Predators, resources, and trophic chains in the regulation of plankton population and biomass in oligothrophic lakes

The relative strength of "top-down" versus "bottom-up" control of plankton community structure and biomass in two small oligotrophic lakes (with and without fish), located near the Polar circle (Russia), has been investigated for two years, 1996 and 1997. The comparative analyses of zooplankton biomass and species abundance showed strong negative effect of fish, stickeback (Pungitius pungitius L.), on the zooplankton community species, size structure and biomass of particular prey species but no effect on the biomass of the whole trophic level. An intensive predation in Verkhneye lake has lead to: 1) sixfold decline in biomass of large cladoceran Holopedium gibberum comparing to the lake lacking predator, 2) shift in the size mode in zooplankton community and the replacement of the typical large grazers by small species--Bosmina longirostris and rotifers. Their abundance and biomass even increased, demonstrating the stimulating effect of fish on the "inefficient" and unprofitable prey organisms. The analysis of contributions of different factors into the cladoceran's birth rate changes was applied to demonstrate the relative impact of predators and resources on zooplankton abundance. An occasional introduction of the stickleback to Vodoprovodnoye lake (the reference lake in 1996) in summer 1997 lead to drastic canges in this ecosystem: devastating decrease of zooplankton biomass and complete elimination of five previously dominant grazer species. The abundance of edible phytoplankton was slightly higher in the lake with fish in 1996 and considerably higher in the lake where fish has appeared in 1997 showing the prevailing "top-down" control of phytoplankton in oligotrophic ecosystem. The reasons of trophic cascade appearance in oligotrophic lakes are also discussed.     

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No previous study of plankton in lakes has estimated the relative contribution of ciliated protozoa to the biomass of the total plankton community, including phytoplankton. In a series of south-central Ontario lakes, ciliates comprise on the order of 5 to 10% of the total planktonic biomass of these relatively oligotrophic lakes and exist there in densities of 20–40 ml⁻¹. Therefore, ciliates constitute an important component of lake ecosystems that should not be ignored in limnological studies of zooplankton abundance and distribution.     

Predictability and possible mechanisms of plankton response to reduction of planktivorous fish

The predictability of plankton response to reductions of planktivorous fish was investigated by comparing the plankton community in three biomanipulated lakes and ten unmanipulated lakes differing in intensity of fish predation. Data collected on total phosphorus, phytoplankton and zooplankton biomass and share of cyanobacteria and large grazers, as well as specific growth rate of phytoplankton, were further used to test some of the proposed underlying response-mechanisms. In the biomanipulated lakes the algal biomass and share of cyanobacteria decreased, specific growth rate of phytoplankton increased, and zooplankton biomass and share of large grazers increased or remained unchanged. This pattern was largely reflected in the differences in food-chain structure between the unmanipulated lakes with highversus those with low fish predation. The qualitative response to planktivorous fish reduction thus seems largely predictable. The biomanipulated lakes differed, however, in magnitude of response: the smallest hypertrophic, rotenone-treated lake (Helgetjern) showed the most dramatic response, whereas the large, deep mesotrophic lake (Gjersjøen), which was stocked with piscivorous fish, showed more moderate response, probably approaching a new steady state. These differences in response magnitude may be related to different perturbation intensity (rotenone-treatmentversus stocking with piscivores), food-chain complexity and trophic state. Both decreased phosphorus concentration and increased zooplankton grazing are probably important mechanisms underlying plankton response to biomanipulation in many lakes. The results provide tentative support to the hypothesis that under conditions of phosphorus limitation, increased zooplankton grazing can decrease algal biomassvia two separate mechanisms: reduction of the phosphorus pool in the phytoplankton, and reduction of the internal C:P-ratio in the phytoplankton cells.

