Seasonal dynamics of crustacean zooplankton, heterotrophic nanoflagellates and bacteria in a shallow, eutrophic lake

1. The seasonal development of crustacean zooplankton, heterotrophic nanoflagellates (HNF) and bacteria was examined in Grosser Binnensee, a shallow, eutrophic lake in northern Germany. The grazing impact of Daphnia on bacteria and nanoflagellates was estimated from field data on population abundances and from clearance rates obtained in laboratory experiments. 2. The seasonal succession of zooplankton showed distinct peaks of Daphnia magna, cyclopopid copepods, Bosmina longirostris and Daphnia galeata and D. hynlina. The population dynamics of Dapfinia had the strongest impact on all sestonic components. Daphnia maxima coincided with clearwater phases, and were negatively correlated with particulate organic carbon (POC), HNF and phytoplankton. Bacterial abundance was only slightly affected although daphnids were at times more important as bacterial consumers than HNF, as estimated from measured bacterial clearance rates. Other crustaceans (copepods, Bosmina) were probably of minor importance as grazers of bacteria and nanoplankton. 3. HNF abundance varied from 550 ml^?1 to more than 30000 ml^?1. HNF appeared to be suppressed by daphnids and reached highest densities when copepods dominated the metazooplankton. The variation in HNF abundance was not reflected in the concentration of heterotrophic bacteria, which fluctuated rather irregularly between 5 and 20 ± 10⁶ ml^?1. Long filamentous bacteria which were probably resistant to protozoan grazing, however, appeared parallel to the development of HNF. These bacterial cells, although small in number, could comprise more than 30% of the total bacterial biomass.     

Particle selection over a broad size range by crustacean zooplankton

1. Preferences for particles over a broad size range were assessed in four crustacean zooplankton species under field conditions. The particles employed were six spherical algae with diameters of 2, 6, 13, 19, 38 and 61 μm. 2. Diaphanosoma brachyurum and Daphnia cucullata preferred the two smallest particles, Chydorus sphaericus the three smallest and Eudiaptomus gracilis the 19 μm particle. 3. These size preferences agree well with earlier determinations based on seston particles in the same lake, indicating consistent size preferences by these species over a broad size range.     

Habitat selection and diel distribution of the crustacean zooplankton from a shallow Mediterranean lake during the turbid and clear water phases

1. The fish fauna of many shallow Mediterranean Lakes is dominated by small‐bodied exotic omnivores, with potential implications for fish–zooplankton interactions still largely unknown. Here we studied diel variation in the vertical and horizontal distribution of the crustacean plankton in Lake Vela, a shallow polymictic and eutrophic lake. Diel sampling was carried out on three consecutive days along a horizontal transect, including an open‐water station and a macrophyte (Nymphaea alba) bed. Since transparency is a key determinant of the predation risk posed by fish, the zooplankton sampling campaigns were conducted in both the turbid (autumn) and clear water (spring) phases. 2. In the turbid phase, most taxa were homogeneously distributed along the vertical and horizontal axes in the three consecutive days. The only exception was for copepod nauplii, which showed vertical heterogeneity, possibly as a response to invertebrate predators. 3. In the clear water phase, most zooplankton taxa displayed habitat selection. Vertically, the general response consisted of a daily vertical migration (DVM), despite the limited depth (1.6 m). Horizontally, zooplankters showed an overall preference for the pelagic zone, independent of the time of the day. Such evidence is contrary to the postulated role of macrophytes as an anti‐predator refuge for the zooplankton. 4. These vertical (DVM) and horizontal (macrophyte‐avoidance) patterns were particularly conspicuous for large Daphnia, suggesting that predation risk from size‐selective predators (fish) was the main factor behind the spatial heterogeneity of zooplankton in the spring. Thus, the difference in the zooplankton spatial distribution pattern and habitat selection among seasons (turbid and clear water phases) seems to be mediated the predation risk from fish, which is directly related to water transparency. 5. The zooplankton in Lake Vela have anti‐predator behaviour that minimises predation from fish. We hypothesise that, due to the distinct fish community of shallow Mediterranean lakes, aquatic macrophytes may not provide adequate refuge to zooplankters, as seen in northern temperate lakes.     

