Feeding in Euchlanis dilatata lucksiana Hauer on filamentous cyanobacteria and a prochlorophyte

Ingestion and assimilation rates of Euchlanis dilatata lucksiana Hauer, isolated from Lake Loosdrecht (The Netherlands) and cultured on lake water (seston < 33 µm), were measured in the laboratory using the ¹⁴C-tracer technique. The five taxa used as tracer foods, together with 6–7 mg C l^–1 of lake seston in each case, included four species of filamentous cyanobacteria (Oscillatoria redekei, O. limnetica, Aphanizomenon flos-aquae, Anabaena PCC 7120) and a prochlorophyte (Prochlorothrix hollandica). Except Anabaena, they are all commonly encountered in eutrophic Loosdrecht lakes, including Lake Loosdrecht, and their dimensions ranged between 150 and 250 µm in length and 2 and 3.5 µm in width. The small and large Euchlanis used as experimental animals had mean lengths of 217–223 µm and 276–305 µm, respectively. Euchlanis fed on all the taxa offered as food. Clearance rates ranged from 51 to 99 µl ind^–1 d^–1 for the large animals and from 22 to 41 µl ind^–1 d^–1 for the small animals. The highest ingestion rate observed, 1.7 µg ind^–1 d^–1, was for the large animals feeding on Aphanizomenon. The daily ration for both size classes far exceeded 100% of body weight, reaching up to 690% for the small animals feeding on Aphanizomenon. The small animals generally appeared to assimilate the ingested food more efficiently (assimilation efficiencies: 37–100%) than the large animals (34–77%). Compared with an earlier study in which only lake seston (<33 µm) was used as food, the specific clearance rates of Euchlanis on the cyanobacteria and Prochlorothrix were generally somewhat lower.     

Modelling phosphorus fluxes in the hypertrophic Loosdrecht Lakes

A dynamic, deterministic model is presented to simulate the phosphorus cycle and plankton growth in the shallow, hypertrophic Loosdrecht Lakes (The Netherlands) before and after restoration measures. Besides inorganic phosphorus (SRP) in both the surface water and the interstitial water, the model comprises three algal groups, zooplankton, fish, detritus, zoobenthos and upper sediment (all modelled both in carbon and in phosphorus). Within the model system, the phosphorus cycle is completely closed. Carbon and phosphorus are described independently, so that the dynamics of the P/C ratios can be modelled. Sediment processes are described in a simplified form.Simulated values are largely within the range of observed ones. The detrital fraction of the seston (=phytoplankton+detritus) varies from 50–60% in summer to about 90% in winter. SRP in the surface water is very low during most of the year. Sensitivity for external phosphorus input is larger for algal and detrital P than for algal and detrital C and chlorophyll-a. So the P/C ratio of the seston decreases following restoration measures, as is observed in the lakes, while the much higher P/C ratios of zooplankton and fish remain constant. Phosphorus mobilisation from the sediment decreases with decreasing external input. Adaptation of the model system to the reduced loading takes place within about two years.Sources of uncertainty in the model include the limited knowledge on selective grazing as well as on mortality and mineralisation processes.     

Dynamics of phytoplankton detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

A study was made of the mortality and aerobic decomposition of light- and phosphorus-limited cultures of Oscillatoria limnetica, a dominant phytoplankton species in shallow, eutrophic Lake Loosdrecht (The Netherlands). When placed in the dark at 20 °C, most cells died and lysed within twelve days. The labile organic matter was completely decomposed within three weeks. Absorbance spectra indicated that blue green algae may contributed significantly to the refractory dissolved substances in the lake. Refractory particulate matter constituted from 7 to 24% of the biomass of O. limnetica, depending on the growth rate before incubation in the dark. The decomposition rate of this fraction was 0.005 d^–1. On a basis of a steady-state model of the dynamics of phytoplankton detritus, the areal organic dry weight concentration of the detritus in the lake is ca. 60 g m^–2. This means the quantities of detritus in the seston and epipelon are about equal.     