     

Phytoplankton structure in different lake types in central Finland

Phyloplankton structure and its relation to physical and chemical properties of the water was studied in 58 central Finnish lakes. The biomass ranged from 0.2 to 14.2 g m⁻³ and the number of taxa per sample ranged from 33 to 152. The lakes were grouped into 5 types according to their trophic state: eutrophic, dyseutrophic, mesotrophic, oligotrophic, and acid oligotrophic lakes. The average biomass in eutrophic lakes was 5.57 g m⁻³, in dyseutrophic 3.54 g m⁻³, 1.23 g m⁻³ in mesotrophic, 0.52 g m⁻³ in oligotrophic and 0.39 g ⁻³ in acid oligotrophic lakes. The average number of taxa per sample in the corresponding lake types were 109. 1, 79.3, 97.9, 90.9 and 43.8, respectively. The phytoplankton communities in eutrophic lakes were characterized by blue-green algae (21.2% of total biomass) and green algae (18.7% of total biomass). In dyseutrophic lakes the proportion of green algae was much smaller (7.2% of total biomass) than in eutrophic lakes, whereas the proportion of diatoms and cryptophytes was higher (28.2 and 20.4% of total biomass, respectively). Chrysophytes dominated in the oligotrophic and mesotrophic lakes (27.3–39.9% of total biomass). The contribution of dinoflagellates to the total biomass was highest in the most oligotrophic acidified lakes and in those lakes the relative proportions of blue-green and green algae were much higher than in the typical oligotrophic lakes. The lakes were also grouped into 8 community types according to the dominating algal group. Cyanophyceae- and Chlorophyceae-types characterized the eutrophic lakes, whereas Chrysophyceae-Dinopheceae-type was typical for most oligotrophic lakes. The other 5 types occurred in mesotrophic and oligotrophic lakes but the physical and chemical properties of these lakes did not differ much.     

A key role of aluminium in phosphorus availability,food web structure,and plankton dynamics in strongly acidified lakes

We studied extracellular acid phosphatase activity (AcPA) of planktonic microorganisms, aluminium (Al) speciation, and phosphorus (P) cycling in three atmospherically acidified (pH of 4.5–5.1) mountain forest lakes: ?ertovo jezero (CT), Prá?ilské jezero (PR), and Ple?né jezero (PL) in the Bohemian Forest (?umava, Böhmerwald). Microorganisms dominated pelagic food webs of the lakes and crustacean zooplankton were important only in PR, with the lowest Al concentrations (193 µg L^?1) due to 3–4 times lower terrestrial input. The lakes differed substantially in Al speciation, i.e., in the proportion of ionic and particulate forms, with the highest proportion of ionic Al in the most acid CT (pH = 4.5). The P concentration in the inlet of PL (mean: 22.9 µg L^?1) was about five times higher than in CT and PR (3.9 and 5.1 µg L^?1, respectively). Average total biomass of planktonic microorganisms in PL (593 µg C L^?1) was, however, only ～2-times higher than in CT and PR (235 and 272 µg C L^?1, respectively). Enormous AcPA (means: 2.17–6.82 µmol L^?1 h^?1) and high planktonic C : P ratios suggested severe P limitation of the plankton in all lakes. Comparing 1998 and 2003 seasons, we observed changes in water composition (pH and Al speciation) leading to a significant increase in phytoplankton biomass in the lakes. The increase in the seston C : P ratio during the same time, however, indicates a progressive P deficiency of the lakes. The terrestrial Al inputs, together with in-lake processes controlling the formation of particulate Al, reduced P availability for planktonic microorganisms and were responsible for the differences in AcPA. At pH < 5, moreover, ionic Al forms caused inhibition of extracellular phosphatases. We postulate that both particulate and ionic Al forms affect P availability (i.e., inhibition of extracellular phosphatases and inactivation of P), specifically shape the plankton composition in the lakes and affect plankton recovery from the acid stress.     

Phosphatase activity in relation to phytoplankton composition and pH in Swedish lakes

SUMMARY. 1. Potential phosphatase activity and phytoplankton from several lakes of different character were compared in order to evaluate the importance of lake water pH and phytoplankton composition for the activity and pH optimum of lake water phosphatases.
2. In oligotrophic lakes, in which phytoplankton biomass was most often dominated by Ochromonadaceae spp., optimum phosphate activity was found at pH values <6. In eutrophic lakes, where species of Cyanophyceae and Bacillariophyceae dominated the phytoplankton biomass, optimum phosphatase activity was found at pH 7.5 or 8.5.
3. The pH optimum of phosphatase activity often differed from the corresponding lake water pH.
4. Experimental variation in phosphorus availability resulted in predictable changes in phosphatase activity. However, specific phosphatase activity, calculated per biomass of phytoplankton, was dependent on plankton species composition.     

