Measurements of phosphorus uptake by macrophytes and epiphytes from the LaPlatte River (VT) using 32P in stream microcosms

1. Phosphorus (P) uptake by macrophytes and epiphytes from the LaPlatte River (VT) was examined in the laboratory by adding ³²PO₄‐P to recirculating stream microcosms.
2. Water, plugs of sediment and plants were removed from the river and placed into the microcosms. ³²PO₄‐P was then added either to the water or the sediment, and its incorporation into plants and epiphytes was monitored over 3 days. Uptake was examined at both ambient (5 μg L^–1) and increased (50 μg L^–1) soluble reactive phosphorus (SRP) concentrations. A computer program was developed to fit curves to the radiotracer data and calculate rate constants for the simultaneous transfer of ³²P among compartments.
3. Both macrophytes and epiphytes removed P from the water, but epiphyte uptake of P was more rapid. Phosphate enrichment stimulated P uptake by both macrophytes and epiphytes. Macrophytes also obtained P from the sediment. The relative contribution of P to macrophytes from the water vs. that from the sediment appeared to vary with SRP in the overlying water. Accurate estimates of rates of P uptake from sediments by macrophytes were difficult to obtain however, due to very low and highly variable unit rate constants for P uptake and uncertainty about the magnitude of the phosphate pool available for uptake.
4. SRP concentrations were greater in the overlying water than in the sediment pore water of stream microcosms in the present study. Numerous reports in the literature have suggested that this condition favours uptake by macrophyte stems and leaves rather than by roots.
5. Phosphate uptake from the water by macrophytes in shallow streams may be more common than for macrophytes in lakes.     

Denitrification in river sediments: relationship between process rate and properties of water and sediment

1. A survey was made of denitrification and nitrous oxide (N₂O) production in river sediments at fifty sites in north‐east England during one season in order to investigate the relationship between rates and environmental factors likely to influence these processes. The sites were chosen to represent a wide range of physical and chemical conditions. Denitrification rate and N₂O production were measured within 5 h of sampling using the slurry acetylene blockage technique.
2. Denitrification rate ranged from less than 0.005–260 nmol N g^–1 DW h^–1, tending to increase in a downstream direction. N₂O production ranged from negative values (net consumption) to 13 nmol N₂O‐N g^–1 DW h^–1 and accounted for 0–115% of the N gases produced.
3. Denitrification rate and N₂O concentration in the sediment were correlated positively with nitrate concentration in the water column, water content of the sediment and percentage of fine (< 100 μm) particles in the sediment.
4. The variation in denitrification rate was satisfactorily explained (64% total variance) by a model employing measurements of water nitrate and water content of sediments. No simple or multiple relationship was found for N₂O production.     

Growth of macrophytes and ecosystem consequences in a lowland Danish stream

SUMMARY. 1. The River Suså is a small, nutrient-rich stream situated in an open landscape with clayish subsoil under intensive cultivation. Discharge was variable daily and seasonally due to low groundwater input. Above-ground development of submerged macrophytes was restricted to late May to November, when water velocity and depth were low. Dominant macrophytes were rooted Potamogeton pectinatus and Sparganium emersum and unrooted Cladophora . Biomass development was closet) related to light availability.
2. Growth rates of macrophytes were linearly related to light availability when self-shading was accounted for. Potamogeton pectinatus grew rapidly m May-June, concentrated the biomass at the water-surface during July-August, and then declined exponentially when the shoots became basally senescent. Sparganium emersum had linear, flexible leaves that were continuously replaced from a basal meristem. Sparganium emersum was less susceptible to high water velocities than Potamogeton pectinatus and the biomass declined later and at lower rates during autumn. Sparganium emersum also regrew after culling that left its meristem intact in the sediment. Unrooted Cladophora developed a high biomass during sunny periods and subsequently disappeared at high discharges. The summer biomass of rooted macrophytes was greater in years with high summer discharge, whereas the biomass of Cladophora and of the epiphytic microbial community was lower due to scouring.
3. Submerged macrophytes played a key role in structure and functioning of the ecosystem. They reduced water velocities two to four fold during summer and promoted extensive organic sedimentation. The biomass of benthic diatoms declined parallel to increased macrophyte shading and sedimentation. In addition, submerged macrophytes formed a large substratum for macroinvertebrates and for a microbial community.     

