Macrophytes moderate the taxonomic and functional composition of phytoplankton assemblages during a nutrient loading experiment

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On body size and habitat selection in rotifers in a macrophye-dominated lake Budzyńskie,Poland

Lake Budzyńskie is shallow, freshwater lake with a well-developed and differentiated macrophytic vegetation. Zooplankton samples were collected from five stations: two of them in submerged macrophytes (Chara and Myriophyllum), one in the zone of floating leaves (Potamogeton), a rush station (Typha) and one in the open water surrounding the vegetation beds. The mean Rotifera densities differed significantly between the lake parts. Furthermore, different habitats were characterised by differences in body size with the exception of the middle body size group (Keratella cochlearis, Polyarthra vulgaris and Trichocerca similis), which was dominated by limnetic representatives. However, in all the other size-dependent groups both stands of submerged macrophytes were characterised by much higher densities than other zones. Additionally, body size within the examined habitats significantly differed. Thus, the size structure of Rotifera communities was directly related to morphological and spatial structures of the substrata. Two groups of habitats were distinguished: the first one consisting of open water and two vegetated zones of less complicated structure (Potamogeton and Typha), and the second of more complex submerged macrophyte species (Chara and Myriophyllum). The differentiation of the architecture of macrophytes affected the nutritional conditions and refuge effectiveness of these habitats.     

Representation of aquatic vegetation change by plant macrofossils in a small and shallow freshwater lake

We explored spatial and temporal relationships between contemporary aquatic vegetation and surface sediment macrofossil remains in a small, shallow, English lake (Green Plantation Pond). The aquatic vegetation of Green Plantation Pond underwent a marked compositional change after 2005 with a shift from Elodea spp.-Potamogeton pusillus-Chara spp. to Ceratophyllum spp.-Chara spp.-Potamogeton crispus dominance. By comparing macrophyte and plant macrofossil distributions at multiple, closely spaced points in Green Plantation Pond for 2000 and 2008–2009, we studied the ability of macrofossils to track this major aquatic vegetation change. Representation of macrophytes by macrofossils was high with 63 and 76 % of extant plant species recorded by macro-remains in the 2000 and 2009 sediment surveys respectively. Nevertheless, plants were both over-represented (Nitella flexilis, Chara spp. and Zannichellia palustris) and under-represented (Ranunculus sect. Batrachium, Potamogeton spp.) in the sediment record in terms of relative macrofossil abundances and the number of occupied sample points. The study also revealed a lack of preservation of Elodea spp. leaf remains in the second (2009) survey compared to the first (2000) probably due to a longer time interval (5 vs. 10 months) between macrophyte and sediment sampling. Nevertheless, the macrofossils reliably recorded both the main shift in the contemporary vegetation (e.g. especially increases in Ceratophyllum spp. and P. crispus abundance) and other more subtle floristic changes (e.g. increases in Myriophyllum spicatum and Lemna spp.) exceptionally well. This study highlights the huge potential of macrofossils for tracking sub-decadal changes in the aquatic vegetation of small, shallow lakes.     

Experimental evidence for enhanced top-down control of freshwater macrophytes with nutrient enrichment

The abundance of primary producers is controlled by bottom-up and top-down forces. Despite the fact that there is consensus that the abundance of freshwater macrophytes is strongly influenced by the availability of resources for plant growth, the importance of top-down control by vertebrate consumers is debated, because field studies yield contrasting results. We hypothesized that these bottom-up and top-down forces may interact, and that consumer impact on macrophyte abundance depends on the nutrient status of the water body. To test this hypothesis, experimental ponds with submerged vegetation containing a mixture of species were subjected to a fertilization treatment and we introduced consumers (mallard ducks, for 8 days) on half of the ponds in a full factorial design. Over the whole 66-day experiment fertilized ponds became dominated by Elodea nuttallii and ponds without extra nutrients by Chara globularis. Nutrient addition significantly increased plant N and P concentrations. There was a strong interactive effect of duck presence and pond nutrient status: macrophyte biomass was reduced (by 50 %) after the presence of the ducks on fertilized ponds, but not in the unfertilized ponds. We conclude that nutrient availability interacts with top-down control of submerged vegetation. This may be explained by higher plant palatability at higher nutrient levels, either by a higher plant nutrient concentration or by a shift towards dominance of more palatable plant species, resulting in higher consumer pressure. Including nutrient availability may offer a framework to explain part of the contrasting field observations of consumer control of macrophyte abundance.     

