Effects of aquatic plant management on stream metabolism and oxygen balance in streams

1. In unshaded, nutrient-rich streams, prolific growth of stream macrophytes often results in flows that over-top the banks and in high primary production and respiration that may result in extreme diel variations in dissolved oxygen. Consequently, water protection authorities commonly remove macrophytes periodically.
2. We investigated the effect of plant removal on stream metabolism and oxygen balance in two Swiss streams with a high macrophyte biomass. We monitored the concentration of dissolved oxygen before and after macrophytes were removed by cutting and dredging, and calculated rates of gross primary production and ecosystem respiration by means of diel oxygen curves.
3. The removal of plants, which had reached a dry biomass of 320–420 g m−2 immediately before plant removal, had a different impact on stream metabolism in the two streams. In the first (plants removed in May), neither primary production nor ecosystem respiration were significantly affected. In the second (plants removed in late July), gross primary production and ecosystem respiration were reduced by about 70%. In this latter stream gross primary production increased in the first 2 weeks after plant removal but never recovered to pre-disturbance levels.
4. The removal of plants coincided with only a moderate increase in nocturnal oxygen concentration (+1 mg L−1). This, and the rapid partial recovery of stream metabolism in the second stream, suggests that an increase in the oxygen concentration after plant cutting is transient in unshaded, nutrient-rich streams.     

Responses of a lake outlet community to light and nutrient manipulation: effects on periphyton and invertebrate biomass and composition

1.  Experimental channels were installed at the outlet of a small Canadian Shield lake to study the role of light and nutrients on the regulation of periphyton and invertebrate biomass and taxonomic composition. Light (93% reduction) and nutrients (four-fold increase of ambient total phosphorus (TP) concentration) were manipulated in a factorial design experiment.
2.  Periphyton chlorophyll a (Chl a ), measured four times during the 12-week experiment, increased due to higher irradiance but was unaffected by phosphorus enrichment. Over the experiment, periphyton biomass was, on average, three times higher in open than in shaded channels.
3.  Algal taxonomic and growth form composition were affected by light and phosphorus enrichment. The proportion of cyanophytes was significantly higher in unenriched, shaded channels (45%) compared to the three other treatments. Single cells and colonial forms were dominant in shaded channels, whereas filamentous and chain-forming algae were prevalent in open channels.
4.  Total invertebrate biomass remained unchanged over time and among treatments. At the beginning of the experiment, all the channels were dominated by Chironomidae and filter feeders (Simuliidae and Hydropsychidae). After 55 days, filter feeders became clearly dominant (60%) in all treatments except in the enriched, open channels where there was a significant shift in the functional group composition toward grazers (snails and oligochaetes). Among filter feeders, Simuliidae increased in shaded channels.     

Structure, biomass, production and depth distribution of periphyton on artificial substratum in shallow lakes with contrasting nutrient concentrations

1. To examine how the vertical distribution of periphytic biomass and primary production in the upper 0–1 m of the water column changes along an inter‐lake eutrophication gradient, artificial substrata (plastic strips) were introduced into the littoral zones of 13 lakes covering a total phosphorus (TP) summer mean range from 11 to 536 μg L^?1. Periphyton was measured in July (after 8 weeks) and September (after 15 weeks) at three water depths (0.1, 0.5 and 0.9 m). 2. Periphyton chlorophyll a concentration and dry weight generally increased with time and the communities became more heterotrophic. Mean periphytic biomass was unimodally related to TP, reaching a peak between 60 and 200 μg L^?1. 3. The proportion of diatoms in the periphyton decreased from July to September. A taxonomic shift occurred from dominance (by biovolume) of diatoms and cyanobacteria at low TP to dominance of chlorophytes at intermediate TP and of diatoms (Epithemia sp.) in the two most TP‐rich lakes. 4. The grazer community in most lakes was dominated by chironomid larvae and the total biomass of grazers increased with periphyton biomass. 5. Community respiration (R), maximum light‐saturated photosynthetic rate (P_max), primary production and the biomass of macrograzers associated with periphyton were more closely related to periphyton biomass than to TP. Biomass‐specific rates of R, P_max and production declined with increasing biomass. 6. Mean net periphyton production (24 h) was positive in most lakes in July and negative in all lakes in September. Net production was not related to the TP gradient in July, but decreased in September with increasing TP. 7. The results indicate that nutrient concentrations alone are poor predictors of the standing biomass and production of periphyton in shallow lakes. However, because periphyton biomass reaches a peak in the range of phosphorus concentration in which alternative states occur in shallow lakes, recolonisation by submerged macrophytes after nutrient reduction may potentially be suppressed by periphyton growth.     