Plant palatability and disturbance level in aquatic habitats: an experimental approach using the snail Lymnaea stagnalis (L.)

1. The palatability of aquatic macrophytes to the snail Lymnaea stagnalis was investigated in the laboratory. Eight species of macrophyte were selected from habitats that differed in either flood disturbance regime or nutrient status.
2. In a non-choice test, single macrophyte species were offered to individual snails. The average amount of plant dry mass consumed per Lymnaea dry mass ranged from 3.6 ± 1.4 (±SE) to 63.6 ± 13.9 mg g^–1 day^–1 across plant species. In a choice test, all eight plant species were presented simultaneously to sets of five snails. The average total consumption was 66.1 ± 3.8 mg g^–1 day^–1 and the maximum average consumption for a single plant was 26.2 ± 3.6 mg g^–1 day^–1.
3. In both tests, the amount consumed by snails differed significantly between the plant species. The species growing in undisturbed habitats were the least consumed. Habitat nutrient status was unrelated to plant palatability.
4. These results suggest that macrophyte species growing in habitats that are rarely disturbed by floods allocate a greater proportion of their resources to resisting herbivory.     

Sources of nutrients to rooted submerged macrophytes growing in a nutrient-rich stream

1. The relative contribution of roots and leaves to nutrient uptake by submerged stream macrophytes was tested in experiments where plants were grown in an outdoor flow-channel system. Water was supplied from a nutrient-rich stream with inorganic nitrogen and phosphorus concentrations typical of Danish streams.
2. Four submerged macrophyte species were tested, Elodea canadensis , Callitriche cophocarpa , Ranunculus aquatilis and Potamogeton crispus, and all species were able to satisfy their demand for mineral nutrients by leaf nutrient uptake alone. This was evident from manipulative experiments showing that removal of the roots had no negative impact on the relative growth rate of the plants. Further, the organic N and P concentrations of the plant tissue was constant with time for the de-rooted plants.
3. Enrichment of water and/or sediment had no effect on the relative growth rate of two species, E. canadensis and C. cophocarpa , indicating that in situ nutrient availability was sufficient to cover the needs for growth. Despite the lack of a response in growth rate, a reduced root/shoot biomass ratio was observed with nutrient enrichment of water and/or sediment, and an increased tissue-P concentration in response to open-water enrichment.
4. The open-water nutrient concentrations of the stream in which the experiments were performed are in the upper part of the range found for Danish farmland streams (the majority of Danish streams). Still, however, the negligible effect of nutrient enrichment on the growth of submerged macrophytes observed suggests that mineral nutrient availability might play a minor role in controlling macrophyte growth in most Danish streams.     

The seasonal biomass and productivity of the submerged macrophytes in a polluted Wisconsin stream

SUMMARY 1. We measured biomass and light/dark bottle productivity of macrophytes in a Wisconsin stream throughout one growing season. Except for a brief period in early spring when a Cladophora glomerata -filamentous algal community was dominant, Potamogeton pectinatus was the dominant macrophyte species in Badfish Creek.
2. Maximum community biomass was 710 g DW m⁻², with a maximum above ground biomass of 620 g DW m⁻² and a maximum below ground biomass of 120 g DW m⁻². Annual productivity was estimated at 1435 g DW m⁻² year⁻¹, with a calculated P/B of 2.01.
3. In situ net production averaged 2.83g C g AFDW⁻¹ h⁻¹ Net positive carbon gain by the P. pectinatus community occurred when water temperatures were above 15°C, and daylength at least 12h. This is correlated to the onset of tuber germination in spring, and the point of maximal biomass decline in autumn.     