The dynamics and role of limnetic zooplankton in Loosdrecht lakes (The Netherlands)

The paper summarizes the results of a ten-year (1981–1991) zooplankton research on the Lake Loosdrecht, a highly eutrophic lake. The main cause of the lake's eutrophication and deteriorating water quality was supply up to mid 1984 of water from the River Vecht. This supply was replaced by dephosphorized water from the Amsterdam-Rhine Canal in 1984. The effects of this and other restoration measures on the lake's ecosystem were studied. Despite a reduction in the external P-load from ca. 1.0 g P m^–2 y^–1 to ca. 0.35 g m^–2 y^–1 now, the filamentous prokaryotes, including cyanobacteria and Prochlorothrix, continue to dominate the phytoplankton.Among the crustacean plankton Bosmina spp, Chydorus sp. and three species of cyclopoid copepods and their nauplii are quite common. Though there was no major change in the composition of abundant species, Daphnia cucullata, which is the only daphnid in these lakes, became virtually extinct since 1989. Among about 20 genera and 40 species of rotifers the important ones are: Anuraeopsis fissa, Keratella cochlearis, Filinia longiseta and Polyarthra. The rotifers usually peak in mid-summer following the crustacean peak in spring. The mean annual densities of crustaceans decreased during 1988–1991. Whereas seston (< 150 µm) mean mass in the lake increased since 1983 by 20–60%, zooplankton (> 150 µm) mass decreased by 15–35%.The grazing by crustacean community, which was attributable mainly to Bosmina, had mean rates between 10 and 25% d^–1. Between 42 and 47% of the food ingested was assimilated. In spring and early summer when both rotifers and crustaceans have their maximal densities the clearance rates of the rotifers were much higher. Based on C/P ratios, the zooplankton (> 150 µm) mass contained 2.5 times more phosphorus than seston (< 150 µm) mass so that the zooplankton comprised 12.5 % of the total-P in total particulate matter in the open water, compared with only 4.5% of the total particulate C. The mean excretion rates of P by zooplankton varied narrowly between 1.5 and 1.8 µg P 1^– d^–1, which equalled between 14 and 28% d^–1 of the P needed for phytoplankton production.The lack of response to restoration measures cannot be ascribed to one single factor. Apparently, the external P-loading is still not low enough and internal P-loading, though low, may be still high enough to sustain high seston levels. Intensive predation by bream is perhaps more important than food quality (high concentrations of filamentous cyanobacteria) in depressing the development of large-bodied zooplankton grazers, e.g. Daphnia. This may also contribute to resistance of the lake's ecosystem to respond to rehabilitation measures.     

A laboratory study of feeding and assimilation in Euchlanis dilatata lucksiana

Feeding of Euchlanis dilatata lucksiana, a brachionid rotifer isolated from the Loosdrecht Lakes (The Netherlands), was examined in the laboratory using ¹⁴C-labelled food. The gut-filling time at a food concentration of 9.6 µg C ml^–1 was about 15 minutes. Animals which were fed on 3 size fractions of the lake seston (< 7 µm, 7–15 µm, and 15–33 µm) showed a clear preference (Jacobs' selectivity index) for the largest size fraction. This fraction was composed predominantly of filamentous cyanobacteria. For animals weighing 0.37–0.49 µm C ind.^–1 the daily ration (daily food consumption per unit body weight × 100) ranged from 50 to 100% at food levels of 2 mg C l^–1 and below, but increased to 150–250% at food concentrations of 5 mg C l^–1 and above. The assimilation efficiency was 100% up to 5 mg C l^–1 of food, but decreased to about 80% at higher food levels.     

Coupling of phytoplankton and detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

An oscillating steady state is described of phytoplankton, dominated by Prochlorothrix hollandica and Oscillatoria limnetica, and sestonic detritus in shallow, eutrophic Lake Loosdrecht (The Netherlands). A steady-state model for the coupling of the phytoplankton and detritus is discussed in relation to field and experimental data on phytoplankton growth and decomposition. According to model predictions, the phytoplankton to detritus ratio decreases hyperbolically at increasing phytoplankton growth rate and is independent of a lake's trophic state. The seston in L. Loosdrecht contains more detritus than phytoplankton as will apply to many other lakes. The model provides a basis for estimating the loss rate of the detritus, including decomposition, sedimentation and hydraulic loss. In a shallow lake like L. Loosdrecht detritus will continue to influence the water quality for years.     

