion rate. N content increased and P content decreased with ingestion rate. As a result, the N : P ratio of fish bodies also increased with ingestion rate. The N : P ratio of nutrients excreted by fish also increased with ingestion rate, counter to predictions of stoichiometric theory, which predicts that excreted N : P ratio is negatively correlated to body N : P. However, this finding can be explained by relaxing the assumption of constant nutrient assimilation rates, and our mass balance data suggest that assimilation rates vary indeed with ingestion rate. 6. Our study provides experimental evidence that translocation of benthic‐derived nutrients by fish can affect the flux of nutrients among habitats, while also suggesting that stoichiometry models need to better incorporate how variable ingestion rates affect nutrient assimilation and excretion rates.     

Effects of macrograzers and light on periphyton stoichiometry

Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.     

Facilitation of clear-water conditions in shallow lakes by macrophytes: differences between charophyte and angiosperm dominance

A number of mechanisms result in a feedback between water clarity and macrophytes and, consequently, the occurrence of alternative stable states in shallow lakes. We hypothesize that bottom-up mechanisms and interactions within the benthic food web are more important in a charophyte-dominated clear-water state, while top-down mechanism and interactions in the planktonic food web prevail at angiosperm dominance. Charophytes, which dominate at lower nutrient concentrations and develop higher densities than most angiosperms, can have a higher influence on sedimentation, resuspension, and water column nutrients. During dominance of dense submerged vegetation like charophytes, zooplankton can be hampered by low food quality and quantity and by high predation pressure from juvenile fish, which in turn are favoured by the high refuge potential of this vegetation. Grazing pressure from zooplankton on phytoplankton can therefore be low in charophytes, but the main feedback in angiosperm-dominated ecosystems. Charophytes offer a higher surface than most angiosperms to periphyton, which favors benthic invertebrates. These support macrophytes by grazing periphyton and constitute a central link in a trophic cascade from fish to periphyton and macrophytes. To test these hypotheses, more experiments and field measurements comparing the effect of charophytes and angiosperms on water clarity are needed.     

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.     

Fish foraging effects on benthic assemblages along a warm-temperate stream: differences among drift feeders, benthic predators and grazers

In streams, physical and biotic conditions change from the headwaters to the mouth, shaping longitudinal patterns in community structure. We examined how fish foraging effects on periphyton and benthic invertebrates changed along a longitudinal gradient of a warm-temperate stream in southwestern Japan. We established three study sites according to changes in the composition of fish feeding guilds (upper site characterized by drifting-invertebrate feeders, Oncorhynchus masou ; middle site by benthic invertebrate feeders, Rhinogobius spp.; lower site by the presence of periphyton grazers, Sicyopterus japonicus ), and performed two manipulative experiments to examine effects of different fish assemblages on periphyton and benthic invertebrate abundances. Results of an exclosure experiment suggested that fishes had no effect on the benthic assemblages at the upper and middle sites whereas fishes reduced the abundances of both periphyton and invertebrates on stone surfaces at the lower site, where both benthic invertebrate feeders and grazers inhabited. A subsequent enclosure experiment showed that the reduction of invertebrate densities at the lower site was caused by the grazers rather than benthic invertebrate feeders. These experimental results suggested that effects of fishes on benthic assemblages are intensified downstream, owing to the occurrence of the grazing fish. Furthermore, observational data based on field sampling suggested that such grazing effects were reflected in longitudinal patterns in periphyton and invertebrate abundances. Overall results emphasize an important role of the grazing fish ( S. japonicus ) in shaping longitudinal patterns in benthic assemblage structure.     