Effects of single Ca(OH)2 doses on phosphorus concentration and macrophyte biomass of two boreal eutrophic lakes over 2 years

1. Two hardwater eutrophic lakes of central Alberta were subjected to single doses of Ca(OH)₂ (74 or 107 mg L^–1). The effects of lime treatment on phosphorus (P) precipitation, sediment P release, and macrophyte biomass were assessed for up to 2 years.
2. In both lakes, sediment P release was reduced to 16 and 27%, respectively, of pre-treatment values by the first winter following treatment. However, sediment P release returned to pre-treatment values during the following year.
3. In contrast to these short-term effects, macrophyte biomass decreased by as much as 80% after lime application and remained there for at least 2 years.
4. Our results indicate that a single dose of Ca(OH)₂ may give short-term (< 1 year) control of P and long-term control (> 1 year) of macrophytes in hardwater eutrophic lakes of Alberta.     

Lime treatment and its effects on the chemistry and biota of hardwater eutrophic lakes

1. The main focus of this study was to investigate the effects of single and multiple moderate doses of lime (slaked lime, Ca(OH)₂, and/or calcite, CaCO₃) on eutrophic hardwater lakes. This information would contribute to strategies to manage phytoplankton and macrophyte biomass in eutrophic lakes.
2. Water chemistry and biota were monitored for up to 7 years after initial lime treatment and results were compared with reference systems.
3. Complementary studies investigated the effect of lime on macrophytes in ponds, irrigation canals and microcosm experiments.
4. When water pH was kept within its natural range (≤ 10), single and multiple lime applications to lakes and ponds controlled macrophyte biomass, without negatively affecting invertebrate communities.
5. Single lime treatments at moderate dosages of lakes and ponds resulted in variable and mostly temporary changes in chlorophyll a (chl a ) and phosphorus (P) concentration. Although sediment P release was reduced in single-dose lakes during the first winter following treatment, reductions appeared temporary.
6. Multiple treatments of lakes and ponds were effective at reducing both chl a and P concentrations over longer periods. Mean winter P release rate was also reduced after initial treatment.
7. In laboratory studies, sediment cores were incubated with eight different treatments to assess P release. Redox-sensitive treatments were no more effective at lowering total P concentration in overlying water than some redox-insensitive treatments. Lime reduced total P concentrations, but was not as effective as treatments with alum.
8. The use of lime in managing macrophyte and phytoplankton biomass in shallow, hardwater lakes and ponds may be preferable over other treatments, because lime is economical and non-toxic as long as pH is kept within a natural range.     

Biomass and oxygen dynamics of the epiphyte community in a Danish lowland stream

SUMMARY. 1. We examined the abundance and oxygen metabolism of epiphytic organisms on the dominant macrophyte, Potamogeton pectinatus , in headwaters of the eutrophic River Suså. Microbenthic algae were abundant in the stream during spring and macrophytes during summer.
2. The low macrophyte biomass in spring supported a dense epiphyte cover whereas the high macrophyte biomass during summer had a thin epiphyte cover of 10–100-fold lower abundance per unit area of macrophyte surface. The epiphyte community was dominated by microalgae in spring and by heterotrophs, probably bacteria, during summer. This seasonal shift was shown by pronounced reductions of the chlorophyll a content (from 2–3% to 0.1–0.7% of organic DW), the gross photosynthetic rate (from 20–85 to 3–15 mg O₂, g-¹ organic DW h⁻¹) and the ratio of gross photosynthesis to dark respiration in the epiphyte community (from 5–18 to 1). The reduced contributions of epiphytic microalgae correlated with reduced light availability during summer.
3. Both the density and the photosynthetic activity of epiphytic algae were low on a stream area basis relative to those of microbenthic algae and macrophytes. Rapid variations in water velocity and extensive light attenuation in water and macrophyte stands probably constrained the development of epiphytic algae. The epiphyte community was more important in overall stream respiration, contributing c. 10% to total summer respiration and c. 20% to summer respiration within the predominantly heterotrophic microbial communities on sediments and macrophyte surfaces.     