Size spectra and abundance of planktonic ciliates within various habitats in a macrophyte-dominated lake (Eastern Poland)

Body size, community structure, abundance and biomass of ciliates were compared in various stands of macrophytes in a macrophyte-abundant shallow lake in Eastern Poland. Samples were collected in belts of Phragmites, Typha, Ceratophyllum, Elodea, Stratiotes and Chara. Additionally, protozooplankton was collected from the open water zone surrounding the vegetation belts. Differences in numbers of ciliate taxa between micro-sites were statistically significant. The highest numbers were found in Chara and Ceratophyllum stands, lower numbers in Stratiotes and Elodea stands and the lowest in the open water, Phragmites and Typha areas. Ciliate biomass was, like density, significantly higher in submerged macrophytes than in emergent macrophytes and open water zones. Based on differences in macrophyte structure, two groups of habitats with similar patterns of size-related ciliate distribution were distinguished. The first group consisted of two vegetated zones of sparse stem structure (Phragmites and Typha) and the open water zone, the second group comprised submerged macrophyte species, which were more dense and complex. Generally, the abundance of ciliates correlated positively with total suspension solid (TSS) and total organic carbon (TOC) concentrations. In the Chara and Ceratophyllum stands, relations between ciliate numbers, TSS and TOC were stronger.     

Effects of fish and nutrient additions on food-web stability in a charophyte-dominated lake

1. The response of major food‐web constituents to combinations of nutrient addition and zooplanktivorous fish abundance was tested during two subsequent years in the shallow charophyte‐dominated lake Naardermeer in the Netherlands, using in situ enclosures. 2. Treatment effects differed sharply between study years. In 1998, when the summer temperature was low (17–21 °C), high algal biomass only developed at high nutrient levels in the presence of fish, but with no major effect on Chara biomass. In 1999, when the summer temperature was relatively high (20–25 °C), algal blooms occurred at high nutrient levels regardless of fish abundance, and were associated with a drastic decline in Chara biomass. 3. Differences between years in temperature and initial zooplankton composition and biomass were likely to contribute to the varying relative importance of top‐down and bottom‐up effects in these enclosure experiments. 4. The results suggest that when nutrient loads are increased towards levels where the macrophyte‐dominated state is being destabilised, a ‘switch’ is more likely to occur in warm summers.     

Long‐term allelopathic control of phytoplankton by the submerged macrophyte Elodea nuttallii

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Relations between trophic state indicators and plant biomass in Florida lakes

We collected quantitative data on macrophyte abundance and water quality in 319 mostly shallow, polymictic, Florida lakes to look for relationships between trophic state indicators and the biomasses of plankton algae, periphyton, and macrophytes. The lakes ranged from oligotrophic to hypereutrophic with total algal chlorophylls ranging from 1 to 241 mg m^–3. There were strong positive correlations between planktonic chlorophylls and total phosphorus and total nitrogen, but there were weak inverse relationships between the densities of periphyton and the trophic state indicators total phosphorus, total nitrogen and algal chlorophyll and a positive relationship with Secchi depth. There was no predictable relationship between the abundance of emergent, floating-leaved, and submersed aquatic vegetation and the trophic state indicators. It was only at the highest levels of nutrient concentrations that submersed macrophytes were predictably absent and the lakes were algal dominated. Below these levels, macrophyte abundance could be high or low. The phosphorus–chlorophyll and phosphorus–Secchi depth relationships were not influenced by the amounts of aquatic vegetation present indicating that the role of macrophytes in clearing lakes may be primarily to reduce nutrient concentrations for a given level of loading. Rather than nutrient concentrations controlling macrophyte abundance, it seems that macrophytes acted to modify nutrient concentrations.     

