Effectiveness of phytoplankton control by large-bodied and small-bodied zooplankton

Employingin situ enclosures containing inocula of the lake zooplankton (mainlyDaphnia galeata, Daphnia cucullata andBosmina spp.) from a moderately eutrophic Lake Ros (Northern Poland) or large-bodiedDaphina magna, the following observations on succession of phytoplankton were made: 1) whereasD. magna could control the density of all the photoplankton size classes, the lake zooplankton could not suppress the large-sized phytoplankters or net phytoplankton; 2) the lake zooplankton was able to control the density of small algae (< 50μm), but its effect on large algae may be opposite: a promotion of net phytoplankton growth by removing small-sized algae which can out-compete net phytoplankton for limited PO₄-P resources (<5μg P l⁻¹). Since efficiency of phytoplankton density control byD. magna decreased with an increase in net phytoplankton abundance, biomanipulation could not be successful without introducing or maintaining a high population of large-bodied cladoceran species before high densities of large algae would make the control of phytoplankton inefficient.     

COMPARISON OF IN SITU PHOTOSYNTHETIC ACTIVITY OF EPIPHYTIC,EPIPELIC AND PLANKTONIC ALGAL COMMUNITIES IN AN OLIGOTROPHIC LAKE,SOUTHERN FINLAND1

The photosynthetic activity of different algal communities at the outer edge of an Equisetum fluviatile L. stand in an oligotrophic lake (Pääjärvi, in southern Finland) was investigated. Production by the algal communities was measured simultaneously using a modified ¹⁴C-method, and the results were related to the volume of algae and the available irradiance. The relative production rate (P/B quotient) of phytoplankton was ca. 3 × that of epiphyton and ca. 20 × that of epipelon. Epiphyton productivity remained almost constant although the algal volume varied greatly, suggesting that the surface layer of the algal community was mainly responsible for the photosynthetic activity. In the littoral area (at 1 m depth) primary production/m² of lake surface by phytoplankton, epiphyton and epipelon was similar but in the littoriprofundal area (2–4 m) phytoplankton production was twice that of epipelon. Primary productivity of epiphyton and epipelon/m² of substratum was about equal to phytoplankton productivity/m³ of water at the same irradiance. This relation provided a means of estimating the relative contributions of the different algal communities to the total algal production in the lake.     

COMBINATION OF FLOW CYTOMETRY AND SINGLE CELL ABSORPTION SPECTROSCOPY TO STUDY THE PHYTOPLANKTON STRUCTURE AND TO CALCULATE THE CHL A SPECIFIC ABSORPTION COEFFICIENTS AT THE TAXON LEVEL1

The phytoplankton community structure of a hypertrophic lake was quantitatively determined with the aid of flow cytometry. The flow cytometry signals were calibrated to obtain cell‐specific information, such as the chl a content and the biovolume per cell. The reliability of this method was tested with laboratory cultures. The results of the phytoplankton structure in a hypertrophic lake with respect to chl distribution in the different algal groups obtained by flow cytometry were compared with the results from HPLC pigment fingerprinting. Both methods yield the percentage contribution of the different algal groups to total chl a. The chl a specific absorption coefficient of the phytoplankton (a^*_Phy) was determined via visible (VIS) spectroscopy of samples taken from a hypertrophic lake (Auensee) in 2003. The results indicated that a^*_Phy of the total cell suspension is dependent on the phytoplankton structure as well as on environmental factors. The linear relationship between a^*_Phy at 675 nm and the product of the chl a content per cell and the biovolume offered the possibility to normalize phytoplankton absorption spectra to acquire the taxon‐specific a^*_Phy. The estimated a^*_Phy (675 nm) values were used to normalize single cell absorption spectra at this wavelength to obtain the a^*_Phy between 400 and 750 nm for representatives of the major algal groups. Our measurements show that the absorption coefficient for the whole phytoplankton community varies within the season. Finally, we used the a^*_Phy and the chl a distribution to calculate the light absorption of each algal group in the hypertrophic lake.     

