E. Van Donk

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.

Impact of the fungicide carbendazim in freshwater microcosms. II. Zooplankton, primary producers and final conclusions

Effects of chronic application of the fungicide Derosal(R) (active ingredient carbendazim) were studied in indoor macrophyte-dominated freshwater microcosms. The concentrations (0, 3.3, 33, 100, 330 and 1000 µg/l) were kept at a constant level for 4 weeks. This paper is the second of a series of two; it describes the effects on zooplankton and primary producers and presents an overall discussion. The zooplankton community was negatively affected by the three highest treatment levels (NOEC(community)=33 µg/l). At higher treatment levels Cladocera taxa were completely eliminated, while Copepod numbers were reduced. Rotatoria taxa decreased (Keratella quadrata and Lecane sp.) or increased in abundance (Testudinella parva) at the highest treatment level only. Due to the reduced grazing pressure, the abundance of some phytoplankton taxa and the chlorophyll-a content of the phytoplankton increased at the three highest treatment levels (NOEC(community)=33 µg/l). This effect was not observed for the periphyton, most probably because the reduced grazing pressure was compensated by the increased abundance of some snail species such as Lymnaea stagnalis and Physella acuta. At the end of the experimental period the biomass of the macrophyte Elodea nuttallii was significantly elevated at the two highest treatment levels. It is hypothesised that carbendazim might have caused, directly or indirectly, the removal of pathogene organisms from the macrophyte.

Long-term responses to fish-stock reduction in small shallow lakes: interpretation of five-year results of four biomanipulation cases in The Netherlands and Denmark

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).

The ecological stoichiometry of toxins produced by harmful cyanobacteria: an experimental test of the carbon-nutrient balance hypothesis

The elemental composition of primary producers reflects the availability of light, carbon and nutrients in their environment. According to the carbon-nutrient balance hypothesis, this has implications for the production of secondary metabolites. To test this hypothesis, we investigated a family of toxins, known as microcystins, produced by harmful cyanobacteria. The strain Microcystis aeruginosa HUB 5-2-4, which produces several microcystin variants of different N:C stoichiometry, was cultured in chemostats supplied with various combinations of nitrate and CO(2). Excess supply of both nitrogen and carbon yielded high cellular N:C ratios accompanied by high cellular contents of total microcystin and the nitrogen-rich variant microcystin-RR. Comparable patterns were found in Microcystis-dominated lakes, where the relative microcystin-RR content increased with the seston N:C ratio. In total, our results are largely consistent with the carbon-nutrient balance hypothesis, and warn that a combination of rising CO(2) and nitrogen enrichment will affect the microcystin composition of harmful cyanobacteria.

Effects of grazing by fish and waterfowl on the biomass and species composition of submerged macrophytes.

Biomanipulation improved water transparency of Lake Zwemlust (The Netherlands) drastically. Before biomanipulation no submerged vegetation was present in the lake, but in summer 1987, directly after the measure, submerged macrophyte stands developed following a clear-water phase caused by high zooplankton grazing in spring. During the summers of 1988 and 1989 Elodea nuttallii was the most dominant species and reached a high biomass, but in the summers of 1990 and 1991 Ceratophyllum demersum became dominant. The total macrophyte biomass decreased in 1990 and 1991. In 1992 and 1993 C. demersum and E. nuttallii were nearly absent and Potamogeton berchtholdii became the dominant species, declining to very low abundance during late summer. Successively algal blooms appeared in autumn of those years reaching chlorophyll-a concentrations between 60–130 µg l−1. However, in experimental cages placed on the lake bottom, serving as exclosures for larger fish and birds, E. nuttallii still reached a high abundance during 1992 and 1993. Herbivory by coots (Fulica atra) in autumn/winter, and by rudd (Scardinius erythrophthalmus) in summer, most probably caused the decrease in total abundance of macrophytes and the shift in species composition.

Differential sensitivity of green algae to allelopathic substances from Chara

Three short-term laboratory experiments were conducted to investigate allelopathic effects of a mixture of Chara globularis var. globularis Thuillier and Chara contraria var. contraria A. Braun ex Kützing on three different green algae. Single phytoplankton species were exposed to filtered water originating from charophyte cultures. Phytoplankton growth was monitored by determination of chlorophyll concentrations in batch cultures. The change in chlorophyll concentration during the experiments was analysed with a logistic growth model, resulting in an estimate of the exponential growth rate and the duration of the lag phase of the single green algae. The results indicate allelopathic effects of Chara on the growth of the green algae Selenastrum capricornutum Printz and Chlorella minutissima Fott et Nováková, whereas Scenedesmus obliquus (Turpin) Kützing did not seem to be affected. The exponential growth rate of S. capricornutum decreased 7% in the presence of water from a charophyte culture, while the growth rate of C. minutissima decreased with 3%. The allelopathic effect of Chara did not increase when the green alga C. minutissima was P-limited. The effect of Chara was different when young sprouts were used. With young sprouts the duration of the lag phase of C. minutissima was extended (25%), whilst for old plants the growth rate of this green alga decreased. Although the inhibiting effect of charophytes on specific phytoplankton species is rather small, the differential sensitivity of the species to Chara might influence the composition and biomass of phytoplankton communities in the field.