Microhabitat–zooplankton relationship in extensive macrophyte vegetations of eutrophic clear-water ponds

The influence of different macrophyte taxa or growth forms on biological and environmental variables is often analysed in one-lake studies. However, the unique combination of non-vegetational characteristics of a waterbody, i.e. its site identity, can be an influential factor in itself, shaping the measured parameters irrespective of the presence or absence of certain macrophyte species. In this situation, the relative strengths of all factors can be determined best in a study that explicitly accounts for differences in the identity of the waterbodies. Several functional macrophyte groups are known to provide a potent microinvertebrate refuge or permanent habitat. The objective of this study was to detect patterns in the zooplankton assemblages associated with different extensive habitats of macrophyte species and to relate these patterns to three major factors: the microhabitat, the pond identity and the seasonality in the warmer months of the year. Five ponds located in the Woluwe catchment of the Brussels-Capital Region (Belgium) were studied monthly for macrophyte and zooplankton characteristics from July until October 2005. The vegetation in the clear ponds was characterized by extensive monospecific stands (Ceratophyllum, Chara, Nitella, Potamogeton, Nuphar and filamentous algae). Zooplankton could be analysed in seven different vegetation types and in the open water zones and contained a total of 17 cladoceran and 27 rotifer genera. Principal components analysis (PCA) ordination of zooplankton communities showed a seasonal gradient and a tendency to group within-pond habitats, although they differed in macrophyte species and habitat structure. Despite the absence of clustering of similar microhabitats across ponds, percent volume infested (PVI), vegetation biomass density and Daphnia length (used as a proxy for fish predation pressure) contributed significantly positive to the Shannon zooplankton biodiversity indices. Moreover, densities of most zooplankton subgroups and of total zooplankton were significantly and positively related to PVI. It is assumed that in eutrophic ponds, extensive, often monospecific macrophyte vegetations provide an ecological environment suitable for both macrophyte-associated species and migrating pelagic zooplankton, thereby maintaining a high microinvertebrate biodiversity.     

Fish community structure response to major habitat changes within the littoral zone of an estuarine coastal lake

Synopsis The littoral environment and fish fauna of Swartvlei, an estuarine lake, was monitored for four years during which major habitat changes occurred. Initially (1979) the zone was dominated by the submerged macrophytes Potamogeton pectinatus, Chara globularis and Lamprothamnium papulosum. This plant community was replaced by filamentous algal mats during 1980 and with the disappearance of these mats in 1981 the littoral zone was transformed into a sandy habitat. There was a highly significant decline in the numbers of fishes in the littoral zone between the macrophyte and sand phases but no significant decrease in fish biomass between the two phases. Analysis of gill net catches revealed an increase in the CPUE of the family Mugilidae between the macrophyte and sand phases but a decline in the CPUE of vegetation associated species such as Monodactylus falciformis and Rhabdosargus holubi over the same period. The increase in mullet stocks during the sand phase was attributed to epipsammic micro-algal production and the input of allochthonous detritus during the 1981 floods. The three fish species diversity indices used in this study showed minor fluctuations between the habitat phases and these variations were related to changes in the equitability of distribution between the individual species within each habitat type. The numbers of fish species recorded during the macrophyte, algal mat and sand phases varied by less than 20%. The resilience of estuarine fishes to major alterations in their environment was illustrated by the fact that all fish species recorded at the beginning of 1979 were present at the end of 1982, despite major habitat and food resource changes.     

Primary producers in a Pampean stream: temporal variation and structuring role

Low current velocities, high nutrient levels, the lack of riparian forest vegetation, and the development of dense and rich macrophyte communities characterize Pampean streams. The objective of this study was to describe the main physical, chemical, and biological characteristics of a headwater Pampean stream as well as to analyze the role of macrophytes and phytobenthos. The study was conducted in a stream considered to be not much disturbed by human activities. Samples of water and organisms (macrophytes, benthic algae and invertebrates) were taken monthly for 14 months in two sampling stations, in fast flow and slow flow sites. Macrophyte biomass and diversity increased in spring and summer, and they decreased in autumn, when the plant community was greatly affected by an important flood. Phytobenthos biomass was lower in late summer, possibly due to the establishment of a dense cover of the floating macrophyte Lemna gibba L. Density of amphipods and gastropods greatly increases in spring and summer, jointly with the macrophyte development. Analysis of correlation showed that current velocity is the most important factor influencing macrophyte biomass and phytobenthos structure, while depth, nutrients, and herbivores are linked factors. Pampean streams could be considered systems dynamically fragile, because habitat heterogeneity is generated by aquatic vegetation, a substratum that varies along time.     