Grazing rates of crustacean zooplankton communities on intact phytoplankton communities in Canadian Subarctic lakes and ponds

Quantitative genetic studies in natural populations are of growing interest to speciation research since divergence is often believed to arise through micro-evolutionary change, caused by natural selection on functional morphological traits. The species flock of cichlid fishes in Africa’s oldest lake, Lake Tanganyika, offers a rare opportunity to study this process. Using the cichlid species Tropheus moorii, we assessed the potential for microevolution in a set of morphological traits by estimating their quantitative genetic basis of variation. Two approaches were employed: (1) estimation of trait heritabilities (h ²) in situ from a sample of wild caught fish, and (2) estimation of h ² from first generation offspring produced in a semi-natural breeding experiment. In both cases, microsatellite data were used to infer pedigree structure among the sampled individuals and estimates of h ² were made using an animal model approach. Although power was limited by the pedigree structures estimated (particularly in the wild caught sample), we nonetheless demonstrate the presence of significant additive genetic variance for aspects of morphology that, in the cichlid species Tropheus moorii, are expected to be functionally and ecologically important, and therefore likely targets of natural selection. We hypothesize that traits showing significant additive genetic variance, such as the mouth position have most likely played a key role in the adaptive evolution of the cichlid fish Tropheus moorii.     

Interactions between phytoplankton and zooplankton in a fertilized lake

The limnology of an oligotrophic lake, Langvatn, situated in Trøndelag county in Central Norway, has been studied for five years (1974–1978). In two years, 1975 and 1976, the lake was fertilized with a general fertilizer to change feeding conditions for the zooplankton. Mean phytoplankton biomass in the epilimnion and primary production for the years (1974–1978) were 417, 618, 1370, 607 and 779 mg m⁻³ and 10.6, 22.2, 49.0, 26.8 and 17.7 g C m⁻² yr⁻¹, respectively. Cladocerans were the dominant herbivore group in 1974 and 1975 and Rotifera in the next three years. The main difference in the interaction between phytoplankton and zooplankton occurred when cladoceran dominance gave way to rotifer dominance. Heavy phytoplankton grazing by cladocerans in 1974 and 1975 stabilized the biomass and maintained it at a low level, which also resulted in a relatively low primary production. The rotifer-dominated community during the years 1976–1978 did not possess the ability to maintain a stable level of algal biomass. Primary production was also relatively high during these years.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

The light-climate of Loch Leven, a shallow Scottish lake, in relation to primary production by phytoplankton

Seasonal changes in incident irradiance and underwater light penetration at Loch Leven from 1968 to 1971 are discussed in relation to the photosynthetic behaviour and crop density of phytoplankton. Light extinction was highest in the blue and lowest in the orange spectral regions, a pattern typical of other turbid waters. Euphotic depth varied between 1·2 and 7·4 m and was on average c. three times the Secchi disc transparency. Underwater light extinction depended chiefly on phytoplankton crop density (estimated as chlorophyll a). Despite the shallowness and wind-exposed situation of the loch there was no evidence of appreciable light extinction due to sediment disturbance. Possible causes of variability in the relationship between the minimum vertical extinction coefficient (k _min) and the concentration of chlorophyll a are discussed. The value of k_s, the increment in k_min per unit increment in algal concentration, was estimated from field data as 0·0086 In units per mg chl a/m² and from laboratory spectroradiometer data as 0·0079 In units per mg chl a/m². These k_s values imply theoretical upper limits for the amount of chlorophyll a in the euphotic zone (Σn _max) of 430 and 468 mg chl a/m², respectively. Observed euphotic chlorophyll a contents (Σn) were sometimes close to these upper limits. Typical photosynthesis/depth profiles are described. Profile area is shown to be related to the logarithm of the ratio between surface-penetrating irradiance (I_o') and the irradiance (I_k) defining the onset of light-saturation of photosynthesis. Standardized profiles, plotted on a common scale of ‘optical depth’, are used to illustrate the relatively minor influence of variations in I_o' and I_k on hourly rates of photosynthesis per unit area. The saturation parameter (I_k) generally increased as photosynthetic capacity (P_max) increased; the temperature-dependence of I_k is explained by the temperature-dependence of the enzyme-controlled (dark) reactions of photosynthesis, which control P_max. A spring peak in the ratio between surface penetrating irradiance (I_o') and I_k is interpreted as a result of a lag in the seasonal increase in water temperature with increase in surface irradiance. The gradient (K') of the linear light-limited region of the photosynthesis-irradiance curve showed little variation and had an average value of 0·31 mg O₂/mg chl a.h per 1 W/m² (PAR). Interactions between mixed depth, underwater light extinction and phytoplankton productivity are discussed; comparisons are made with other shallow, optically deep lakes.     