Snail–periphyton interactions in a prairie lacustrine wetland

This study examined the effects of nutrients and macrophytes on snail grazers and periphyton in a prairie wetland food web. Snails (Gyraulus circumstriatus) and periphyton in large enclosures in a lacustrine wetland, Delta Marsh, MB, Canada were subjected to two experimental treatments, nutrient addition (nitrogen, phosphorus) and macrophyte exclusion (using a porous geotextile carpet) during July and August. Snail biomass and periphyton biomass (on both artificial substrata and submerged macrophytes) increased over time in all treatments, representing seasonal growth. Snail biomass was three times higher on macrophytes than on artificial substrata. In response to nutrient addition, snail biomass was significantly elevated over time on macrophytes but not on artificial substrata. Conversely, periphyton biomass was higher on artificial substrata but not on macrophytes in response to nutrient addition. Snail biomass and periphyton biomass on artificial substrata showed no response to macrophyte exclusion. Snail biomass on all substrata was inversely correlated with turbidity, whereas periphyton biomass showed no relationship with turbidity. Timing of nutrient additions to wetlands may influence whether the response occurs primarily in phytoplankton or in periphyton and macrophytes.     

The influence of macrophyte beds on plankton communities and their export from fluvial lakes in the St Lawrence River

1. To determine the influence of macrophyte beds on plankton abundance within fluvial lakes of the St Lawrence River, planktonic components (macrozooplankton, heterotrophic bacteria, and phytoplankton as chlorophyll- a [Chl- a ]) were sampled in Lake St Francis and Lake St Pierre during summer 1998. We tested the hypothesis that the abundance of planktonic components was higher within macrophyte beds in comparison to the more rapidly flushed open water areas of the fluvial lakes.
2. The large cross channel variation in zooplankton biomass was indeed correlated with the presence of dense beds of submerged macrophytes. Total macrozooplankton biomass was nine-fold greater within the beds (mean=180 μg L−1 dry mass) than in either the open water or areas with only sparse vegetation (mean=20 μg L−1 dry mass).
3. Chl- a and heterotrophic bacterial abundance were also higher in the beds, but only slightly so. There was no difference in total phosphorus or dissolved organic carbon concentrations between areas of dense vegetation, sparse vegetation or open water.
4. Macrophyte beds on the margins of the fluvial lakes allow the development of high planktonic abundance relative to the fast flowing central channel. Macrozooplankton biomass was much higher at the outflows of the lakes (∼50 μg L−1 dry mass) in comparison to the inflows (<20 μg L−1 dry mass). The increase is due to the transfer of organisms from submerged macrophyte beds into the central channel in the downstream quarter of the two lakes where the marginal littoral waters enter central channel waters.
5. Along rivers, shallow fluvial lakes appear to act as sources of plankton which is exported downstream during years of extensive littoral macrophyte development.     

Periphyton-macroinvertebrate interactions in light and fish manipulated enclosures in a clear and a turbid shallow lake

In a clear and a turbid freshwater lake the biomasses of phytoplankton, periphytic algae and periphytonassociated macrograzers were followed in enclosures with and without fish (Rutilus rutilus) and four light levels (100%, 55%, 7% and < 1% of incoming light), respectively. Fish and light affected the biomass of primary producers and the benthic grazers in both lakes. The biomass of primary producers was generally higher in the turbid than the clear lake, and in both lakes fish positively affected the biomass, while shading reduced it. Total biomass of benthic grazing invertebrates was higher in the clear than in the turbid lake and the lakes were dominated by snails and chironomids + ostracods, respectively. While light had no effect on the biomass of grazers in the clear lake, snail breeding was delayed in the most shaded enclosures and presence of fish reduced the number of snails and the total biomass of grazers. In the turbid lake ostracod abundance was not influenced by light, but was higher in fish-free enclosures. Density of chironomids correlated positively with periphyton biomass in summer, while fish had no effect. Generally, light-mediated regulation of primary producers was stronger in the turbid than in the clear lake, but the regulation did not nambiguously influence the primary consumers. However, regulation by fish of the benthic grazer community was stronger in the clear than in the turbid lake, and in both lakes strong top-down effects on periphyton were seen. The results indicate that if present-day climate in Denmark in the future is found in coastal areas at higher latitudes, the effect of lower light during winter in such areas will be highest in clear lakes, with typically lower fish biomass and higher invertebrate grazer density.     