Zooplankton biomass dynamics in oligotrophic versus eutrophic conditions: a test of the PEG model

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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.     

Two types of maternal care for juveniles observed in Caprella monoceros Mayer, 1890 and Caprella decipiens Mayer, 1890 (Amphipoda: Caprellidae)

Spatial and seasonal patterns in phytoplankton and zooplankton communities of Lake St. Clair from June through September, 1984 are described. Phytoplankton biomass averages 586 µg l^-1 with the Diatomae and Chrysophyceae predominating. Zooplankton biomass averages 663 µg l^- with small bosminid Cladocera being the most abundant organisms. Lake St. Clair zooplankton biomass is second only to that of Lake Erie amongst the St. Lawrence Great Lakes. Biomass size spectra are typical in structure for mesotrophic lakes but low explained variance in the annual normalized spectrum is indicative of a perturbed system. Since 1972/1973 there appears to have been a slight decrease in zooplankton abundance in the lake accompanied by a shift from dominance of rotifers to dominance of cladocerans. We hypothesize that high flushing rate and seasonal variability coupled with contaminant loadings have resulted in a plankton community reduced in taxonomic diversity and dominated by small-bodied species.     

Cascading effects of generalist fish introduction in oligotrophic lakes

Introduction of strictly planktivorous fish to lakes can alter plankton communities via cascading interactions in food webs. Less is known about the large-scale and long-term effects resulting from the introduction of fish with more generalist feeding habits, and the extent to which these effects depend on lake trophic status. Paleolimnological records of three oligotrophic lakes in Maine, USA were used to analyze the response of plankton communities to the introduction of white perch (Morone americana), a fish that often numerically dominates fish assemblages and switches from strict planktivory to a more generalist diet during ontogeny. After white perch introduction, cladoceran ephippia size increased up to 50 %, suggesting that the most important role of this generalist fish, with respect to water quality, is as a piscivorous trophic link. Algal standing crop declined by a quarter to over a half of pre-introduction levels, suggesting that top-down effects of white perch reduced phytoplankton biomass. In comparing these oligotrophic lakes to prior work in a eutrophic system, white perch introduction had similar effects on zooplankton body size; however, cascading effects to phytoplankton were only observed in low productivity lakes.     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

The role of plankton, zoobenthos, and sediment in organic matter degradation in oligotrophic and eutrophic mountain lakes

Intensity of organic matter degradation, assessed by the respiratory electron transport system (ETS) activity, was studied in microplankton, zooplankton, chironomid larvae as the dominant group of the macrobenthos, and sediment in mountain lakes of different trophic levels in summer months. The highest ETS activities per unit of surface were observed in sediments. Significantly lower activities were observed in microplankton, and lower still in zooplankton, and chironomids. The total ETS activity m^–2 was higher in eutrophic lakes (Jezero na Planini pri Jezeru and Krnsko jezero) than in oligotrophic ones (Zgornje Kriko jezero, Spodnje Kriko jezero, Jezero v Ledvicah). The contributions of communities investigated to total ETS activity m^–2 differed between lakes of different trophic level. Estimation of respiratory carbon loss through different components revealed that the most of the organic matter was oxidized in sediments of mountain lakes. The respiratory carbon losses were higher through zooplankton than through microplankton in all lakes. Carbon losses through plankton components and sediments were significantly lower in oligotrophic than in eutrophic lakes. The contribution of respiratory carbon loss through chironomids to total carbon loss m^–2 was higher in oligotrophic than in eutrophic lakes. Therefore, it seems that contributions of microplankton and zooplankton to mineralization processes increase, and contributions of chironomids and sediment surface decrease with increasing trophic level of the lakes.     