Long-term phytoplankton changes in a shallow hypertrophic lake,Albufera of Valencia (Spain)

A long-term phytoplankton study was carried out in the Albufera of Valencia, a shallow hypertrophic lake (surface area 21 km², mean depth 1 m, total inorganic nitrogen load 155 g m^-2 y^-1, total inorganic phosphate load 15 g m^-2 y^-1) from 1980 to 1988. The lake functions as a reservoir for the surrounding rice cultivation. From 1940's to 1988, its phytoplankton assemblage has been altered from a mesotrophic to a hypertrophic character, as consequence of the increasing pollution. For 1980–88, annual variations in the phytoplankton were less pronounced than seasonal changes. The hypertrophic and morphometric features of the lake favoured the stability of the phytoplankton assemblage and chlorophyll a levels during the study period. Seasonal and horizontal distribution of the total phytoplankton abundance and biomass were highly influenced by the hydrological cycle of the lagoon. Compared with other shallow nutrient rich lakes, the Albufera of Valencia is similar to the shallow hypertrophic lakes of the Netherlands.     

Disturbance events affecting phytoplankton biomass,composition and species diversity in a shallow,eutrophic, temperate lake

The seasonal changes in phytoplankton biomass and species diversity in a shallow, eutrophic Danish lake are described and related to different disturbance events acting on the phytoplankton community.Both the spring diatom maximum and the summer bloom of the filamentous blue-green alga, Aphanizomenon flos-aquae (L.) Ralfs, coincided with low values of phytoplankton species diversity and equitability. Diatom collapse was mainly due to internal modifications as nutrient depletion (Si, P) caused by rapid growth of phytoplankton, and increased grazing activity from zooplankton. A large population of Daphnia longispina O.F. Müller in June effectively removed smaller algal competitors, thus favouring the development of a huge summer bloom (140 mm³ l^–1) of Aphanizomenon flos-aquae. Heavy rainfall and storms in late July increased the loss of Apahnizomenon by out-flow and disturbed the stratification of the lake. These events caused a marked decline in phytoplankton biomass but had no effect on species diversity. A second storm period in late August circulated the lake completely and was followed by a rapid increase in phytoplankton diversity, and a change in the phytoplankton community structure from dominance of large, slow-growing K-selected species (Aphanizomenon) to small, fast-growing r-selected species (cryptomonads).     

Can suspension feeding by bivalves regulate phytoplankton biomass in Lake Waccamaw, North Carolina?

Suspension feeding by bivalves has been hypothesized to control phytoplankton biomass in shallow aquatic ecosystems. Lake Waccamaw, North Carolina, USA is a shallow lake with a diverse bivalve assemblage and low to moderate phytoplankton biomass levels. Filtration and ingestion rates of two relatively abundant species in the lake, the endemic unionid, Elliptio waccamawensis, and an introduced species, Corbicula fluminea, were measured in experiments using natural phytoplankton for durations of 1 to 6 days. Measured filtration and ingestion rates averaged 1.78 and 1.121 ind.^–1 d^–1, much too low to control phytoplankton at the observed phytoplankton biomass levels and growth rates. Measured ingestion rates averaged 4.80 and 1.50 µg chlorophyll a ind.^–1 d^–1, too low to support individuals of either species. The abundance of benthic microalgae in Lake Waccamaw reaches 200 mg chlorophyll a m^–2 in the littoral zone and averages almost an order of magnitude higher than depth-integrated phytoplankton chlorophyll a. Total microalgal biomass in the lake is therefore not controlled by suspension feeding by bivalves.     

Whole-lake food-web manipulation as a means to study community interactions in a small ecosystem

Whole-lake food-web manipulation was carried out in the hypertrophic Lake Zwemlust (The Netherlands), with the aim of studying the effects on the lake's trophic status and to gain an insight into complex interactions among lake communities. Before manipulation this small (1.5 ha) and shallow (1.5 m) lake was characterized byMicrocystis blooms in summer and high chlorophyll-a concentrations were common (ca. 250 μg 1⁻¹). In March 1987 the planktivorous and benthivorous fish species in the lake were completely removed (ca. 1000 kg ha⁻¹), a new simple fish community (pike and rudd) was introduced and artificial refuges were created. The effects of this manipulation on the light climate, nutrient concentrations, phytoplankton, zooplankton, fish, macrophytes, and macrofauna were monitored during 1987, 1988 and 1989. Community interactions were investigated in phytoplankton bioassays and zooplankton grazing experiments. After the manipulation, despite the still high P and N loads to the lake (ca. 2.2 g P m⁻² y⁻¹ andca. 5.3 g N m⁻² y⁻¹), the phytoplankton density was low (Chl-a<5μg l⁻¹), due to control by large-sized zooplankton in spring and N-limitation in summer and autumn. A marked increase in the abundance of macrophytes and filamentous green algae in 1988 and 1989, as well as N loss due to denitrification, contributed to the N limitation of the phytoplankton. Before manipulation no submerged macro-vegetation was present but in 1988, the second year after manipulation, about 50% of the lake bottom was covered by macrophytes increasing to 80% in 1989. This led to substantial accumulation of both N and P, namely 76% and 73% respectively of the total nutrients in the lake in particulate matter. Undesirable features of the increase in macrophytes were: 1) direct nuisance to swimmers; and, 2) the large scale development of snails, especiallyL. peregra, which may harbour the parasite causing ‘swimmers' itch’. But harvesting of only about 3% of the total macrophyte biomass from the swimmers' area, twice a year, reduced the nuisance for swimmers without adversely affecting the water clarity.     