Stratification and dynamics of microbial loop communities in Lake Fryxell, Antarctica

1. Lake Fryxell, situated in the McMurdo Dry Valleys, Antarctica, offers the opportunity to study microbial loop processes in the absence of crustacean zooplankton and other higher organisms. This is the first study of Lake Fryxell to provide detailed temporal and vertical variations of microbial loop organisms.
2. Protozoan communities are concentrated around the chemocline (9–10 m) in Lake Fryxell. Phototrophic nanoflagellates (PNAN), heterotrophic nanoflagellates (HNAN) and ciliates formed deep maxima of 14 580, 694 and 58 cells mL−1 respectively. Although abundance and biomass at the chemocline was high, diversity of protozoa was low, Plagiocampa accounting for> 80% of the total ciliate biomass.
3. In the mixolimnion (4.5–8 m), protozoa were less abundant, but more diverse, with 24 ciliate morphotypes being identified within this region of the water column. Inter-annual variability of protozoan biomass and abundance was greater in the mixolimnion than at the chemocline due to more variable nutrient and prey concentrations.
4. Physicochemical gradients in Lake Fryxell were very stable because the perennial ice cover reduced wind driven currents. As a consequence, ciliate species occurred in distinct depth strata, Monodinium being most abundant directly beneath the ice cover, Askenasia having maximum abundance at 8 m and Plagiocampa dominating ciliate biomass at the chemocline. The lack of vertical mixing reduced seasonal successions of PNAN and ciliate species. Three cryptophyte species dominated the PNAN community at all times (>79% of total biomass).     

Fish-mediated indirect effects in a littoral food web

The strength of the direct effect by scraping cladocerans and the indirect effect of nutrient regeneration by filtering herbivorous cladocerans on periphyton growth was investigated in a littoral food web. Ten enclosures were erected in a lake in an area with artificial vegetation. Five enclosures were stocked with juvenile perch ( Perca fluviatilis ) and five lacked fish. In addition, a reference area in the artificial vegetation was sampled. The mesh size of the net surrounding the cages was chosen to allow an inflow of phytoplankton into the cages from the surrounding water. The periphyton and filtering herbivorous cladoceran biomasses were highest in the fish-free treatment. There was no difference in phytoplankton biomass between treatments despite the large difference in filtering herbivorous cladoceran biomass, suggesting that the inflow of phytoplankton into enclosures completely compensated for the loss due to filtering. The reference area and the enclosure with fish showed the same patterns in developments with respect to filtering herbivorous cladocerans and periphyton. Scraping cladoceran biomass was higher in the fish-free treatment resulting in a positive correlation between scraping cladoceran and periphyton biomass. Our results suggest that the positive indirect effect of filtering herbivorous cladoceran nutrient regeneration on periphyton was stronger than the negative direct effect of grazing by scraping cladocerans on periphyton in this semi-open system, and that pelagic production by phytoplankton may foster periphyton growth in the littoral habitat via filtering herbivorous cladocerans. Furthermore, heterogeneity within trophic levels involving primary producers of different growth forms such as phytoplankton and periphyton may enhance the potential for compensatory responses via nutrient recycling.     

