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.     

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.     

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 eutrophic Loosdrecht Lakes: Current ecological research and restoration perspectives

Features of the Loodsrecht Lakes, with emphasis on the main lake, are discussed with reference to restoration.Characteristics of the present situation are: (1) very low water transparency-Secchi-disc readings around 0.3 m occur in all seasons; (2) relatively small seasonal changes in sestonic matter; (3) important input of resuspended particles into the seston; (4) predominance of filamentous blue-green algae for most of the year; (5) relative scarcity of crustacean zooplankton, while rotifers are abundant; (6) poor development of littoral communities, and absence of benthic producers. The blue-green algae maintain high population density at very low growth rates: rates of loss are low. The zooplankton grazing rate is low due to inefficient filtering, but predation of larger crustaceans by fish may also be important. Studies on epipelon indicated that loss by deposition may be largely compensated by resuspension.Starting in 1984, the external phosphorus loading was markedly reduced. Results for 1984 and 1985 indicate that complementary measures are needed in order to improve water quality. Action should be directed towards increasing the phytoplankton turnover rates. Accelerated specific growth rate can be expected to accompany lower biomass, more successful competition by other algal groups, and enhancement of grazing pressure.Considering the shallowness of the system, promotion of littoral development and return of submerged vegetation may be important in establishing a new equilibrium of the system.     

Colonization and succession of macroinvertebrates in recently freshened Lake Volkerak-Zoom (The Netherlands)

Macroinvertebrates were studied during the first five years after closure and freshening of the lake system Volkerak-Zoom, two lakes interconnected by a channel. Before embankment, in 1987, these lakes were part of the tidal eastern Scheldt estuary in the south-western part of the Netherlands. Four months after embankment, the first freshwater species were found. In general, the colonization process followed the same pattern as was reported for newly created reservoirs. In Lake Volkerak-Zoom differences in colonization time were observed between distinguished depth zones. Within the chironomid assemblages, dominance of Chironomus muratensis was followed by a dominance of Procladius, Glyptotendipes and Cladotanytarsus. The succession was related to changes in phytoplankton concentrations. Although in 1991 changes in macrofauna composition were less pronounced as compared to previous years, succession was still going on.     

Control of phosphorus loading and flushing as restoration methods for Lake Veluwe,The Netherlands

As a result of high nutrient loading Lake Veluwe suffered from an almost permanent bloom of the blue-green algaOscillatoria agardhii Gomont. In 1979, the phosphorus loading of the lake was reduced from approx. 3 to 1 g P.m^–2.a^–1. Moreover, since then the lake has been flushed during winter periods with water low in phosphorus. This measure aimed primarily at interrupting the continuous algal bloom. The results of these measures show a sharp decline of total-phosphorus values from 0.40–0.60 mg P.l^–1 (before 1980) to 0.10–0.20 mg P.l^–1 (after 1980). Summer values for chlorophylla dropped from 200–400 mg.m^–3 to 50–150 mg.m^–3.The increase in transparency of the lake water was relatively small, from summer values of 15–25 cm before the implementation of the measures to 25–45 cm afterwards. The disappointing transparency values may be explained by the decreasing chlorophylla and phosphorus content of the algae per unit biovolume. Blue-green algae are gradually loosing ground. In the summer of 1985 green algae and diatoms dominated the phytoplankton for the first time since almost 20 years. To achieve the ultimate water quality objectives (transparency values of more than 100 cm in summer), the phosphorus loading has to be reduced further.     

Long-term changes in amphipod population dynamics in a temperate estuary following ecosystem restoration

The Mondego estuary (Portugal) has suffered severe ecological stress over the last two decades, as manifested in the replacement of seagrasses by opportunistic macroalgae, degradation of water quality and increased turbidity. A restoration plan was implemented in 1998, which aimed to reverse the eutrophication effects, and especially to restore the original natural seagrass (Zostera noltii) community. This article explores the long-term changes in Ampithoe valida and Melita palmata (Amphipoda) populations in response to eutrophication (with consequent seagrass loss and macroalgal proliferation) and to the subsequent restoration plan (with progressive seagrass recovery and macroalgal biomass decline). Until the early 1990s, high densities of A. valida and M. palmata were recorded in the Mondego estuary, especially during the occurrence of the macroalgal bloom and during all the periods in which green macroalgae were available. After the implementation of the restoration plan, species abundance, biomass and production levels decreased considerably due to the progressive decline of green macroalgae. This implied the virtual disappearance of the amphipod population, mainly A. valida. Distinct behaviours displayed by the two species could be related to different food strategies and habitat preferences. Ampithoe valida showed feeding preferences for ephemeral softer, filamentous or bladed algae (e.g. Ulva sp.) due to its high caloric content, using the Z. noltii bed only as a habitat for protection against predators or shelter from wave action. On the other hand, M. palmata did not suffer a strong decline in its population density, biomass and production, which may indicate that this species is probably not a primary consumer of green macroalgae and may readily shift to alternative ecological niches. Handling editor: P. Viaroli     