Diel patterns of zooplankton grazing in a shallow lake

A diel survey of in situ species-specific zooplankton clearancerates (with radioactively labelled Chlorella cells) was conductedin the shallow lake of Cr?teil characterized by small-sizedplanktonic forms (algae<10µm and zooplankton <1.3mm). Experiments were performed every 4h at two depths (1 and4m). Power functions relating individual filtering rates tobody length were established for the three most abundant cladoceransand for calanoids (nauplii being included in this feeding group),for each depth and time. No filtering penodicity was observedin Ceriodaphnia spp., adults and copepodites of Eudiaptomusgracilis and Eurytemora velox, and nauplii. On the contrary,clear nocturnal filtering peaks were obtained for Daphnia spp.and for Diaphanosoma brachyurum, these being more pronouncedfor the larger individuals at 1 m deep. The observed diel periodicitycannot be explained by variations in physico-chemical parametersor food concentration alone. The ecological significance ofthis phenomenon in the polymictic lake of Cr?teil is discussedin the light of previously published data and the hypothesesaccounting for it.     

Long-term changes in crustacean zooplankton and water quality in a shallow, eutrophic Chinese lake densely stocked with fish

From surveys made in 1962–1963, 1973–1974, 1979–1996 at two Stations in Lake Donghu, a shallow eutrophic water body near Wuhan, P. R. China, the authors, derive long-term changes in species composition, standing crop and body-size of planktonic crustaceans. The species number decreased from the 1960s to the 1990s. The cladocerans dropped from 46 (1960s) to 26 (1980s) to 13 (1990s); the copepods decreased from 14 (1960s) to 10 (1980s) to 7 (1990s). From the mid-1980s on, the dominant crustaceans also changed: Daphnia hyalina and D. carinata ssp. were replaced by Moina micrura and Diaphanosoma brachyurum at Stations 1 and 2, respectively; Cyclops vicinus replaced Mesocyclops leuckarti.Densities and biomass of Cladocera decreased markedly after 1987. Annual average densities and biomass of cladocerans were statistically differences between 1962–1986 and 1987–1996 (P > 0.01). Annual average densities of Daphnia (Station 1 + Station 2) were negatively correlated with fish yield .Since the 1980s, annual average body length of Cladocera and Calanoida decreased, while annual average body length of Cyclopoida increased. In the same years, average body length of copepods was lower during May–October than during January–April and November–December.A 12-yr data analysis showed annual average concentration of chlorophyll^-a (Chl^-a) to be negatively correlated with annual average density of Daphnia, whilst lake transparency was positively correlated with annual average densities of Daphnia. The results imply that, since Daphnia feeds efficiently on phytoplankton, it could decrease concentration of Chl^-a, and enhance water transparency.     

Sedimentation of phytoplankton populations dominated by Microcystis in a shallow lake

The sedimentary flux of phytoplankton was measured using sedimenttraps in a shallow hypertrophic lake (Lake Kasumigaura), whereMicrocystis bloomed, from June to November 1983 The sedimenttraps were set at 0.5, 1.5 and 3.0 m depth in Takahamairi Bay(3.5 m depth). Microcystis spp. (including M.aerugmosa and M.viridis)in the traps were rare until early August, but increased thereafter.Sinking rates of Microcystis were 0.0045, 0.020 and 0.24 m day^–1in June–August, September and October respectively, whichwere far lower than those of Melosira (0.2–1.7 m day^–1)and Syncdra (0.2–1.0 m day^–1). The total sedimentaryfluxes of POC and that of algal carbon during the study periodwere 283.2 and 96.7 gC m^–2 which were 59.5% and 20.3%of the gross primary production (475.8 gC m^–2) respectively.The sedimentary flux of living algae measured by algal countswas large in June but small in August and September. On theother hand, the flux of detritus obtained by subtracting totalalgal carbon from POC was small in June and July but large inAugust and September. Therefore diatoms, which appeared mostlyin June, tended to sink as live algae, while Microcystis sankas detritus after being decomposed or consumed in the waterIt was concluded from the results of carbon budget calculationsand the respiration rate of the 1- to 20-µm fraction thatthe activity of decomposers or consumers increased greatly inthe short period at the end of the bloom of Microcystis.     