Temporal and geographical variation in lake trophic status in the English Lake District: evidence from (sub)fossil diatoms and aquatic macrophytes

1. A sediment core (representing 250–300 years) was taken from each of three lakes of conservation interest and contrasting trophic status in the English Lake District: Wastwater, Bassenthwaite Lake and Esthwaite Water. Lithostratigraphic analyses, radiometric dating and analysis of fossil diatoms were carried out.
2. Transfer functions, based on the diatoms, were used to reconstruct total phosphorus (TP) and, thus, eutrophication at the study lakes. In Wastwater, changes in lake pH were also reconstructed.
3. The lakes were also classified according to their present macrophyte flora, the latter being compared with previous records.
4. The fossil diatoms of Wastwater were continuously dominated by taxa typical of oligotrophic, circumneutral waters, indicating that the lake has not been enriched or acidified in the last 250 years. The aquatic macrophyte flora has probably remained unchanged since before the Industrial Revolution.
5. The diatom assemblages of both Bassenthwaite Lake and Esthwaite Water began to change in the mid-1800s. Further change occurred from the 1960s, at the onset of a recent period of eutrophication. These two lakes have experienced continued nutrient enrichment throughout the 1970s, 80s and 90s, largely associated with increasing phosphorus inputs from sewage effluent. There is no evidence of any recovery in response to recent reductions in external nutrient loads.
6. Only in Esthwaite Water has the change in aquatic macrophytes been pronounced.
7. Palaeolimnological reconstruction is useful in determining background conditions and natural variation in lake ecosystems.     

The response of two submerged macrophytes and periphyton to elevated temperatures in the presence and absence of snails: a microcosm approach

Global warming may affect snail–periphyton–macrophyte relationships in lakes with implications also for water clarity. We conducted a 40-day aquaria experiment to elucidate the response of submerged macrophytes and periphyton on real and artificial plants to elevated temperatures (3°C) under eutrophic conditions, with and without snails present. With snails, the biomass and length of Vallisneria spinulosa leaves increased more at the high temperature, and at both temperatures growth was higher than in absence of snails. The biomass of periphyton on V. spinulosa as well as on artificial plants was higher at the highest temperature in the absence but not in the presence of snails. The biomass of Potamogeton crispus (in a decaying state) declined in all treatments and was not affected by temperature or snails. While total snail biomass did not differ between temperatures, lower abundance of adults (size >1 cm) was observed at the high temperatures. We conclude that the effect of elevated temperature on the snail–periphyton–macrophyte relationship in summer differs among macrophyte species in active growth or senescent species in subtropical lakes and that snails, when abundant, improve the chances of maintaining actively growing macrophytes under eutrophic conditions, and more so in a warmer future with potentially denser growth of periphyton.     

Are epiphytes a significant component of intertidal Zostera noltii beds?

The role of epiphytes in an intertidal Zostera noltii seagrass bed in Marennes-Oléron Bay was assessed in comparison with the other main benthic primary producers (Z. noltii, microphytobenthos) at two bathymetric levels and on a seasonal basis. Assemblage and biomass of epiphytes were studied using scanning electron microscopy (SEM). Z. noltii and its detrital matter followed a typical seasonal pattern: microphytobenthos was present in large quantities throughout the year representing 21% of the total biomass while detrital matter, above-ground parts and below-ground parts accounted for 65, 9 and 5%, respectively. Only two species of epiphytic diatoms, Cocconeis scutellum and Cocconeis placentula, were observed on seagrass leaves. Epiphyte biomass was very low, representing on average less than 0.001% of that of microphytobenthos or leaves. This low epiphyte biomass is linked with the absence of macroalgae and also with the low biovolume of Cocconeis, which formed a monolayer of cells on leaves. This can be explained by the severe conditions of the intertidal and the high leaf turn-over of Z. noltii leaves.     

SPECIES COMPOSITION,BIOMASS, AND NUTRIENT CONTENT OF PERIPHYTON IN THE FLORIDA EVERGLADES1

Periphyton biomass, nutrient dynamics in the biomass, and species composition were studied in two Florida Everglades sloughs from August 1991 to August 1992. Periphyton biomass on macrophytes was strongly season-dependent. Maximum biomasses, 1180, 161, and 59 g dry mass.m^?2 on Eleocharis vivipara, E. cellulosa, and Nymphaea odorata, respectively, occurred in summer and early autumn; winter and spring periphyton biomass was very low (practically not measurable). Periphyton was dominated by blue-green algae (cyanobacteria) during the summer and autumn; diatoms dominated during the winter and spring. Green algae occurred mostly during the summer and autumn, but their growth was sparse and did not contribute significantly to periphyton biomass. Nitrogen-to-phosphorus ratios in the periphyton were very high (59–121:1), suggesting phosphorus limitation of periphyton growth. The periphyton contained large concentrations of calcium (up to 22.3% on dry mass basis) especially in late summer and autumn.     