Poor nutritional quality of primary producers and zooplankton driven by eutrophication is mitigated at upper trophic levels

Eutrophication and rising water temperature in freshwaters may increase the total production of a lake while simultaneously reducing the nutritional quality of food web components. We evaluated how cyanobacteria blooms, driven by agricultural eutrophication (in eutrophic Lake Köyliöjärvi) or global warming (in mesotrophic Lake Pyhäjärvi), influence the biomass and structure of phytoplankton, zooplankton, and fish communities. In terms of the nutritional value of food web components, we evaluated changes in the ω‐3 and ω‐6 polyunsaturated fatty acids (PUFA) of phytoplankton and consumers at different trophic levels. Meanwhile, the lakes did not differ in their biomasses of phytoplankton, zooplankton, and fish communities, lake trophic status greatly influenced the community structures. The eutrophic lake, with agricultural eutrophication, had cyanobacteria bloom throughout the summer months whereas cyanobacteria were abundant only occasionally in the mesotrophic lake, mainly in early summer. Phytoplankton community differences at genus level resulted in higher arachidonic acid, eicosapentaenoic acid (EPA), and docosahexaenoic acid (DHA) content of seston in the mesotrophic than in the eutrophic lake. This was also reflected in the EPA and DHA content of herbivorous zooplankton (Daphnia and Bosmina) despite more efficient trophic retention of these biomolecules in a eutrophic lake than in the mesotrophic lake zooplankton. Planktivorous juvenile fish (perch and roach) in a eutrophic lake overcame the lower availability of DHA in their prey by more efficient trophic retention and biosynthesis from the precursors. However, the most efficient trophic retention of DHA was found with benthivorous perch which prey contained only a low amount of DHA. Long‐term cyanobacterial blooming decreased the nutritional quality of piscivorous perch; however, the difference was much less than previously anticipated. Our result shows that long‐term cyanobacteria blooming impacts the structure of plankton and fish communities and lowers the nutritional quality of seston and zooplankton, which, however, is mitigated at upper trophic levels.     

The Contribution of Ciliated Protozoa to Zooplankton Biomass in an Acidic, Subtropical Lake

Ciliated protozoa accounted for up to 50% of the mean daily zooplankton biomass in McCloud Lake, a small (5 ha), oligotrophic, acidic (pH 4.7) lake in north-central Florida. Food resources (algae and bacterioplankton) were limiting for crustacean and rotifer zooplankton during much of the year. Myxotrophic ciliates were a dominant component of the planktonic food web. Stentor niger , an uncommon species in the plankton of lakes, dominated the ciliate assemblage and usually comprised >90% of total ciliate biomass. Stentor niger always contained high densities of photosynthetic zoochlorellae and contributed an estimated 30% to the total autotrophic biomass.     

<Emphasis Type="Italic">Coelastrum pascheri</Emphasis> sp. n., a new green alga from lakes of the Bohemian Forest

Coelastrum pascheri sp. n. is described. It is similar to Coelastrum morus W. et G. S. West sensu Skuja (1930) in the morphology of the free processes, but differs from it in cell size (it is significantly smaller, 13–16 µm) and in the number of cells in a cenobium (usually 4). The new alga was found in the littoral of some lakes: ?erné jezero (?erné Lake), Grosser Arbersee and Kleiner Arbersee in the Bohemian Forest (?umava, Böhmerwald) and also in Sphagnum within a small puddle in the Upolínová meadow in the Slavkovský les Mts.     

Phosphorus decrease and climate variability: mediators of synchrony in phytoplankton changes among European peri-alpine lakes

1. In an attempt to discern long‐term regional patterns in phytoplankton community composition we analysed data from five deep peri‐alpine lake basins that have been included in long‐term monitoring programmes since the beginning of the 1970s. Local management measures have led to synchronous declines in phosphorus concentrations by more than 50% in all four lakes. Their trophic state now ranges from mesotrophic to oligotrophic. 2. No coherence in phytoplankton biomass was observed among lakes, or any significant decrease in response to phosphorus (P)‐reduction (oligotrophication), except in Lakes Constance and Walen. 3. Multivariate analyses identified long‐term changes in phytoplankton composition, which occurred coherently in all lakes despite the differing absolute phosphorus concentrations. 4. In all lakes, the phytoplankton species benefiting from oligotrophication included mixotrophic species and/or species indicative of oligo‐mesotrophic conditions. 5. A major change in community composition occurred in all lakes at the end of the 1980s. During this period there was also a major shift in climatic conditions during winter and early spring, suggesting an impact of climatic factors. 6. Our results provide evidence that synchronous long‐term changes in geographically separated phytoplankton communities may occur even when overall biomass changes are not synchronous.     