Interactive effects of nutrients and contaminants from pulp mill effluents on riverine benthos

Bleached kraft mill effluents (BKME) contain both nutrients(nitrogen and phosphorus) and contaminants that can have a stimulatoryor inhibitory effect on riverine food webs. For the Northern RiverBasins Study (NRBS), separation of the effects of nutrients andcontaminants in BKME was an important issue because field biomonitoringwas unable to isolate these impacts. We addressed this problem by usingriverside mesocosm experiments, along with field observations, todetermine the effects of BKME on benthic food webs of the upperAthabasca River near Hinton, Alberta, Canada. Response variablesincluded periphyton biomass, benthic insect density and composition, andthe biomass of abundant invertebrates (i.e., caddisfly, mayfly andstonefly taxa). Experiments were conducted during autumn when nutrientenhancement was hypothesized to increase autochthonous primaryproduction as a result of low river discharge and high water clarity.Treatments included reference river water, 1% BKMEconcentrations, and nutrients (nitrogen plus phosphorus) equivalent tothe 1% BKME treatment. Although pulp mill effluents can inducesublethal toxicity in benthic biota, BKME concentrations equivalent tolevels in the Athabasca River did not cause measurable toxicity.Relative to reference water treatments, BKME-associated increases inavailable phosphorus produced enrichment effects including increases inperiphyton and insect biomass, and invertebrate abundance. Insect familyrichness was not affected by the treatments. The development andapplication of a field-based artificial stream system for the NRBSimproved our mechanistic understanding of the effects of BKME on benthicbiota of nutrient and contaminant stressors. By combining artificialstream results with field observations, we were also able to link thismechanistic understanding of stressor effects directly to impacts in theriver. We conclude that the response to BKME in the Peace-AthabascaBasin is largely one of nutrient enrichment rather than that oftoxicity.     

Importance of sediments in understanding nutrient cyclings in lakes

Inorganic and organic nutrients are continuously transported to lake bottoms by sedimentation. By various biological, physical, chemical and mechanical processes quantities of certain nutrients can be brought back to the free water again. This cycling between the sediments and water may occur according to various schemes dependent on lake type and bottom conditions. Lake morphology, temperature regimes, trophic level and sediment type can all strongly influence the size of nutrient pools and rates of turnover.The various activities of bacteria, benthic algae, macrophytes, benthic invertebrates and fish, in conjunction with influences of temperature, pH-values, Eh-values, water content, organic matter and elemental sediment composition, lead to the extremely complex nature of nutrient cycling. Three essential components of aquatic ecosystems are discussed, namely carbon, nitrogen and phosphorus.The objective of this paper is to illustrate in condensed form the heterogeneous nature of nutrient cycling processes. In addition, the importance of sediments in understanding nutrient cycling is discussed from a water management perspective.     

Periphyton-macroinvertebrate interactions in light and fish manipulated enclosures in a clear and a turbid shallow lake

In a clear and a turbid freshwater lake the biomasses of phytoplankton, periphytic algae and periphytonassociated macrograzers were followed in enclosures with and without fish (Rutilus rutilus) and four light levels (100%, 55%, 7% and < 1% of incoming light), respectively. Fish and light affected the biomass of primary producers and the benthic grazers in both lakes. The biomass of primary producers was generally higher in the turbid than the clear lake, and in both lakes fish positively affected the biomass, while shading reduced it. Total biomass of benthic grazing invertebrates was higher in the clear than in the turbid lake and the lakes were dominated by snails and chironomids + ostracods, respectively. While light had no effect on the biomass of grazers in the clear lake, snail breeding was delayed in the most shaded enclosures and presence of fish reduced the number of snails and the total biomass of grazers. In the turbid lake ostracod abundance was not influenced by light, but was higher in fish-free enclosures. Density of chironomids correlated positively with periphyton biomass in summer, while fish had no effect. Generally, light-mediated regulation of primary producers was stronger in the turbid than in the clear lake, but the regulation did not nambiguously influence the primary consumers. However, regulation by fish of the benthic grazer community was stronger in the clear than in the turbid lake, and in both lakes strong top-down effects on periphyton were seen. The results indicate that if present-day climate in Denmark in the future is found in coastal areas at higher latitudes, the effect of lower light during winter in such areas will be highest in clear lakes, with typically lower fish biomass and higher invertebrate grazer density.     

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

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