Chlorophyll content of seston in a regulated Rocky Mountain river,Idaho, USA

The chlorophyll content of seston at four sites in a regulated mid-order Rocky Mountain river, Henry's Fork of the Snake River, Idaho, USA, was examined. Spectrometry was used to determine the amount of chlorophyll a, b, c and in seston and potential organic matter sources including macrophytes, algae, plankton, and terrestial plant litter. The amount of chlorophyll in seston varied among size fractions, sites, and seasons. Coarse seston (6–1 mm) contained the most chlorophyll a (2.3 mg g^–1 ashfree dry mass [AFDM]) followed by ultra fine seston (53–0.3 µm; 1.9 mg g^–1 AFDM), very coarse seston (> 6 mm, 1.8 mg g^–1 AFDM), very fine seston (250–53 µm; 1.3 mg g^–1 AFDM), and fine seston (1–0.25 mm; 0.7 mg g^–1 AFDM). Chlorophyll content of coarse seston was similar at all sites reflecting a common source, aquatic macrophyte debris. Chlorophyll content of coarse and fine seston were highest in fall reflecting the importance of phenology of aquatic plants on sestonic pigment levels. Very fine seston from below a reservoir contained more chlorophyll than seston from downstream and tributary sites suggesting a reservoir source. Terrestial plant litter was chlorophyll depleted compared to autochthonous materials and seston. Most seston was autochthonously-derived from a variety of macrophyte, algal, and planktonic sources. The chlorophyll content of Henry's Fork seston was higher than that reported for similar rivers. Island Park Dam moderates river temperature and flow, enhances autotrophy, and accounts for the consistently high sestonic chlorophyll levels.     

A laboratory study of phosphorus and nitrogen excretion of Euchlanis dilatata lucksiana

Phosphorus (PO₄-P) and nitrogen (NH₄-N) excretion rates of Euchlanis dilatata lucksiana, a rotifer, isolated from Lake Loosdrecht (The Netherlands) and cultured in the lake water at 18–19 °C, were measured in the laboratory.In a series of experiments, the effects of experiment duration on the P and N excretion rates were examined. The rates measured in the first half-hour were about 2 times higher for P and 2–4 times for N than the rates in the subsequent three successive hours which were quite comparable.Eight experiments were carried out in triplicate, 4 each for P and N excretion measurements, using animals of two size ranges: 60–125 µm and > 125 µm. The specific excretion rates varied from 0.06 to 0.18 µg P.mg^–1 DW.h^–1 and 0.21 to 0.76 µg N.mg^–1 DW.h^–1. Generally an inverse relationship was observed between the specific excretion rates and the mean individual weight. The excretion rates of Euchlanis measured by us are lower than those reported for several other rotifer species, most of which are much smaller than Euchlanis.Extrapolating the excretion rates of Euchlanis to the other rotifer species in Lake Loosdrecht, and accounting for their density, size and temperature, rotifer excretion appears to be a significant, potential nutrient (N,P) source for phytoplankton growth in the lake. The excretion rates for the rotifers appear to be about two thirds of the total zooplankton excretion, even though the computed rotifer mean biomass is about one-third of the total zooplankton biomass.     

Phosphorus in the sediment of the Loosdrecht lakes and its implications for lake restoration perspectives