Spatial distribution of epiphytic chironomid larvae in a shallow macrophyte-dominated lake: effect of macrophyte species and food resources

Four submerged macrophytes were analysed with their respective assemblages of chironomid and algal taxa. Studies were made bimonthly, from January to November, in a shallow macrophyte-dominated lake. The periphytic algal community was represented by chlorophytes, diatoms and cyanobacteria; diatoms showed the highest relative abundance on the macrophyte species during the study. Algal biomass was affected by the season, and in January the highest values were found on Stratiotes aloides L., Potamogeton lucens L., Myriophyllum spicatum L.; in July on Chara aculeolata Kütz. The lowest algal biomass, on most of the macrophytes studied, was found in September. Larvae of Dicrotendipes sp., Diplocladius cultriger Kieffer, Endochironomus albipennis gr., Endochironomus impar (Walker), Glyptotendipes sp. and Paratanytarsus austriacus Kieffer were dominant within the chironomid community. The spatial distribution of chironomids between macrophytes species was related to periphytic algae biomass and certain environmental variables (temperature, water transparency, biomass of chlorophytes and diatoms). Analysis of chironomid diet and the results of canonical correspondence analysis confirmed this relationship. The strongest chironomid-algae relationships were observed on S. aloides and P. lucens.     

Seasonal dynamics of macrophytes and phytoplankton in shallow lakes: a eutrophication‐driven pathway from plants to plankton?

1. Seasonal relationships between macrophyte and phytoplankton populations may alter considerably as lakes undergo eutrophication. Understanding of these changes may be key to the interpretation of ecological processes operating over longer (decadal‐centennial) timescales. 2. We explore the seasonal dynamics of macrophytes (measured twice in June and August) and phytoplankton (measured monthly May–September) populations in 39 shallow lakes (29 in the U.K. and 10 in Denmark) covering broad gradients for nutrients and plant abundance. 3. Three site groups were identified based on macrophyte seasonality; 16 lakes where macrophyte abundance was perennially low and the water generally turbid (‘turbid lakes’); 7 where macrophyte abundance was high in June but low in August (‘crashing’ lakes); and 12 where macrophyte abundance was high in both June and August (‘stable’ lakes). The seasonal behaviour of the crashing and turbid lakes was extremely similar with a consistent increase in nutrient concentrations and chlorophyll‐a over May–September. By contrast in the stable lakes, seasonal changes were dampened with chlorophyll‐a consistently low (<10–15 μg L^?1) over the entire summer. The crashing lakes were dominated by one or a combination of Potamogeton pusillus, Potamogeton pectinatus and Zannichellia palustris, whereas Ceratophyllum demersum and Chara spp. were more abundant in the stable lakes. 4. A long‐term loss of macrophyte species diversity has occurred in many shallow lakes affected by eutrophication. One common pathway is from a species‐rich plant community with charophytes to a species‐poor community dominated by P. pusillus, P. pectinatus and Z. palustris. Such compositional changes may often be accompanied by a substantial reduction in the seasonal duration of plant dominance and a greater tendency for incursions by phytoplankton. We hypothesise a slow‐enacting (10–100 s years) feedback loop in nutrient‐enriched shallow lakes whereby increases in algal abundance are associated with losses of macrophyte species and hence different plant seasonal strategies. In turn such changes may favour increased phytoplankton production thus placing further pressure on remaining macrophytes. This study blurs the distinction between so‐called turbid phytoplankton‐dominated and clear plant‐dominated shallow lakes and suggests that plant loss from them may be a gradual process.     

Differences in tolerance of pondweeds and charophytes to vertebrate herbivores in a shallow Baltic estuary

It has been suggested that herbivorous waterfowl may be important in shaping aquatic plant communities in shallow wetlands. As such, a shift from canopy forming pondweeds to bottom-dwelling charophytes in a formerly turbid pondweed dominated lake has been partly attributed to waterfowl herbivory. Here we study the separate and combined effects of both belowground herbivory in spring by whooper swans and Bewick ‘s swans, and grazing in summer by waterfowl and fish on the community composition in a shallow Baltic estuary during one year. The macrophyte community was dominated by charophytes (mainly Chara aspera) with Potamogeton pectinatus and Najas marina present as subdominants. Other species were rare. Both spring and summer herbivory had no effect on total plant biomass. However, P. pectinatus was more abundant in plots that were closed to spring and summer herbivores. N. marina was more abundant in grazed plots, whereas Chara spp. biomass remained unaffected. Probably belowground propagules of both C. aspera and P. pectinatus were consumed by swans but since C. aspera bulbils were numerous it may have compensated for the losses. P. pectinatus may not have fully recovered from foraging on tubers and aboveground biomass. Our results are in line with other studies in Chara dominated lakes, which found no effect of grazing on summer aboveground Chara biomass, whereas several studies report strong effects of herbivory in lakes dominated by P. pectinatus.     