Restoring lakes by using artificial plant beds: habitat selection of zooplankton in a clear and a turbid shallow lake

1. Return of large‐bodied zooplankton populations is of key importance for creating a shift from a turbid to a clear‐water state in shallow lakes after a nutrient loading reduction. In temperate lakes, recovery is promoted by submerged macrophytes which function as a daytime refuge for large zooplankton. However, recovery of macrophytes is often delayed and use of artificial plant beds (APB) has been suggested as a tool to enhance zooplankton refuges, thereby reinforcing the shift to a clear‐water state and, eventually, colonisation of natural plants. 2. To further evaluate the potential of APB in lake restoration, we followed the day–night habitat choices of zooplankton throughout summer in a clear and a turbid lake. Observations were made in the pelagic and littoral zones and in APB in the littoral representing three different plant densities (coverage 0%, 40% and 80%). 3. In the clear lake, the zooplankton (primarily Daphnia) were mainly found in the pelagic area in spring, but from mid‐May they were particularly abundant in the APB and almost exclusively so in mid‐June and July, where they appeared in extremely high densities during day (up to 2600 ind. L⁻¹). During night Daphnia densities were overall more equally distributed between the five habitats. Ceriodaphnia was proportionally more abundant in the APB during most of the season. Cyclopoids were more abundant in the high APB during day but were equally distributed between the five habitats during night. 4. In the turbid lake, however, no clear aggregation was observed in the APB for either of the pelagic genera (Daphnia and Bosmina). This may reflect a higher refuge effect in the open water due to the higher turbidity, reduced ability to orient to plant beds and a significantly higher fish density (mainly of roach, Rutilus rutilus, and perch, Perca fluviatilis) in the plant beds than in the clear lake. Chydorus was found in much higher proportions among the plants, while cyclopoids, particularly the pelagic Cyclops vicinus, dominated in the pelagic during day and in the pelagic and high density plants during night. 5. Our results suggest that water clarity is decisive for the habitat choice of large‐bodied zooplankton and that introduction of APB as a restoration measure to enhance zooplankton survival is only a useful tool when water clarity increases following loading reduction. Our results indicate that dense APB will be the most efficient.     

The microbial food web structure of a hypertrophic warm-temperate shallow lake, as affected by contrasting zooplankton assemblages

The composition of zooplankton is known to affect the structure of the microbial trophic web. The zooplankton of the hypertrophic Laguna Chascomús (Argentina) is generally dominated by rotifers and cyclopoids copepods. An unusual dominance by small-cladocerans was observed after a massive winter fish kill in 2007. We hypothesized that small-cladocerans would increase the grazing pressure on heterotrophic flagellates (HF), reducing the degree of coupling between HF and picoplankton. The aim of this study was to investigate the microbial food web structure under two contrasting zooplankton assemblages. The lake was sampled every other week between 2007 and 2009. The abundances of heterotrophic bacteria (HB) and picocyanobacteria (Pcy) laid among the highest values reported for aquatic systems (>10⁸ and 10⁷ cells ml^?1, respectively). Pcy averaged 53% of total picoplanktonic biomass. When small-cladocerans dominated zooplankton HF reached the higher abundance (>10⁵ cells ml^?1) and picoplankton showed the opposite pattern, while the proportion of grazing resistant morphologies (i.e. microaggregates of Pcy) was higher. In contrast, when rotifers dominated, HF abundance decreased and picoplankton increased. Our data suggest that the degree of HF–HB coupling was affected by changes in zooplankton dominance. In contrast to our initial hypothesis, the present results suggest that large numbers of rotifers (>5,000 ind. l^?1) are more efficient than small-cladocerans at controlling HF populations.     

Grazing of bacteria and phytoplankton by heterotrophic nanoflagellates in a Baltic Sea sample

Grazing by heterotrophic nanoflagellates on bacteria and phytoplankton was studied in a laboratory experiment, using a natural pelagic community originating from the Tvärminne sea area off the southern coast of Finland. Water was prescreened to remove larger grazers. Four experimental treatments were used: light and dark, with and without added nutrients. The growth of the large heterotrophic flagellates was stimulated by increased production of < 3 m phytoplankton. Clearance rates for heterotrophic nanoflagellates were estimated and were found to be within the range of previously reported values.     