Interactions between Phytoplankton,Zooplankton, and Fish in the Nutrient Rich Shallow Lake Hjarbæk Fjord,Denmark

In the shallow, nutrient-rich dammed-up lake Hjarbæk Fjord the balance between phytoplankton, zooplankton, and fish was completely disturbed during the summer period. Extensive growth of small blue-green algae (Oscillatoria cfr. trichoides, Pseudanabaena galeata) and chlorococcal green algae (Scenedesmus spp., Monoraphidium contortum, Actinastrum hantzschii) induced a pH rise to 10.5 in July. Fish were driven into the rivers, and many that did not escape died. After the fish had disappeared daphniae multiplied explosively and grazed away the major part of the phytoplankton. Ammonia concentrations rose and oxygen concentrations dropped to noxious levels for highly sensitive fish in August–September. Larger phytoplankton species (Aphanizomenon flos-aquae, Pediastrum boryanum, Pandorina morum, a. o.) were not grazed by the daphniae and gave rise to a maximum of copepods in September. Fish returned to the lake in October, when pH, ammonia, and oxygen levels had normalized.     

Bottom-up effects of bream (Abramis brama L.) in Lake Balaton

Enclosures (17 m³) were used in the mesotrophic area of Lake Balaton to determine the impact of benthivorous bream (Abramis brama L.) on the lower trophic levels during summers of 1984–86. In enclosures with a fish biomass similar to the biomass in the eutrophic area of the lake, the number of phytoplankton species was highest. In enclosures with a low fish biomass the phytoplankton was dominated by the greens. A high biomass of bream in the mesotrophic basin caused bacterial production corresponding to that of the eutrophic part of the lake. Crustaceans were dominated by copepods and were unable to control phytoplankton peaks. Bottom-up effects of bream were more obvious than top-down effects and seem to be more important in the possible control of water quality.     

The trophic state of Lake Ladoga as indicated by late summer phytoplankton

As a part of the joint Russian-Finnish evaluation of human impact on Lake Ladoga, we studied the phytoplankton of the lake in order to find biological indicators for eutrophication. A second aim of the investigation was intercalibration of sampling and phytoplankton counting techniques between the Russian and Finnish laboratories. Phytoplankton samples were collected from 27 sampling stations in the lake and from the rivers Volkhov and Neva in 9–13 August 1993. In surface water samples the phytoplankton fresh weight biomass varied in the range 218–3575 mg m^–33. Highest biomass values were encountered in Sortavala Bay, and lowest ones in the western central part of the lake. Phytoplankton species composition varied considerably in the lake; blue-green and green algae predominated near-shore areas and Cryptophyceae in the offshore stations. Canonical correspondance analysis revealed close grouping of eutrophy indicating communities, dominated mainly by greens and blue-greens, in the most nutrient-rich parts of Lake Ladoga, the Volkhov and Svir Bays. Samples from the vicinity of the inflows of Vuoksi and Burnaya Rivers and off Pitkaranta formed a separate group, dominated by diatoms, most of which were typical to mesotrophic or eutrophy lakes. As judged by phytoplankton biomass values and chlorophyll a concentrations, Lake Ladoga may generally be classified as mesotrophic. Eutrophicated areas are found in the northern archipelago of the lake and in the areas influenced by large rivers.     