In 1984 the external phosphorus load of the shallow eutrophic Loosdrecht lakes was reduced from 3.3 to 1.0 mg m^–2 d^–1. The effect of phosphorus release from the sediment on lake restoration was investigated. Diffusive release under aerobic conditions (20 °C) decreased from 1 mg m^–2 d^–1 in 1984 to 0.3 mg m^–2 d^–1 in 1990. The generation of inorganic phosphorus due to mineralization during summer equals 3 mg m^–2 d^–1, which is much higher than the measured rate of diffusive release. Despite that, the phosphorus release is hardly stimulated by anaerobic conditions, which indicates that only a small amount of phosphorus is adsorbed by ferric iron in the top sediment layer. This apparent discrepancy is probably caused by the uptake of inorganic phosphorus uptake during resuspension and the loss of inorganic phosphorus with downward seepage.The estimated removal of phosphorus due to downward seepage of 0.8 mg m^–2 d^–1 agrees well with the average phosphorus retention in the lake. This indicates that sediment burial and diagenesis are unimportant mechanisms for withdrawing phosphorus from the nutrient cycle.Between 1982 and 1991 the total phosphorus content of the upper 2 cm of the sediment decreased from 0.94 to 0.60 g kg^–1 DW. At present, about 20% of total phosphorus in this layer is potentially bioavailable, but largely incorporated in easily degradable organic matter. This pool is much smaller in deeper layers. Based on the estimated and measured rates and pool sizes, the annual average phosphorus cycle in the lakes was modelled to evaluate the effects of various restoration measures. The main predictions of the model are: 1) further reduction of the external load may cause a gradual decrease of the total phosphorus concentration in the lake water; 2) dredging and iron addition, without reduction of the external load, may give a rapid improvement followed by a slow return to the present situation; and 3) reduction of the external load, combined with a cut off of downward seepage will not improve the water quality.     

Effects of diet and heavy metals on growth rate and fertility in the deposit-feeding snail Potamopyrgus jenkinsi (Smith) (Gastropoda: Hydrobiidae)

Evidence for the influence of food type and heavy metals on shell growth and fertility is presented for a freshwater population of the snail P. jenkinsi. When fed an excess of lettuce or lamb heart (protein source), growth rates were higher for lettuce. Highest growth rates occurred at a diet of lettuce plus lamb heart. Fertility was favoured by a diet of lamb heart. When fed an excess of lettuce, the EC₅₀ growth values were 16 µg Cd l^–1, 13 µg Cu l^–1, and 103 µg Zn l^–1 in lake water; snail fertility was inhibited at 25 µg Cd l^–1 and 30 µg Cu l^–1. A diet of lake detritus spiked with Cd or Cu resulted in a decrease of approximately 50% in growth rates, when compared with growth on non-spiked detritus. Spiked detritus lost metals into lake water. Food type positively interacted with metal stress, both for growth rate and fertility. The assessment of inhibitory effects of detritus contaminated either in the field or, notably, by spiking, and serving as food source for deposit feeders is hampered by sampling problems in the field and by redistribution processes of pollutants between particles and water in laboratory-scale experiments.     

Studying the phosphorus release from the Loosdrecht Lakes sediments,using a continuous flow system

Phosphorus release from the Loosdrecht Lakes sediments was studied, using a continuous flow reactor. The summer release maxima were 4 mg P.m^–2.d^–1 in 1984 and 1.4 mg P.m^–2.d^–1 in 1985. Temperature and downward seepage controlled release rates to a great extent, the pH of the overlying water being only of minor importance. From these results it could be concluded that release processes might be driven by mineralization of particulate organic phosphorus in the sediment. Pore water studies in the sediments of the release reactor confirmed this hypothesis. From the profiles phosphorus dissolution rates were calculated.     

Feeding habits and food resources of filter-feeding Trichoptera in a regulated mountain stream

A year-round study was conducted to examine feeding habits and food resources of the filter-feeding Trichoptera Arctopsyche grandis and Brachycentrus occidentalis along a regulated mountain stream gradient. There was a well defined longitudinal species replacement with A. grandis reaching maximum densities 2.3 kilometers below the impoundment, and concomitant with its decline downstream was an increase in B. occidentalis. At all sampling sites the < 75 µm organic seston fraction usually consisted primarily of diatoms (>70%, by areal estimate on microscope slides), whereas the 75–250 µm and > 250 µm seston fractions were predominantly composed of detritus (> 80 %). B. occidentalis larvae consumed primarily detritus and diatoms (> 70 % of the diet), while A. grandis ingested a variety of materials with animals, detritus and/or filamentous algae often constituting > 80% of the diet. Animal material was over-represented in the diets of both species when compared with amounts in the seston. Feeding habits provided partial explanations for the distinct longitudinal distribution patterns of filter-feeding Trichoptera observed in the regulated river.     