A 2-year experimental study on nutrient and predator influences on food web constituents in a shallow lake of north-west Spain

1. A 2‐year study was carried out on the roles of nutrients and fish in determining the plankton communities of a shallow lake in north‐west Spain. Outcomes were different each year depending on the initial conditions, especially of macrophyte biomass. In 1998 estimated initial ‘per cent water volume inhabited’ (PVI) by submerged macrophytes was about 35%. Phytoplankton biomass estimated as chlorophyll a was strongly controlled by fish, whereas effects of nutrient enrichment were not significant. In 1999 estimated PVI was 80%, no fish effect was observed on phytoplankton biomass, but nutrients had significant effects. Water temperatures were higher in 1998 than in 1999. 2. In the 1998 experiment, cladoceran populations were controlled by fish and cyanobacteria were the dominant phytoplankton group. There were no differences between effects of low (4 g fresh mass m^?2) and high (20 g fresh mass m^?2) fish density on total zooplankton biomass, but zooplankton biomass was higher in the absence of fish. With the high plant density in 1999, fish failed to control any group of the zooplankton community. 3. Total biovolume of phytoplankton strongly decreased with increased nutrient concentrations in 1998, although chlorophyll a concentrations did not significantly change. At higher nutrient concentrations, flagellate algae became more abundant with likely growth rates that could have overcompensated cladoceran feeding rates. This change in phytoplankton community composition may have been because of increases in the DIN : SRP ratio. Both chlorophyll a concentration and total phytoplankton biovolume increased significantly with nutrients in the 1999 experiment. 4. A strong decline of submerged macrophytes was observed in both years as nutrients increased, resulting in shading by periphyton. This shading effect could account for the plant decline despite lower water turbidity at the very high nutrient levels in 1998.     

The role of microphytobenthos on shallow coastal lagoons: a modelling approach

Ria Formosa is a Region of Restricted Exchange given its limited connection to coastal water circulation. Furthermore, it is subject to several anthropogenic activities that can lead to an increase in nutrients and potentially to eutrophication. Previous studies have shown the importance of the benthic compartment, specifically the microphytobenthos (MPB) in this shallow coastal lagoon. The dCSTT–MPB model [new version of the dynamic Comprehensive Studies Task Team (dCSTT) model] here described couples the benthic and pelagic compartments. Due to the shallowness of the system, the benthic microalgae are one of the most important primary producers of the system. Preliminary results of the model show a large biomass of benthic microalgae, which strongly influences the pelagic chlorophyll concentration by resuspension. However, algae concentrations in the water column are relatively small due to the high flushing rate of the lagoon. The MPB community is mainly supported by nutrients in the pore water. A sensitivity analysis (SA) has revealed that the factors associated with the benthic compartment were the most important and sensitive to changes. Porosity, benthic chlorophyll recycling, loss of MPB due to grazing and the yield of microphytobenthic chlorophyll from nitrogen were some of the most sensitive parameters, as well as the ones associated with decay of particulate organic nitrogen. The development of our dCSTT–MPB model has itself provided insights into benthic function.     

Variation in growth rates of the zebra mussel, Dreissena polymorpha, within Lake Wawasee

1. Field experiments conducted in Lake Wawasee in 1995 and 1996 measured the response of shell growth of Dreissena polymorpha to environmental gradients.
2. Shell growth decreased with initial shell length in four mussel size classes ranging between 8 and 22 mm, and decreased with depth, with mussels in shallow water (<4 m) having growth rates nearly twice those of mussels in deeper water (4–7 m).
3. Growth occurred early in the spring–summer period (May–June) with relatively little shell added later in the summer (July–September), and varied significantly among sites within Lake Wawasee, but not between the 2 years of this study.
4. Rank order of sites was consistent for both years implying that environmental conditions responsible for variation in shell growth were stable within Lake Wawasee.
5. Cage design did not have a significant effect on mussel shell growth nor did the distance of growth cages above the bottom (0.5–0.75 m above the bottom versus directly on the bottom).
6. This study demonstrates the sensitivity of adult mussel growth to subtle variation in environmental conditions occurring within and among lakes, with potential consequences for mussel population dynamics and community structure and function.     