Why biomanipulation can be effective in peaty lakes

The effects of fish stock reduction (biomanipulation) was studied in an 85 ha shallow peaty turbid lake. The lake cleared in a 4-week period in April–May 2004, which demonstrated that biomanipulation can be effective in peaty lakes. We demonstrated that it is possible to reduce the fish stock to <25 kg ha⁻¹ benthivorous fish and <15 kg ha⁻¹ planktivorous fish, sufficiently low to switch the lake from a turbid to a clear state. Knowledge of lake morphology, fish stock, fish behaviour, and a variety of fishing methods was necessary to achieve this goal. It is expected that continuation of fisheries to remove young of the year planktivorous species is needed for several years, until macrophytes provide sufficient cover for zooplankton and can compete with phytoplankton. Cladocerans developed strongly after fish removal. The clearing of the lake coincided with a sudden decrease of filamentous cyanobacteria and suspended detritus, and a strong increase of Bosmina. We assume that Bosmina was able to reduce filamentous prokaryotes and detritus. After the disappearance of the cyanobacteria, Bosmina disappeared too. After the clearing of the lake Daphnia dominated in zooplankton and apparently was able to keep phytoplankton levels low. In our case, wind resuspension did not prevent biomanipulation from being successful. No correlation between windspeed and turbidity was found, neither in an 85 ha nor in a 230 ha shallow peaty lake. Regression analysis showed that on average 50% of the amount of suspended detritus can be explained by resuspension by fish and 50% by phytoplankton decomposition. The main goal of this biomanipulation experiment, clear water and increased submerged plant cover in a shallow peaty lake, was reached.     

The Seasonal Dynamics and Trophic Relations of the Plankton Components in Lake Peipsi (Peipus)

The seasonal dynamics of the biomass and production of phyto-, zoo- and bacterioplankton was investigated during the vegetation periods (from May to November) in 1985 and 1986 in the pelagial of the large eutrophic lake Peipsi (Estonia). The average values of productions per vegetation period for the investigation years were as follows: phytoplanktion − 203.5 gC · m⁻²; bacterioplankton − 37.9 gC · m⁻²; filter-feeding zooplankton − 20.6 gC · m⁻² and predatory zooplankton − 1.5 gC · m⁻². The herbivorous zooplankton production constituted 10.1% of primary production. This ratio indicates a direct relationship between zoo- and phytoplankton in the food chain — filtrators are feeding mostly on living algae and the detrital food chain seems of little importance. The dominance of large forms (Melosira sp., Aphanothece saxicola), in the phytoplankton during the major part of the vegetation period is assumed to be a result of high grazing pressure on small algae. Zooplankton grazing was investigated in situ in a specially constructed twin bathometer. Experimental measurements revealed, that zooplanktion presence in the experimental vessel actually stimulated the phytoplankton growth in many cases — the negative grazing values have been registered. That could be caused by the stimulation effect of nutrients (N, P), excreted by the concentrated zooplankton in the grazing chamber, which led to an increase of the nongrazed phytoplankton production. Bacteria have satisfied the zooplankton food requirements on average by 11%. Grazing on bacteria increased, when grazing on phytoplankton was somehow disturbed.     

The recovery of a very shallow eutrophic lake, 20 years after the control of effluent derived phosphorus

1. Monitoring at fortnightly to monthly intervals of a very shallow, lowland lake over 24 years has enabled the time course of recovery from nutrient enrichment to be investigated after high external P loading of the lake (>10 g P m^?2 year^?1) was reduced between 1977 and 1980. 2. The lake showed a relatively rapid response during the spring and early summer, with a reduction in phytoplankton biomass occurring after 5 years when soluble reactive phosphorus concentration was <10 μg L^?1. 3. However, during the later summer the response was delayed for 15 years because of sustained remobilisation of phosphorus from the sediment. The greater water clarity in spring and a gradual shift from planktonic to benthic algal growth may be related to the reduction in internal loading after 15 years. 4. Changes in the phytoplankton community composition were also observed. Centric diatoms became less dominant in the spring, and the summer cyanobacteria populations originally dominated by non‐heterocystous species (Limnothrix/Planktothrix spp.) almost disappeared. Heterocystous species (Anabaena spp. and Aphanizomenon flos‐aquae) were slower to decline, but after 20 years the phytoplankton community was no longer dominated by cyanobacteria. 5. There were no substantial changes in food web structure following re‐oligotrophication. Total zooplankton biomass decreased but body size of Daphnia hyalina, the largest zooplankton species in the lake, remained unchanged, suggesting that the fish population remained dominated by planktivorous species. 6. Macrophyte growth was still largely absent after 20 years, although during the spring water clarity may have become sufficient for macrophytes to re‐establish.     