A continuous-flow system for measuring in vitro oxygen and nitrogen metabolism in separated stream communities

SUMMARY. 1. This paper describes a continuous-flow system that we have used extensively for measurement of oxygen exchange and nitrogen transformations in different communities of lowland streams, i.e. the water, the sediment, and the macrophyte—biofilm community. The system, which is set up on the stream bank, consists of several macrophyte and sediment chambers equipped with a double-flow system that ensures an internal water velocity close to that in the stream and which, by continuously renewing the water, mimics diel fluctuation in stream temperature and water chemistry. Water temperature and dissolved oxygen are measured continuously and the data stored on magnetic tape. Water samples are collected in a refrigerator and analysed using standard chemical procedures.
2. The application of the system is illustrated using results obtained during a 2-day summer experiment in a shallow macrophyte—rich stream. The biological processes in the stream were mainly associated with the macrophyte—biofilm community and the sediment, those in the water being negligible. Oxygen release was confined to the macrophyte—biofilm community, the sediment consuming oxygen both by day and by night. Whole-system gross production and dark respiration occurred at similar rates (6–7g O₂ m⁻² day⁻¹), net balance being about zero. Inorganic nitrogen was consumed both by the sediment and to a greater extent by the macrophytes, the diel average consumption being 1g N m⁻² day⁻¹.
3. The sum of the activity in the macrophyte and sediment chambers corresponded to the overall activity of the stream section as determined by upstream/downstream mass balance. This indicates that the results obtained with the continuous-flow chambers realistically describe the oxygen and the nitrogen metabolism of the stream.     

Influence of invasive macrophytes on channel morphology and hydrology in an open tropical lowland stream, and potential control by riparian shading

1. The catchments of many tropical lowland streams in far north Queensland have been extensively cleared for the cultivation of sugar cane to the extent where very little of the native riparian vegetation remains. Stream channels are often choked by a matrix of introduced pasture grass ( Brachiaria mutica , or para grass) and accumulated sediment from cropland erosion.
2. Detailed transects across Bamboo Creek, a fourth order cane-land stream, revealed an estimated sediment load of 20 000 t km^–1. This has resulted in an estimated 85% reduction in the predicted bankful discharge of the original stream channel. Channel capacity has been reduced from 2.3 times to 0.3 times the predicted Q₅₀ flood discharge of 140 m³ s^–1.
3. Shade cloth treatments of 50% and 90% across the stream were used to mimic the effect of shading by riparian vegetation. Three months of shading resulted in a substantial reduction in the height and standing biomass of para grass in both shade treatments, compared to open plots (0% shade). The most dramatic effect was in the 90% treatment, where a mean reduction of 63% in height and 52% in total biomass was recorded. This was despite high net primary production of para grass in the open plots of 2.8 g dry wt m^–2 day^–1, which resulted in a overall increase of 11% and 28% in plant height and total biomass, respectively.
4. These data suggest that restoration of native riparian vegetation will be an effective long-term means of controlling invasive macrophytes in disturbed cane-land stream channels. Reduction of excessive macrophyte growth and the mobilisation of accumulated sediment are essential to the restoration of natural hydrological and ecological processes.     

Invertebrate grazing during the regenerative phase affects the ultimate structure of macrophyte communities

1. Although the biomass of freshwater macrophytes consumed by invertebrate herbivores (excluding crayfish) is usually low, there is growing evidence that invertebrates do exert a structuring effect on macrophyte communities. To explain this, we postulated that the effect of invertebrates may be concentrated on macrophytes during their regenerative phase.
2. We tested this hypothesis by means of a mesocosm-based experiment, in which we investigated the effects of different densities of pond snails [ Lymnaea stagnalis (L.)] on macrophytes regenerating from the natural sediment propagule bank.
3. After 2 months, a diverse macrophyte community had established in the absence of snails, mainly from sexual propagules. Under moderate snail grazing (4 individuals m⁻²), the ultimate biomass of macrophytes was similar, but its species composition differed dramatically. Only a few unpalatable taxa, such as Ceratophyllum demersum and Nymphaeaceae, persisted. Moreover, the relative success of macrophytes regenerating from vegetative rather than sexual propagules improved. Under higher snail grazing (20 m⁻²), all macrophytes disappeared before the end of the experiment.
4. These results confirm that snails at natural densities can have a strong effect on the ultimate structure of macrophyte communities by selectively consuming some species at a juvenile stage. Therefore, the regenerative phase can be seen as a window of opportunity for invertebrate grazers, which can have a qualitative effect on communities that is disproportionate to the biomass consumed.     