An assessment of the importance of emergent and floating-leaved macrophytes to trophic status in the Loosdrecht lakes (The Netherlands)

The potential importance of the six major emergent and floating-leaved macrophyte species in recycling of sediment phosphorus in the Loosdrecht lakes was studied. Representative plant samples were collected at the time of maximum biomass, and analysed for biomass and carbon, nitrogen and phosphorus contents. Species cover was determined by aerial photography.Total cover in the seven lakes studied ranged between 2 and 26 percent. For the four main species, biomass per unit area increased with lake trophic status. Consistent differences in C, N and P contents per unit biomass were not observed. Although cover values were small, significant amounts of C, N and P were contained in the macrophytes when compared with maximum sestonic content.Potential P loads from macrophyte decay were calculated. In Lake Loosdrecht, the P load represented 15 percent of current external P inputs. The potential importance of macrophyte decay to P recycling in the other lakes is greater.Decay of macrophyte species at the end of the growing season appears to affect autumnal nutrient and chlorophyll a levels in the water column of some lakes. The re-establishment of submerged species following lake restoration may increase the importance of this pathway in the lakes.     

Monthly mass balances for compartments of the Loosdrecht Lakes system: Approach and preliminary results

Water quality modelling in the framework of the Water Quality Research Loosdrecht Lakes requires mass balances covering periods of one month or less for at least three of the lake compartments of the Loosdrecht Lakes system: the Loosdrecht Lakes, L. Vuntus and L. Breukeleveen. The hydrological system of the Loosdrecht area underlying these mass balances is described. Data available on the various mass balance items are surveyed, as well as the methods considered to calculate those items that do not lend themselves to direct measurement.The system will be modelled by means of a series of interdependent spreadsheets. Calibration of this model to the 1983–1986 data set should render it suitable to calculate the mass balances retrospectively,i.e. in the period 1975–1982. The model in its present form cannot be used to predict mass balances.Masses of Cl, P and N stored in the three main lake compartments of interest at consecutive sampling dates in 1983 and 1984 are graphically presented and compared with roughly estimated biannual balances of the same substances for the aggregate of the three lake compartments.Not unexpectedly, the distribution within the system of the conservative substance Cl is a function of the differences between the lake compartments in mixing ratios of high-Cl to low-Cl inputs.The amounts of the nutrients P and N stored in the lake water follow a rather erratic course. Though it is possible to idealize this to a seasonal cycle, the two-year period dealt with by this study is too short to identify such a cycle with certainty. From the large residual terms of the biannual nutrient balances it is obvious that the not directly quantifiable processes like sedimentation, release from the sediment and (presumably) denitrification that are lumped into those residual terms, are of major importance to the amounts of nutrients in the lake water.     

Vertical distributions of zooplankton and population dynamics of Daphnia in a meromictic lake

Vertical distributions of zooplankton were studied in relation to profiles of temperature, oxygen and chlorophyll a in Roi Lake, a small meromictic lake in central Alberta. Zooplankton were distributed fairly evenly through the oxygenated part of the water column in early summer, but a gradual descent of several species became evident in June. The vertical distribution of chlorophyll was dominated by a huge peak at the 8- to 9-m-deep chemocline. the location of a plate of photosynthetic sulfur bacteria. Ambient concentration of chlorophyll was a poor predictor of the numbers of zooplankton and the fecundity of Daphnia pulicaria at different depths, and per capita birth rates of Daphnia were usually highest in the surface waters. The reproductively disadvantageous restriction of daphnids to deep water by late summer and their catastrophic decline in the face of high ambient concentrations of chlorophyll suggest that factors other than temperature and food supply are important in influencing the dynamics and distribution of zooplankton in this lake.     

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.     

Restoration of Botshol (The Netherlands) by reduction of external nutrient load: Recovery of the Characean community

The formerly rich characean community in Botshol with six species of which the rareNitellopsis obtusa andChara hispida dominated at many sites, decreased to only two species,Chara globularis andC. connivens, in the period 1980–1988. The macrophyteNajas marina also remained at some sites, and the aquatic mossFontinalis antipyretica and the filamentous algaVaucheria dichotoma predominated at many sites. These phenomena may have been due to eutrophication by the inlet of polluted water. This process of eutrophication was stopped by restoration measures in 1989, resulting in a lower phosphorus concentration (ca. 0.024 mg l^–1) and a higher water transparency. Immediately after these measures the Characeae community increased strongly in abundance and number of species. During the summer of 1990, and especially of 1991, a spectacular growth occurred ofChara connivens. Chara connivens was often accompanied byChara hispida. Other species with scattered occurrence wereChara aculeolata, C. aspera, C. contraria andC. Globularis. The reasons for the shift in dominance fromNitellopsis obtusa toChara connivens are discussed. One of the reasons may be the recent higher chloride content which is one of the consequences of the restoration measures.     

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.     