In situ phosphorus release experiments in the Warnow River (Mecklenburg,northern Germany)

Results are presented of in situ benthic phosphorus release experiments in an undercut bank of an impounded river. Due to high sedimentation of phytoplankton biomass high oxygen consumption rates between 259.4 and 947.0 mg O₂ m^–2 d^–1 developed, leading to almost anaerobic conditions and phosphorus releases between 175.2 and 236.3 mgP m^–2 d^–1 over a period of 18 days.In a second series of experiments the water column overlying the sediment was aerated, resulting in much lower P release rates (1.1 to 32.9 mgP m^–2 d^–1) over a period of 30 days. The influence of pH and nitrate was studied by adjusting pH and adding NO₃ ^– to the overlying water. Increasing pH positively affected P release rates and enhanced NO₃ ^– levels led to an increase of benthic P release, too.     

Nitrogen and phosphorus accumulation and cycling by Nymphoides peltata (Gmel.) O. Kuntze (Menyanthaceae)

In 1980, the seasonal changes in nitrogen and phosphorus concentration of various plant parts of Nymphoides peltata (Gmel.) O. Kuntze, together with aspects of nitrogen and phosphorus cycling by this species were studied in an oxbow lake of the river Waal (The Netherlands). The nitrogen and phosphorus stores of the water, seston, sediment and macrophyte compartments were assessed each month.The underground Nymphoides structures had high nitrogen and phosphorus concentrations before and after the main growing season, while during summer the aboveground plant parts had high nutrient contents. Nymphoides peltata accumulated maximum amounts of nitrogen (334 mmol m⁻²) and phosphorus (56.6 mmol m⁻²) in July. The upper layers of the bottom appeared to be an enormous nutrient reservoir (94–99% of total) of which the largest part was not directly available to Nymphoides. Nutrient uptake from the sediments by N. peltata is suggested by the fact that the bottom and/or interstitial water of the sample station devoid of rooted macrophytes, contained higher concentrations of nitrogen and phosphorus than that of the Nymphoides stands. The annual flux of nutrients from Nymphoides to the detritus compartment was estimated to be ca. 1200 mmol nitrogen and 164 mmol phosphorus per m² of littoral. During breakdown of the detritus there was a relatively fast net conversion of organically bound nitrogen and phosphorus to inorganic forms, especially at higher temperatures.Nymphoides has the potential to function as an important nitrogen and phosphorus pump, which regenerates sediment nutrients.     

Zooplankton structure in the Loosdrecht lakes in relation to trophic status and recent restoration measures

A five-year zooplankton study (1982–86) on three shallow and highly eutrophic lakes in the Loosdrecht area (The Netherlands) did not reveal any significant changes following the considerable reduction in external P-loading (from about 1.0 g to 0.3 g P m^–2 year^–1) since mid-1984.The recent annual fluctuations in the rotifer and crustacean densities are within the range of those found before the restoration measure became operative. A decrease in the average size of the crustaceans and an absence of large-bodied forms reflects an increased fish predation rather than a change in the quality or quantity of their sestonic food ( < 150 µm) which continues to be dominated by filamentous cyanobacteria and Prochlorothrix hollandica, a prochlorophyte discovered in these lakes recently.     

Aspects of aerobic mineralization during spring in Lake Vechten with special reference to the 14C-labelling technique

Aerobic mineralization, i.e. seston respiration, microbial breakdown of detritus and microbial assimilation-dissimilation of photosynthetically derived D(issolved) O(rganic) C(arbon) was measured in concentrated samples from the pelagic zone of Lake Vechten. The samples were described by cell numbers of dominant algae prior to concentrating by centrifuge with continuous rotor. The concentrated samples were incubated in the laboratory at in situ temperature and a light intensity of 30 W · m^–2 for measuring primary production and photosynthetically derived DOC. After filtration the particulate fraction was incubated in unlabelled lake water far measuring respiratory production of DI¹⁴C. Portions of the same particulate fraction were sterilized and incubated in unlabelled sample concentrate for measuring microbial breakdown of detritus. The seston respiration amounted to 19–30% loss of the particulate fraction over 15–45 h. Microbial breakdown of detritus amounted to 28–40% loss of the particulate fraction over 24–168 h. In both cases P(articulate) O(rganic) ¹⁴C(arbon) was transformed to mainly DI¹⁴C. Microbial assimilation-dissimilatioh of photosynthetically derived DOC could not be measured reliably.     