Resistance and resilience of ecosystem metabolism in a flood-prone river system

1. Gross primary production (GPP) and ecosystem respiration (ER) were analysed for 18 months in two reaches of the River Thur, a prealpine river in Switzerland. The upper reach at 655 m above sea level (a.s.l.) is bedrock constrained, has a high slope (0.60%) and a catchment area of 126 km2. The lower reach at 370 m a.s.l. has a more extensive hyporheic zone, a lower slope (0.17%) and a catchment of 1696 km2.
2. In both reaches, temporal patterns of stream metabolism reflected the occurrence of bed-moving spates. Average reductions of GPP and ER by spates were 53 and 24% in the upper reach, and 37 and 14% in the lower reach, respectively. The greater resistance of ER than GPP in both reaches shifted the ecosystem metabolism towards heterotrophy (decrease of the ratio of GPP to ER (P/R)) following spates.
3. Recovery of GPP was significantly faster in the lower reach and exhibited distinct seasonal variation (positive correlation with incident light). The differences in stability (both resistance and resilience) between reaches reflected differences in geomorphic settings and disturbance regime.
4. Stepwise regression analysis was used to explore the potential influence of season, disturbance and prevailing environmental conditions on stream metabolism in each reach. Time since spate plus temperature explained 73 and 86% of variation in ER and GPP, respectively, in the upper reach and 55% of variation in ER in the lower reach. Season plus prevailing environmental conditions explained 67% of variation in GPP in the lower reach.
5. To test how the perception of stability may change with increasing scale of observation, the disturbance regimes of 12 sites were compared with the disturbance regime of the entire Thur catchment. The analysis suggests that stream metabolism at the catchment scale is far more resistant to high flow events than at the reach scale.     

Plant community structure determines primary productivity in shallow,eutrophic lakes

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Top‐down and bottom‐up control of periphyton by benthivorous fish and light supply in two streams

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Bimodality in stable isotope composition facilitates the tracing of carbon transfer from macrophytes to higher trophic levels

Even though the suitability of macrophytes to act as a carbon source to food webs has been questioned by some studies, some others indicate that macrophyte-derived carbon may play an important role in the trophic transfer of organic matter in the food web of shallow lakes. To evaluate the importance of macrophytes to food webs, we collected primary producers—macrophytes and periphyton—and consumers from 19 South American shallow lakes and analyzed their carbon stable isotopes composition (δ¹³C). Despite the diversity of inorganic carbon sources available in our study lakes, the macrophytes’ δ¹³C signatures showed a clear bimodal distribution: ¹³C-depleted and ¹³C-enriched, averaging at ?27.2 and ?13.5‰, respectively. We argue that the use of either CO₂ or HCO₃ ^? by the macrophytes largely caused the bimodal pattern in δ¹³C signals. The contribution of carbon from macrophytes to the lake’s food webs was not straightforward in most of the lakes because the macrophytes’ isotopic composition was quite similar to the isotopic composition of periphyton, phytoplankton, and terrestrial carbon. However, in some lakes where the macrophytes had a distinct isotopic signature, our data suggest that macrophytes can represent an important carbon source to shallow lake food webs.     

The Contribution of Gastropod Biomass in Macrobenthic Communities of a Karstic River

This study evaluated the standing crop biomass of freshwater snails and its spatial distribution according to four substrate types (boulders, cobbles, gravel covered with periphyton and gravel covered with macrophytes). The study site, in the Kupa River of the NW Dinarid area of Croatia, is in a karstic region. Depending on habitat, the gastropod wet biomass varied from 6.4% to 43.8% of the total macroinvertebrate biomass per unit area. The gastropods are the dominant component of biomass on boulder substrates in the upper region of the river, and on gravel substrates covered with macrophytes, in the lower course. It was found that gastropods are also the dominant component of the functional group of scrapers. The average annual gastropod biomass constitutes more than 80% of scrapers at sampling sites in the upper and lower regions of the river.