Production potential of aquatic plants in systems mixing floating and submerged macrophytes

1. To investigate whether the areal production of aquatic plant systems can be increased by growing floating and submerged plants together, the floating fern Azolla filiculoides was grown at six densities and two nutrient availabilities in the presence and absence of the submerged macrophyte Elodea canadensis .
2. High nutrient availability did not affect the total areal biomass production, but raised the internal N concentration of Azolla and Elodea by 34% and 152%, respectively, and the internal P concentration by 50% and 378%.
3. High Azolla density reduced Elodea production, whereas Elodea did not affect the production of Azolla . A maximal total production of 4.0 g DW m⁻² day⁻¹ was obtained at an Azolla density at and above full surface cover, when Elodea contribute with less than 10% to the total. This contribution did not raise the combined production of Azolla and Elodea significantly above the production of Azolla grown alone. Thus, maximum production was not enhanced by growing floating and submerged macrophytes together.     

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.     

Effects of CO2 enrichment on mineral accumulation and nitrogen relations in a submersed macrophyte

1. Six- to eight-week greenhouse experiments with independent control of pH and dissolved CO₂ evaluated the potential for CO₂ enrichment to stimulate the accumulation of Al, Fe, P and N in shoots of Vallisneria americana , particularly at pH 5. These minerals were provided only as they occurred in natural lake sediments.
2. The effect of CO₂ enrichment at pH 5 v pH 7.3 on growth and tissue N concentration was also determined.
3. CO₂ enrichment at pH 5 effected 5.5- and 7-fold increases in total shoot accumulation of Al and Fe, respectively. In a two-way factorial experiment, CO₂ enrichment yielded 6- to 11-fold greater total shoot P accumulation in plants grown on less and more fertile sediments, respectively.
4. In a three-way factorial experiment, CO₂ enrichment stimulated Vallisneria growth, especially at pH 5, and resulted in a 31–58% reduction in tissue [N] for different pH × sediment combinations. These are greater reductions than previously reported. It also increased total shoot N accumulation up to 6-fold, and there were significant interactions with pH and sediment source: the CO₂ enrichment effect on shoot N accumulation was greater at pH 5 than at pH 7.3, and it was greater with the more fertile sediment at pH 5.
5. Water chemistry (pH and/or [CO₂]) and sediment fertility thus both indirectly influenced the accumulation of sediment-derived minerals in macrophyte shoots within the water column.     

Effects of CO2 enrichment on mineral accumulation and nitrogen relations in a submersed macrophyte

1. Six- to eight-week greenhouse experiments with independent control of pH and dissolved CO₂ evaluated the potential for CO₂ enrichment to stimulate the accumulation of Al, Fe, P and N in shoots of Vallisneria americana , particularly at pH 5. These minerals were provided only as they occurred in natural lake sediments.
2. The effect of CO₂ enrichment at pH 5 v pH 7.3 on growth and tissue N concentration was also determined.
3. CO₂ enrichment at pH 5 effected 5.5- and 7-fold increases in total shoot accumulation of Al and Fe, respectively. In a two-way factorial experiment, CO₂ enrichment yielded 6- to 11-fold greater total shoot P accumulation in plants grown on less and more fertile sediments, respectively.
4. In a three-way factorial experiment, CO₂ enrichment stimulated Vallisneria growth, especially at pH 5, and resulted in a 31–58% reduction in tissue [N] for different pH × sediment combinations. These are greater reductions than previously reported. It also increased total shoot N accumulation up to 6-fold, and there were significant interactions with pH and sediment source: the CO₂ enrichment effect on shoot N accumulation was greater at pH 5 than at pH 7.3, and it was greater with the more fertile sediment at pH 5.
5. Water chemistry (pH and/or [CO₂]) and sediment fertility thus both indirectly influenced the accumulation of sediment-derived minerals in macrophyte shoots within the water column.     