Impacts of sediment type on the performance and composition of submerged macrophyte communities

To restore deteriorated lake ecosystems, it is important to identify environmental factors that influence submerged macrophyte communities. While sediment is a critical environmental factor for submerged macrophytes and many studies have examined effects of sediment type on the growth of individual submerged macrophytes, very few have tested how sediment type affects the growth and species composition of submerged macrophyte communities. We constructed submerged macrophyte communities containing four co-occurring submerged macrophytes (Hydrilla verticillata, Myriophyllum spicatum, Ceratophyllum demersum and Chara fragilis) and subjected them to three sediment treatments, i.e., clay, a mixture of clay and quartz sand at a volume ratio of 1:1 and a mixture at a volume ratio of 1:4. Compared to the clay, the 1:1 mixture treatment greatly increased overall biomass, number of shoot nodes and shoot length of the community, but decreased its diversity. This was because it substantially promoted the growth of H. verticillata within the community, making it the most abundant species in the mixture sediment, but decreased that of M. spicatum and C. demersum. The sediment type had no significant effects on the growth of C. fragilis. As a primary nutrient source for plant growth, sediment type can have differential effects on various submerged macrophyte species and 1:1 mixture treatment could enhance the performance of the communities, increasing the overall biomass, number of shoot nodes and shoot length by 39.03%, 150.13% and 9.94%, respectively, compared to the clay treatment. Thus, measures should be taken to mediate the sediment condition to restore submerged macrophyte communities with different dominant species.     

Herbivory in omnivorous fishes: effect of plant secondary metabolites and prey stoichiometry

1. Many animals that consume freshwater macrophytes are omnivorous (i.e., they include both plant and animal matter in their diet). For invertebrate omnivorous consumers, selection of macrophyte species depends partly on the presence of secondary metabolites in plants, plant carbon/nutrient balances and/or physical structure of plants. However, little is known about the mechanisms influencing consumption of macrophytes in aquatic vertebrates. 2. For two fish species, the omnivorous rudd (Scardinius erythrophthalmus) and herbivorous grass carp (Ctenopharyngodon idella), feeding preferences were determined in three choice experiments. We tested (i) whether the presence of secondary metabolites and macrophyte stoichiometry affects macrophyte species selection by fish, (ii) the importance of macrophyte stoichiometry by manipulating the macrophytes experimentally and (iii) the rate of herbivory when the most palatable macrophyte is offered simultaneously with a common animal prey. 3. In a choice experiment with five species of submerged macrophytes (Callitriche sp., Chara globularis, Elodea nuttallii, Myriophyllum spicatum and Potamogeton pectinatus), Myriophyllum was clearly consumed least by both fishes, which strongly correlated with the highest phenolic concentration of this macrophyte. Additionally, a significant negative relationship was found between consumption and C : N ratio of the five macrophytes. The two most consumed macrophytes also had the lowest dry matter concentration (DMC). 4. In a second choice experiment, the C : N ratio of the least (Myriophyllum) and most (Potamogeton) palatable plants was manipulated by growing the macrophytes under fertilised and unfertilised conditions and subsequently feeding them to rudd. The avoidance of consumption of the chemically defended Myriophyllum by rudd was partly alleviated by the lowered C : N ratio. 5. The third choice experiment showed that both fishes preferred animal prey (the amphipod Gammarus pulex) over the most palatable macrophyte (Potamogeton) when offered simultaneously. The C : N ratio of the amphipods was about half that of the lowest C : N ratio measured in the macrophytes. Consumption by the fishes could not clearly be related to C : P or N : P ratios of prey items in any of the experiments. 6. We conclude that omnivorous fish avoid macrophytes that are chemically defended. However, when these defences are only minor, stoichiometry (C : N ratio) in combination with DMC may be a determining factor for consumption by vertebrate facultative herbivores.     

Summer algal communities and primary productivity in fish ponds

The paper presents data on primary productivity and phytoplankton communities in new experimental ponds which received the following treatments; ammonium nitrate and triplesuperphosphate, triplesuperphosphate, cracked corn (10% crude protein) and Auburn No. 3 fish feed (36% crude protein). Comparative data on algal communities were also obtained from production ponds which received feeds or fertilizers. Basic ecological data on macro-algae are also presented.