Top-down effects of crustacean zooplankton on pelagic microorganisms in a mesotrophic lake

A series of single-factor in situ experiments was conductedin a mesotrophic lake in Brandenburg, North Germany, to studythe predatory impact of Eudiaptomus graciloides (adults, copepodites,nauplii), cyclopoid copepods (adult Diacyclops bicuspidatus,Thermocyclops oithonoides) and daphnids (adult Daphnia hyalina,Daphnia cucullata) on the microbial community (bacteria, autotrophicpicoplankton, flagellates, ciliates). All zooplankton speciestested reduced the ciliate community significantly and ingestionrates were always higher for ciliates in the 20–55 µmsize category as compared to smaller ciliates (10–20 µm).Adult E.graciloides, which exhibited the highest predatory impacton ciliates, differed from cyclopoids and daphnids by theirability to decimate ciliates to very low abundances. Ingestionrates of ciliates by the crustacean zooplankton followed thesequence E.graciloides > daphnids = cyclopoids = copepodites.While top-down control was evident for ciliates, top-down effectsdown to the autotrophic picoplankton and flagellates were mostlyrestricted to Daphnia-dominated treatments. Top-down effectswere never strong enough to produce negative bacterial growthrates. For all zooplankton tested, clearance rates for ciliatesexceeded those for phytoplankton. Besides the potential of thecrustacean zooplankton to influence the structure of ciliatecommunities, ciliates may contribute to the energy demands ofcopepods and daphnids, especially when phytoplankton resourcesare limited.     

Spatial and temporal distribution of fish and zooplankton in a shallow lake

Summary 1. We performed both a large- and a small-scale echo sounding study on the spatial and temporal distribution of fish (mainly roach, Rutilus rutilus and perch, Perca fluviatlis ), as well as a small-scale study of zooplankton distribution in the small, shallow and eutrophic Lake Hanebjerg in Denmark. In the small-scale study, sampling was conducted in open water as well as in the edge zone immediately outside two different types of vegetation.
2. Fish daytime abundances differed between the northern and the southern parts of the lake and, on a small scale, small fish aggregated in the edge zones during day, preferably outside dense emergent vegetation. Copepods avoided emergent vegetation, while cladocerans showed no habitat preference. Both small fish and cladoceran numbers were found to be higher during night than day.
3. The relative abundance (number per sample) of cladocerans in the edge zone immediately outside vegetation was negatively correlated with the relative abundance of fish in that zone. There was no correlation between cladoceran and fish abundance in open water, or between the relative abundance of copepods and fish.
4. The presence of pelagic piscivores in combination with avoidance behaviour of both fish and zooplankton is a likely explanation for the observed distribution of small fish and cladocerans in Lake Hanebjerg. Both small- and large-scale distribution patterns may be dependent on the type and distribution of complex structure in the lake. Even in a small lake, large-scale patterns may affect the interpretation of small-scale data.     

Composition,density and feeding of crustacean zooplankton community in a shallow,temperate lake (Lake Balaton,Hungary)

Composition density and filtering rates of crustacean zooplankton were studied in the open water and among the macrophytes of the oligo-mesotrophic part of Lake Balaton from 1981 to 1983. From the individual filtering rates of the different populations and the densities community grazing rates were derived.Copepoda made up 79–90% of crustacean plankton community in the open lake and 95–97% of it in littoral zone. Among them the nauplii dominated. At the end of summer 1982, when Anabaenopsis was in bloom, the filter-feeding species (Eudiaptomus, Daphnia) practically disappeared, being replaced by cyclopoids. Daphnia had the highest filtering rates followed by those of Eudiaptomus and Diaphanosoma. Among copepods, the filtering rates in decreasing order were: ovigerous > all adults > copepodites > nauplii. > . The filtering rates of the different species varied both seasonally as well as from year to year. In 1983, when the concentration of organic seston decreased, filtering rate increased compared with those in the earlier years. During the water bloom in 1982, the rates decreased by 70% on the average.The community grazing rate was very low (3% per day) in the open lake and among macrophytes, both in 1981 and 1982; also the share of crustacean zooplankton in grazing was very low. In 1983, together with the improving of water quality, the community grazing rate increased 4-fold. In 1981 and 1983 the rates were influenced by water temperature but in 1982 by seston concentration.     