Cyanobacteria and cyanotoxins in the source water from Lake Chivero,Harare, Zimbabwe,and the presence of cyanotoxins in drinking water

The phytoplankton community and cyanotoxins in Lake Chivero (formerly Lake McIlwaine) and the presence of cyanotoxins in treated drinking water were investigated between 2003 and 2004. A typical seasonal succession of Cyanobacteria species occurred from January to April, Bacillariophyta from May to July, and Cryptophyta and Chlorophyta from August to December. Microcystis aeruginosa and M. wesenbergii, known producers of the toxin microcystin, and the non-toxic cyanobacterium M. novacekii dominated during summer. The highest concentrations of microcystins and lipopolysaccharide endotoxins occurred when cyanobacterial biomass was highest. Lipopolysaccharide endotoxin concentrations in the lake ranged between 8 and 3 200 Endotoxin Units (EU) ml^?1. Microcystin concentrations in treated water were below the recommended safe limit for drinking water. Lipopolysaccharide endotoxin concentrations in treated water ranged from 0.15 to 11 EU ml^?1. The phytoplankton community comprised non-microcystin-producing species for the greater part of the study period.     

Seasonal dynamics of phytoplankton and phytobenthos,and associated chemical interactions,in a shallow upland lake (Malham Tarn,northern England)

Seasonal changes of phytoplankton were followed over 3 years (1985–87) in a shallow, unstratified and calcareous upland lake.The phytoplankton was of low to moderate abundance and generally dominated by phytoflagellates. Seasonality involved a winter minimum of abundance, a spring maximum of diatoms, and often brief increases in summer that included blue-greens, especially the colonial Gloeotrichia echinulata. Some components were of benthic origin. Seasonal growth of the main component of the phytobenthos, Chara globularisvar. virgata, caused a regular summer depletion in lake water of Ca²⁺ and HCO₃ ^- (alkalinity) by associated CaCO₃ deposition, and a more extreme (and unusual) depletion of K⁺. Chemical analysis of Chara biomass and of underlying sediments indicated a large benthic nutrient stock, much surpassing that represented by the phytoplankton. Growth in this biomass, and the magnitude of water-borne inputs, influenced the removals of Ca²⁺, K⁺ and inorganic N. The phytoplankton was probably limited by a low-P medium, to which co-precipitation of phosphate with CaCO₃ may have contributed. A vernal depletion of Si was probably limiting to diatom growth, and appeared to be mainly induced by benthic rather than planktonic diatoms. Examples of long-term change in composition of the phytoplankton and phytobenthos are noted and discussed in relation to the interaction of these components, nutrient enrichment, and possible alternative stable states.     

Macrophyte-related shifts in the nitrogen and phosphorus contents of the different trophic levels in a biomanipulated shallow lake

Lake Zwemlust, a small highly eutrophic lake, was biomanipulated without reducing the external nutrient loading, and the effects were studied for four years. In this paper we pay special attention to the shifts in relative distribution of nitrogen and phosphorus in the different trophic levels and to the changes in growth limitation of the autotrophs.Despite of the high external nutrient loads to the lake (ca 2.4 g P m^–2 y^–1 and 9.6 g N m^–2 y^–1), the effects of biomanipulation on the lake ecosystem were pronounced. Before biomanipulation no submerged vegetation was present in the lake and P and N were stored in the phytoplankton (44% N, 47% P), fish (33% N, 9% P) and in dissolved forms (23% N, 44% P). P and N contents in sediments were not determined. In the spring and summer following the biomanipulation (1987), zooplankton grazing controlled the phytoplankton biomass and about 90% of N and P were present in dissolved form in the water. From 1988 onwards submerged macrophyte stands continue to thrive, reducing the ammonium and nitrate concentrations in the water below detection levels. In July 1989 storage of N and P in the macrophytes reached 86% and 80%, respectively. Elodea nuttallii (Planchon) St.John, the dominant species in 1988 and 1989, acted as sink both for N and P during spring and early summer, withdrawing up to ca 60% of its N and P content from the sediment. At the end of the year only part of the N and P from the decayed macrophytes (ca 30% of N and 60% of P) was recovered in the water phase of the ecosystem (chiefly in dissolved forms). The rest remained in the sediment, although some N may have been released from the lake by denitrification.In summer 1990 only 30% of the N and P was found in the macrophytes (dominant species Ceratophyllum demersum L.), while ca 30% of N and P was again stored in phytoplankton and fish.     