The relation of biotic and abiotic interactions to eutrophication in Tjeukemeer,The Netherlands

During the summer months of 1974–1985 chlorophyll-a and total P concentration, biomass of Daphnia hyalina, smelt Osmerus eperlanus, bream Abramis brema and pikeperch Stizostedion lucioperca, water temperature and water intake from lake IJsselmeer were monitored in Tjeukemeer. During this period there were manipulations with the bream and pikeperch stocks as a consequence of the termination of a gill-net fishery in 1977, and larval smelt immigrated each year from the large lake IJsselmeer and contributed largely to the yearly smelt recruitment.The correlation matrix of the nine variables mentioned above showed a positive correlation between bream and chlorophyll-a, but surprisingly a negative one between smelt and chlorophyll-a. The latter can only be explained when smelt is the dependent variable. In a multi-linear regression there was a negative effect of temperature, chlorophyll a and pikeperch on smelt and a positive effect of water intake. Daphnia hyalina was negatively influenced by the biomass of smelt and the water intake of lake IJsselmeer. The positive relation of Daphnia hyalina and chlorophyll-a was probably related to better survival chances of D. hyalina in an Oscillatoria-rich environment when smelt is the most important predator. An increasing biomass of bream coincided with higher total-P levels and probably contributed to higher chlorophyll-a levels.     

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.     

Recolonization and restoration of upper intertidal Fucus gardneri (Fucales,Phaeophyta) following the Exxon Valdez oil spill

The Exxon Valdez oil spill in March 1989 and subsequent cleanup caused injury to intertidal Fucus gardneri populations especially in the upper intertidal. A survey in 1994 in Prince William Sound, Alaska showed that the upper boundary of Fucus populations at oiled sites was still an average of 0.4 m lower than the upper boundary at unoiled sites.Restoration of severely damaged Fucus populations was started on a small-scale at a heavily oiled rocky site in Herring Bay, Prince William Sound. Experiments employed mats of biodegradable erosion control fabric to act as a substratum for Fucus germlings and to protect germlings from heat and desiccation stress. A series of plots was covered with mats made from a resilient coconut-fiber fabric in June 1993. Half of the mats were inoculated with Fucus zygotes. A series of uncovered control plots was also monitored. There was no enhancement of Fucus recruitment on the rock surfaces under the mats. Dense populations of Fucus developed on the surface of all of the mats by the summer of 1994. The natural rock surfaces in the control plots, both inoculated and not, were barren of macroscopic algal cover. By September 1994, the juvenile thalli on the mats were approximately 2 cm in length. Inoculating the mats had an effect only in the upper region of the intertidal. It is expected that the thalli will become fertile during the 1995 season. These thalli may serve as a source of embryos to enhance the recovery of new Fucus populations in this high intertidal area.     

Biomanipulation and its feasibility for water quality management in shallow eutrophic water bodies in The Netherlands

Biomanipulation as a tool for lake restoration is discussed mainly using literature data. It is based on the exploitation of the interactions both within and between the trophic levels in an aquatic ecosystem. Important among the interactions are: competition for light and nutrients between aquatic macrophytes and phytoplankton and among different phytoplankton species; grazing by planktonic and benthic filter feeders; and size-selective predation by fish. In several case studies biomanipulation has proved to be successful in restorating mildly eutrophic small waterbodies. However, for long-term stability of the restored ecosystems supplementary measures like reducing the external nutrient loadings are needed. The feasibility of the different biomanipulation measures to improve the water quality in shallow Dutch lakes is discussed. Preliminary results on biomanipulation experiments in enclosures withOscillatoria agardhii and the benthic filter feederDreissena polymorpha are given.     

Phytoplankton growth rates in the freshwater tidal reaches of the Schelde estuary (Belgium) estimated using a simple light-limited primary production model

During the course of 1996, phytoplankton was monitored in the turbid, freshwater tidal reaches of the Schelde estuary. Using a simple light-limited primary production model, phytoplankton growth rates were estimated to evaluate whether phytoplankton could attain net positive growth rates and whether growth rates were high enough for a bloom to develop. Two phytoplankton blooms were observed in the freshwater tidal reaches. The first bloom occurred in March and was mainly situated in the most upstream reaches of the freshwater tidal zone, suggesting that it was imported from the tributary river Schelde. The second bloom occurred in July and August. This summer bloom was situated more downstream in the freshwater tidal reaches and appeared to have developed within the estuary. A comparison between phytoplankton growth rates estimated using a simple primary production model and flushing rate of the water indicated that no net increase in phytoplankton biomass was possible in March while phytoplankton could theoretically increase its biomass by 20% per day during summer. Chlorophyllaconcentrations at all times decreased strongly at salinities between 5–10 psu. This decline was ascribed to a combination of salinity stress and light limitation. Phytoplankton biomass and estimated annual net production were much higher in the freshwater tidal zone compared to the brackish reaches of the estuary (salinity > 10 psu) despite mixing depth to euphotic depth ratios being similar. Possible reasons for this high production include high nutrient concentrations, low zooplankton grazing pressure and import of phytoplankton blooms from the tributary rivers.