An analysis of long-term winter data on phytoplankton and zooplankton in Neusiedler See,a shallow temperate lake,Austria

In the last 40 years, the shallow steppe lake, Neusiedler See, was ice covered between 0 and 97 days. The North Atlantic Oscillation (NAO) as well as the Mediterranean Oscillation affected the lake and its conditions during winter. Both climate indices correlated negatively with the duration of ice cover and the timing of ice-out. Average winter phytoplankton biomass increased from less than 0.2 (0.05–0.84) mg FM l^?1 in the late 1960s/beginning of 1970s to 3.1 (1.72–5.61) mg FM l^?1 in the years 2001–2004. The increase in annual winter biomass of phytoplankton was associated with a significant shift in the composition of the algal assemblage. In the winter 1997/1998, diatoms contributed between 40 and 80% to the phytoplankton biomass while in 2006/2007 cyanoprokaryotes contributed 46%. Mean chlorophyll-a concentrations during winter were significantly correlated with those of total phosphorus (P_tot). Together with cold-water species (rotifer Rhinoglena fertöensis), perennial, eurythermal ones (copepod Arctodiaptomus spinosus) contributed to the zooplankton community. High zooplankton numbers were encountered when rotifers, particularly when densities of Rhinoglena fertöensis were high (r ² = 0.928). Zooplankton abundance and biomass varied from year to year but correlated positively with Chl-a (biomass ? r ² = 0.69; numbers ? r ² = 0.536). Winter zooplankton populations were primarily influenced by winter conditions, but in early winter also by survival of autumn populations, i.e., the more adults of Arctodiaptomus spinosus survived into winter, the higher was the zooplankton biomass in early winter. Phyto- and zooplankton dynamics in shallow lakes of the temperate region seem to critically depend on the biomass in autumn and on winter conditions, specifically on ice conditions and thus are related to climate signals such as the NAO.     

Responses to food web manipulation in a shallow waterfowl lake

The effects of fish stock reduction have been studies in 3 Dutch lakes (Lake Zwemlust, Lake Bleiswijkse Zoom and Lake Noorddiep) and 1 Danish lake (Lake Væng) during 4–5 years. A general response id described. The fish stock reduction led in general to a low fish stock, low chlorophyii-a, high transparency and high abunuance of macrophytes. Large Daphnia became abundant, but their density decreased, due to food limitation and predation by fish. The total nitrogen concentration became low due to N-uptake by macrophytes and enhanced denitrification. In Lake Bleiswijkse Zoom the water transparency deteriorated and the clear water state was not stable. The fish stock increased and the production of young fish in summer was high. lear water occurred only in spring. Large daphnids were absent in summer and the macrophytes decreased.In Lake Zwemlust, Lake Væng and Lake Noorddiep the water remained clear during the first five years. In summer of the sixth year (1992) transparency decreased in Lake Zwemlust (with high P-concentration of 1.0 mg P l^-1). Also in Lake Væng (with a low nutrient concentration of 0.15 mg P.^-1) a short term turbid stage (1.5 month) occurred in summer 1992 after a sudden collapse of the macrophytes. Deterioration of the water quality seems to start in summer and seems related to a collapse in macrophytes. At a low planktivorous fishstock (e.g. Lake Væng)thhe duration of the turbid state is shorter. than in presence of a high planktivorous fish biomass (e.g. Lake Zwemlust, and later years of Lake Bleiswijkse Zoom).     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.     

Factors controlling production of phytoplankton and bacteria under ice in a humic, boreal lake

The factors controlling pelagic primary and bacterial productionof a humic, boreal lake in winter were investigated, combininglaboratory and field experiments where some of the predictedconsequences of the climate change, i.e.the increased load ofphosphate-phosphorus and dissolved organic matter (DOM), weresimulated. In situ incubations were performed in eight acrylictubes lowered underneath the ice cover for 1–4 months.In the lake, production of phytoplankton (0.03–0.33 µgC l^–1 day^–1) was lower than that of bacteria (0.2–2.3µg C l^–1 day^–1) from the end of January tomid-April. Later in spring, the light conditions improved dueto the disappearance of snow and finally ice itself, and primaryproduction was revived. The importance of light as the factorcontrolling primary production in winter was confirmed in laboratoryexperiments where additions of phosphorus and DOM did not enhancethe primary production. These same enrichments resulted in highergrowth rates and production of bacteria. However, bacterioplanktonwas simultaneously controlled by heterotrophic flagellates,as in the laboratory experiments the exclusion of flagellatesalways resulted in higher yields of bacteria of increased cellsize. There was also some evidence that the quality of substratesis of importance for microbial activities. The importance oftemperature was highlighted with the significant correlation(r² = 0.59) between bacterial production and temperature withinthe range 0.6–2.1C. Thus, on the boreal zone the possiblewarming of climate may lead to enhanced activities in winter,but under unchanged light conditions no effects on primary productionwill be expected.