1. All nutrient additions to experimental ponds resulted in higher levels of gross photosynthesis and greater concentrations of chlorophyll a than were found in the control treatments. Fertilization with both nitrogen and phosphorus gave the highest values. Chlorophyll a and gross photosynthesis were higher in ponds receiving high protein content feed (Auburn No. 3) than in ponds to which low protein content feed (corn) was applied.
2. Persistent blooms of blue-green algae occurred in ponds receiving nitrogen and phosphorus fertilization. Phosphorus only fertilization produced blooms of blue-greens, but these blooms did not persist as in the ponds to which nitrogen was also added. Control ponds were dominated by green algae. Blue-green algae were seldom abundant in feed treatments.
3. Production ponds had high level of gross photosynthesis and large concentrations of chlorophyll a.
4. Many of the production ponds which received feed applications developed heavy blooms of blue-green algae.
5. The major species of blue-green algae observed in the present study were Oscillatoria sp., Raphidiopsis curvata, Anacystis nidulans, A. aeruginosa, Spirulina sp., and Anabaena circinalis. Heterocyst bearing forms, which can presumably fix nitrogen, were seldom noted in ponds that received continuous additions of nitrogen from fish feeds.
6. Macro-algae are abundant in many fish ponds. Data illustrating the competition of macro-algae with phytoplankton are presented.

     

Limnology and macrophyte vegetation of a deep,clear limestone lake,Loch Borralie

Summary

A bathymetric survey shows Loch Borralie to have a maximum depth of 36 m. It is one of the most transparent lakes in Britain and is unstratified and poor in nutrients and phytoplankton, although the interstitial water of its calcareous muds is rich in nitrogen and phosphorus. Forty-four percent of the loch surface is densely colonised by macrophytes.

The littoral to 2 m bears very sparse Chara aspera var. lacustris and the zone from 2 to 4.5 m is dominated by tall Hippuris vulgaris, Myriophyllum spicatum and four Potamogeton spp., for which Chara and Nitella (principally C. globularis and N. flexilis) form an understorey. Three Potamogeton species persist to 6.5 m in an undulating Chara- Nitella ‘meadow’ which continues, alone, to between 12 and 15 m depth. Loch Borralie is the most deeply and possibly the most extensively colonised lake in the United Kingdom.     

Impacts of a submerged plant (Elodea canadensis) on interactions between roach (Rutilus rutilus) and its invertebrate prey communities in a lake littoral zone

1. Using 5‐m² field enclosures, we examined the effects of Elodea canadensis on zooplankton communities and on the trophic cascade caused by 4–5 year old (approximately 16 cm) roach. We also tested the hypothesis that roach in Elodea beds use variable food resources as their diet, mainly benthic and epiphytic macroinvertebrates, and feed less efficiently on zooplankton. Switching of the prey preference stabilises the zooplankton community and, in turn, also the fluctuation of algal biomass. The factorial design of the experiment included three levels of Elodea (no‐, sparse‐ and dense‐Elodea) and two levels of fish (present and absent). 2. During the 4‐week experiment, the total biomass of euplanktonic zooplankton, especially that of the dominant cladoceran Daphnia longispina, decreased with increase in Elodea density. The Daphnia biomass was also reduced by roach in all the Elodea treatments. Thus, Elodea provided neither a favourable habitat nor a good refuge for Daphnia against predation by roach. 3. The electivity of roach for cladocerans was high in all the Elodea treatments. Roach were able to prey on cladocerans in Elodea beds, even when the abundance and size of these prey animals were low. In addition to cladocerans, the diet of roach consisted of macroinvertebrates and detrital/plant material. Although the biomass of macroinvertebrates increased during the experiment in all Elodea treatments, they were relatively unimportant in roach diets regardless of the density of Elodea beds. 4. Euplanktonic zooplankton species other than Daphnia were not affected by Elodea or fish and the treatments had no effects on the total clearance rate of euplanktonic zooplankton. However, the chlorophyll a concentration increased with fish in all the Elodea treatments, suggesting that fish enhanced algal growth through regeneration of nutrients. Thus, our results did not unequivocally show that Elodea hampered the trophic cascade of fish via lowered predation on grazing zooplankton. 5. In treatments with dense Elodea beds (750 g FW m^?2), chlorophyll a concentration was always low suggesting that phytoplankton production was controlled by Elodea. Apparently, the top‐down control of phytoplankton biomass by zooplankton was facilitated by the macrophytes and operated simultaneously with control of phytoplankton production by Elodea.