Factors influencing biomass and nutrient content of the submersed macrophyte Egeria densa Planch. in a pampasic stream

We identified factors influencing biomass and nutrient content in E. densa in an enriched pampean stream of Argentina. Physical (current velocity, temperature), chemical (pH, conductivity, dissolved oxygen, nutrient content in water and sediments), and biological variables (biomass and nutrient content of E. densa, biomass of periphyton and other macrophytes) were estimated at each sampling occasion, and mean monthly values estimated. Biomass and nutrient content in E. densa were correlated with these physical-chemical and biological variables. Biomass was positively correlated with ammonium in stream water (P<0.05) and sediment total nitrogen (P<0.01). Nitrogen showed a positive relationship with ammonium (P<0.01), and a negative one with nitrate and periphyton biomass (P<0.05). Phosphorus was positively correlated with soluble reactive phosphorus (P<0.01). The growth of other macrophyte species in the stream seemed to influence E. densa biomass, probably through competition for light. Current velocity was low and not significantly related with E. densa biomass, however, a flood at the beginning of the study washed the macrophyte stand downstream.     

蓝藻胁迫条件下不同基质类型对苦草和伊乐藻生长的影响

为探讨蓝藻胁迫条件下沉水植物生长与基质营养含量的关系,研究了添加相同含量蓝藻后2 种不同基质(较贫瘠的黄色粘土和较肥沃的黑色淤泥)对苦草和伊乐藻2 种沉水植物生长的影响。结果表明:与伊乐藻相比,苦草的生物量在粘土条件下高于淤泥条件,而基质类型对伊乐藻生物量没有显著影响;苦草的最大叶片长度(用于表征株高)、无性系新株数目及其干物质重和伊乐藻的株高、分株数目和分枝干物质重也是在粘土条件下高于淤泥条件;苦草的最大根长在粘土条件下显著高于淤泥条件(p<0.05)。研究结果表明在蓝藻胁迫的条件下,高营养含量的基质不利于沉水植物的生长,并且对根生型沉水植物苦草的影响要大于假根沉水植物伊乐藻。     

Ceratophyllum demersum hampers phytoplankton development in some small Norwegian lakes over a wide range of phosphorus concentrations and geographical latitude

1. Data on submerged and floating-leafed macrophytes, phytoplankton, nutrients (N, P) and calcium were collected from twenty-four small lakes ( 1 km²) over a wide range of latitudes in Norway. The majority of the investigated lakes were mesotrophic or eutrophic, and most of the lakes were markedly affected by diffuse and point-source runoff from agriculture. According to their macrophyte species composition, the majority of the lakes can be classified as Potamogeton lakes or Chara lakes, or a combination of these.
2. This study is consistent with the 'two alternative stable states' hypothesis. We observed clearwater lakes with dense macrophyte cover over a wider range of total P concentration than has been reported previously: from 30 to more than 700 mg P m^–3. The clearwater state was only observed in lakes with mean depths of less than 1.9 m.
3. Most clear lakes with high cover of submerged vegetation showed indications of N limitation.
4. In this study nearly all the macrophyte-dominated lakes with P concentrations above 30 mg m^–3 had dense stands of Ceratophyllum demersum (L.). This indicates that Ceratophyllum may also play an important role in stabilizing and maintaining a clearwater state at high P concentrations.     

Ceratophyllum demersum hampers phytoplankton development in some small Norwegian lakes over a wide range of phosphorus concentrations and geographical latitude

1. Data on submerged and floating-leafed macrophytes, phytoplankton, nutrients (N, P) and calcium were collected from twenty-four small lakes ( 1 km²) over a wide range of latitudes in Norway. The majority of the investigated lakes were mesotrophic or eutrophic, and most of the lakes were markedly affected by diffuse and point-source runoff from agriculture. According to their macrophyte species composition, the majority of the lakes can be classified as Potamogeton lakes or Chara lakes, or a combination of these.
2. This study is consistent with the 'two alternative stable states' hypothesis. We observed clearwater lakes with dense macrophyte cover over a wider range of total P concentration than has been reported previously: from 30 to more than 700 mg P m^–3. The clearwater state was only observed in lakes with mean depths of less than 1.9 m.
3. Most clear lakes with high cover of submerged vegetation showed indications of N limitation.
4. In this study nearly all the macrophyte-dominated lakes with P concentrations above 30 mg m^–3 had dense stands of Ceratophyllum demersum (L.). This indicates that Ceratophyllum may also play an important role in stabilizing and maintaining a clearwater state at high P concentrations.