Zooplankton allochthony is spatially heterogeneous in a boreal lake

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Phytoplankton productivity in findley lake

Findley Lake is a dimictic, oligotrophic, subalpine lake located in the western Cascade Mountains, Washington. The lake is snow covered for most of the year so that the growing season was 3.5 months in 1971 and 4.5 months in 1972. Rapid melt of the lake's snow cover in summer allowed the sudden development of a phytoplankton productivity maximum (as measured by the ¹⁴C tracer method) of 86 mg m⁻² hr⁻¹ and a peak of 48 mg chlorophyll a per m¹ within two weeks of surface clearing in 1972, followed by a rapid decline of productivity and biomass. Annual production (between 10 October, 1971 and 21 October, 1972) was 36 g/m² in the 27.5 m water column. Autotrophic carbon assimilation during the snow-covered period was insignificant. The total production for the lake in 1972 was 530 kg carbon. The concentration of available nitrogen (NO₂ + NO₃ + NH₃ as N) at 15 m ranged from 12 to 76 mg/m³ while PO₄-P ranged from 0.5 to 8.3 mg/m³. In vitro nutrient enrichment experiments with natural phytoplankton communities from the lake indicated that while N and P together were growth limiting, P alone produced a growth response while N alone did not. Contributions to production from net-, nanno-, and ultraplankton were determined by fractional filtration of ¹⁴C-labeled phytoplankton samples. The nannoplankton, dominated by diatoms, accounted for 58% to 94% of productivity.     

Ecology of Dictyosphaerium pulchellum Wood (Chlorophyta, Chlorococcales) in a shallow, acid, forest lake

This study relates to the ecology of Dictyosphaerium pulchellum Wood in Delamere Lake in Cheshire, UK. Dictyosphaerium pulchellum is a cosmopolitan, green colonial phytoplankton species that occasionally forms dense, monospecific populations in lakes. Delamere Lake is a small, shallow, acid lake (mean pH, 4.5) with very high phytoplankton biomass (annual mean chlorophyll a, 290 μg l⁻¹) and devoid of any significant cladoceran population, the efficient grazers of phytoplankton. A predominantly unicellular form of D. pulchellum was the dominant species in Lake Delamere, and it comprised on average ca. 80% (maximum >99%) of the lake phytoplankton biovolume. Laboratory and lake experiments were conducted on this species showed that its pH tolerance varied between 2.4 and 10.7, and its optimum tolerance range between 3.3 and 8.5 depending on other environmental variables. Low pH was not responsible for the unicellular habit of this alga, but a very high nutrient regime could be an important factor. Bioassays revealed that in Delamere Lake this species was limited by nitrogen, but nitrogen did not hamper high growth in the lake. Dictyosphaerium pulchellum can persist at low light levels, tolerate CO₂-deficiency and can grow in polyhumic water with water colour around 300 mg Pt l⁻¹, but probably not in darker waters. The dominance of D. pulchellum in Delamere Lake is apparently due to a combination of several factors: its ability to tolerate both low pH and high turbidity, exploit high nutrient conditions, absence of effective grazing pressure by zooplankton and being a superior competitor.     

Factors regulating summer phytoplankton in a highly eutrophic Antarctic lake

Lakes from Maritime Antarctica are regarded as systems generally inhabited by metazoan plankton capable of imposing a top-down control on the phytoplankton during short periods, while lakes from Continental Antarctica lacking these communities would be typically controlled by scarcity of nutrients, following a bottom-up model. Otero Lake is a highly eutrophic small lake located on the NW of the Antarctic Peninsula, which has no metazoan plankton. During summer 1996, we studied the density, composition and vertical distribution of the phytoplankton community of this lake with respect to various abiotic variables, yet our results demonstrated neither light nor nutrient limitation of the phytoplankton biomass. Densities of heterotrophic nanoflagellates (HNAN) and ciliates from three different size categories were also studied. Extremely low densities of HNAN (0–155 ind. ml^–1) could be due to feeding competition by bacterivore nanociliates and/or predation by large ciliates. A summer bloom of the phytoflagellate Chlamydomonas aff. celerrima Pascher reached densities tenfold those of previous years (158.10³ ind. ml^–1), though apparently curtailed by a strong peak of large ciliates (107 ind. ml^–1) which would heavily graze on PNAN (phototrophic nanoflagellates). Top-down control can thus occur in this lake during short periods of long hydrologic residence time.