Clearance rates of bacteria by the rotifer Filinia longiseta (Ehrb.) measured using three tracers

The clearance rates (CRs) of bacteria by Filinia longiseta were measured at 19°C, both in situ in Lake Loosdrecht and in the laboratory during summer. The tracer particles used in the field were: (1) 0.51 µm fluorescent microspheres, and (2) fluorescently labelled bacteria (FLB). A third type of tracer particle, natural [methyl-³H]-thymidine-labelled bacteria (< 1.2 µm), were used as a radiotracer in a laboratory experiment. The uptake of the first two tracer-particle types was measured by microscopic examination of the rotifer guts. In the third case, the uptake of radioactivity was determined by liquid scintillation counting. The rate of uptake of the microspheres decreased 10 min after the start of the experiment, probably because the gut passage time was exceeded. Using a 5 min feeding time, the rate of uptake of microspheres was higher than that of the FLB, though the variation in the uptake in both cases was high. The ingestion rates and CRs of bacteria by F. longiseta based on the fluorescent tracers were: microspheres, 5115 bact.ind^–1 h^–1 and 0.368 µl ind^–1 h^–1; FLB, 2252 bact.ind^–1 h^–1 and 0.162 µl ind^–1 h^–1. The mean CR using the thymidine-labelled natural bacteria and a 10 min feeding time was 0.179 µl ind^–1 h^–1. Thus, the CR based on the microsphere method was twice as high as for the other two methods.     

Growth pattern and cellular nitrogen and phosphorus contents of the dinoflagellate Peridinium penardii (Lemm.) Lemm. causing a freshwater red tide in a reservoir

The population growth pattern and related changes in both the nitrogen and phosphorus contents in the cell of the dinoflagellate Peridinium penardii (Lemm.) Lemm., which formed a freshwater red tide in a reservoir, were studied in situ. An exponential increase with time in population density was found. A specific growth rate of 0.25 d^–1 was observed. The cellular content of phosphorus per cell decreased from 6.0 × 10^–5 µg to 9.2 × 10^–6 µg along with an increase in population density from 8.0 × 10² cells ml^–1 to 2.5 × 10⁴ cells ml^–1. A prominent change in the cellular nitrogen did not occur. Decreasing cell content and continuous uptake of phosphorus were advantageous for P. penardii to form a freshwater red tide under P-limited conditions.     

Water quality research in the Loosdrecht lakes: proposals for the follow-up restoration measures

Reduction of the external phosphorus load in 1984 did not lead to recovery of Loosdrecht lakes. Therefore the Provincial Water Authorities of the Province of Utrecht proposed, after consulting the research group Water Quality Research Loosdrecht lakes (WQL), measures to reduce the present external phosphorus loading from 0.35 to 0.1 g Pm^–2y^–1. Mathematical modelling suggested that such a reduction would benefit the water quality of the lakes. The results of the WQL investigation did not lead to additional measures (dredging, biomanipulation, chemomanipulation).     

Primary production of phytoplankton and standing crops of zooplankton and zoobenthos in hypertrophic Lake Teganuma

Primary production of phytoplankton and standing crops of zooplankton and zoobenthos were intensively surveyed in Lake Teganuma during May 1983–April 1984. The annual mean chlorophyll a concentrations were as high as 304 µg · l^–1–383 µg · l^–1. The daily gross primary production of phytoplankton was high throughout the year. The peak production rate was recorded in August and September, when blue-green algae bloomed. The annual gross primary production was estimated as 1450 g C · m^–2 · y^–1, extremely high as compared with other temperate eutrophic lakes. Zooplankton was predominantly composed of rotifers. The annual mean standing crop of zooplankton was 0.182 g C · m^–2 around the middle between the inlets and the outlet and was lower than in most other temperate eutrophic lakes. Zoobenthos was mostly composed of Oligochaeta and chironomids. The annual mean standing crop of zoobenthos ranged from 0.052 g C · m^–2 to 0.265 g C · m^–2, the lowest values among temperate eutrophic lakes, which is in contrast to the high primary production.