Lake Chapo: a baseline study of a deep, oligotrophic North Patagonian lake prior to its use for hydroelectricity generation: II. Biological properties

The present paper focuses on the biological properties of Lake Chapo prior to the installation of a hydroelectric power station. Most of the biological parameters indicate that the lake is oligotrophic, i.e., it has very low values of chlorophyll a and primary production. The phytoplankton community of 73 species shows a predominance of only a few species in terms of density and biomass. The zooplankton community is poor in species, with one of the lowest densities measured in the North Patagonian lakes. The fish community is dominated by two introduced salmonid species. Comparison with other North Patagonian lakes reveals that Lake Chapo is the most oligotrophic.     

Spatial patterns and relationships between phytoplankton, zooplankton and water quality in the Saimaa lake system, Finland

The interactions between phytoplankton and zooplankton were studied in two large lakes in the Saimaa lake system, Finland. Both are subjected to substantial waste water loading, and exhibit a clear trophic gradient between the loaded and unloaded areas. The phytoplankton and zooplankton were compared in terms of composition, abundance and biomass at 34–39 stations located in different parts of the lakes. At least four mechanisms were thought to affect the composition of plankton communities: (1) the amount of nutrients (trophic gradient), (2) grazing of algae by herbivores, (3) the effect of the algal species composition on feeding by zooplankters (large, colonial algae in the more loaded parts of the lakes) and (4) the regeneration and reorganization of nutrients.     

PICOPHYTOPLANKTON BIOMASS IN RELATION TO LAKE TROPHIC STATE AND THE TN:TP RATIO OF LAKE WATER IN JAPAN1

Picophytoplankton biomass and its contribution to total phytoplankton biomass were investigated in relation to the nutrient concentration and total N: total P ratio of the epilimnetic waters of 42 Japanese lakes during the warm season in 1991 (April–October). Picophytoplankton biomass (as chlorophyll a) in meso-, eu-, and hypertrophic lakes was significantly higher than those observed in oligotrophic lakes. However, picophytoplankton biomass increased significantly with increased total P concentrations in all systems excluding hypertrophic lakes. Picophytoplankton contribution to total chlorophyll a content was significantly higher in oligo- and mesotrophic lakes than in eu- and hypertrophic lakes and was inversely correlated with total P concentrations in lake water. Picophytoplankton contribution to the total phytoplankton biomass was positively (r = 0.54, n = 42, P = 0.0003) correlated with the total N: total P ratio of lake waters. Each lake trophic type, with the exception of hypertrophic lakes, showed this trend, although the correlation was not significant. We suggest that picophytoplankton contribution is influenced by the total N: total P ratio rather than lake trophic state; however, picophytoplankton were of little importance in hypertrophic lakes.     

Stable isotope analysis of the origins of zooplankton carbon in lakes of differing trophic state

Carbon stable isotope analysis was carried out on zooplankton from 24 United Kingdom lakes to examine the hypothesis that zooplankton dependence on allochthonous sources of organic carbon declines with increasing lake trophy. Stable isotope analysis was also carried out on particulate and dissolved organic matter (POM and DOM) and, in 11 of the lakes, of phytoplankton isolates. In 21 of the 24 lakes, the zooplankton were depleted in ¹³C relative to bulk POM, consistent with previous reports. δ¹³C for POM showed relatively little variation between lakes compared to high variation in values for DOM and phytoplankton. δ¹³C values for phytoplankton and POM converged with increasing lake trophy, consistent with the expected greater contribution of autochthonous production to the total organic matter pool in eutrophic lakes. The difference between δ¹³C for zooplankton and that for POM was also greatest in oligotrophic lakes and reduced in mesotrophic lakes, in accordance with the hypothesis that increasing lake trophic state leads to greater dependence of zooplankton on phytoplankton production. However, the difference increased again in hypertrophic lakes, where higher δ¹³C values for POM may have been due to greater inputs of ¹³C-enriched organic matter from the littoral zone. The very wide variation in phytoplankton δ¹³C between lakes of all trophic categories made it difficult to detect robust patterns in the variation in δ¹³C for zooplankton. Received: 2 November 1998 / Accepted: 3 December 1999