Reduced digestibility of UV-B stressed and nutrient-limited algae by Daphnia magna

Daphnia magna was fed the green alga Selenastrum capricornutum, cultured under four different growth conditions: (1) phosphorus limitation, (2) nitrogen limitation, (3) UV-B irradiation, and (4) no nutrient limitation, no UV-B irradiation. Contrary to non-limited algal cells, nutrient-limited cells were not efficiently assimilated. Especially, P-limited cells passed through the gut mostly intact, while N-limited cells were partly assimilated. Also, algae exposed to moderate doses of UV-B radiation (0.3 mW cm-2 of UV312) were less efficiently assimilated after being grazed. Digestibility of the algae decreased with increased UV-B exposure time. Nutrient-limited and UV-B stressed algal cells increased in volume and became granular in appearance. These changes in the algal cells, combined with changed cell wall properties, most probably reduced their digestibility.

First attempt to apply whole-lake food-web manipulation on a large scale in The Netherlands

Lake Breukeleveen (180 ha, mean depth 1.45 m), a compartment of the eutrophic Loosdrecht lakes system, was selected to study the effects of whole-lake foodweb manipulation on a large scale. In Lake Loosdrecht (dominated by filamentous cyanobacteria), due to water management measures taken from 1970–1984 (sewerage systems, dephosphorization) the external P load has been reduced from 1.2 g m−2 y−1 to 0.35 g m−2 y−1. The water transparency (Secchi-depthca. 30 cm), however, has not improved. The aim of the food-web manipulation in Lake Breukeleveen was not only to improve the light climate of the lake, but also to study if the successfull effects observed in small lakes (a few ha) can be upscaled. In March 1989 the standing crop of planktivorous and bentivorous fish populations was reduced by intensive fishery, fromca. 150 kg ha−1 toca. 57 kg ha−1. The lake was made unaccessible to fish migrating from the other lakes and it was stocked with large-sized daphnids and 0+ pike. However, water transparency did not increase in the following summer and autumn 1989, which is in contrast with great improvement in the light conditions previously observed in smaller lakes. The main explanations for the negative outcome in Lake Breukeleveen are: 1) the rapid increase of the planktivorous fish biomass and carnivorous cladocerans, predating on the zooplankton community; 2) suppression of the large daphnids by the high concentrations of filamentous cyanobacteria; 3) high turbidity of the lake due to resuspension of bottom material induced by wind, unlike in smaller lakes, and thus inability of submerged macrophytes to develop and to stabilize the ecosystem.

The first biomanipulation conference: a synthesis.

At the First Biomanipulation Conference held in Amsterdam (8–11 August 1989), studies presented considered mainly trophic interactions in lakes, enclosures and laboratory systems. Studies on the interactions between phytoplankton and zooplankton emphasized the edibility of the phytoplankton in relation to the zooplankton size structure and the trophic state. Most lake experiments involved 50–100% reduction in fish standing stock or alternatively heavy stocking with piscivorous fish. The most dramatic effects of biomanipulation were found in shallow, eutrophic lakes which exhibited radical changes in ecosystem structure because of changes in light climate and consequently, luxuriant development of macrophytes. There was still much controversy about the top-down effects in relation to trophic state, especially those concerning the role of fish and zooplankton in the development and succession of phytoplankton. Many experiments showed indirect effects within the food web, emphasizing the importance of feedbacks and the complexity of the food web rather than the simplicity of the food chain. The stabilizing effects of refugia for zooplankton and fish on the ecosystem were stressed. Shallow lakes responded generally more rapidly to biomanipulation and this was most successfully accomplished when TP concentration was<50µg l−1, even though in a few cases at 10–20 fold higher TP concentrations (mostly PO4-P lakes) the results achieved could be maintained for two or more summers. For a guaranteed success of the measures an almost complete removal of fish appeared to be indispensible; moreover in many cases removal of benthivorous fish appeared to be even more important than that of planktivorous fish.

Hydrophyte-macroinvertebrate interactions in Zwemlust, a lake undergoing biomanipulation

In two years after biomanipulation of Lake Zwemlust (The Netherlands), macrophytes (helophytes, elodeids) and filamentous algae developed luxuriantly in the lake. They influenced the structure of macroinvertebrate communities inhabiting them. Macrophytes and algae, by changing environmental and trophic conditions, also affected the composition of macrozoobenthos. Vascular plants served as an important source of food for zoobenthos and phytofauna, mainly after they were decomposed. Filamentous algae were consumed readily alive by many animals. Invertebrates appeared to be important as a potential nutrient source for hydrophytes.