Atmospheric nitrogen-deposition may intensify phosphorus limitation of shallow epilithic periphyton in unproductive lakes

1. We tested whether increasing atmospheric nitrogen (N) deposition along a north–south gradient intensifies epilithic phosphorus (P) limitation in oligotrophic Swedish lakes from the north to the south. We examined the epilithic community at a shallow depth from seven northern and six southern Swedish lakes, and also compared the results with a lake located geographically between the two groups. We determined lake nutrient state, epilithic nutrient ratios and epilithic algal composition, as well as grazer N : P ratios, grazer-epilithon N : P imbalance, and N : P cycling ratios.
2. Epilithic communities appear to be generally more N-limited in the northern lakes and more P-limited in the southern lakes. Lake water total N (Tot-N) and epilithic N : P ratios were lower in northern than in southern lakes and the proportion of N₂-fixing cyanobacteria was higher in northern than in southern lakes.
3. Gastropod grazers had lower N : P imbalances and cycled less N relative to P in northern than in southern lakes.
4. Atmospheric N-deposition showed a strong positive correlation with lake water Tot-N and a much weaker positive correlation with epilithon N : P ratios. Atmospheric N-deposition also correlated negatively with the proportion of N₂-fixing cyanobacteria.
5. There are indications that increased atmospheric N-deposition towards the south might intensify P-limitation of epilithic algae and invertebrate grazers, although more studies are needed to show the strength and generality of our findings.     

Spatial-temporal characteristics of epilithic algae succession on artificial substrata in relation to water quality in Erhai Lake,Yunnan Province,China

In this study, two sites with different nutritional levels at Erhai Lake were selected to perform an epilithic succession experiment in order to better understand the successional dynamic process and associated controlling factors, providing important insights into the role of epilithon in ecological functions and facilitating eutrophic lake recovery planning. The environmental parameters, epilithic biomass and epilithic diversity were qualitatively and quantitatively assessed. Our results showed that faster biomasses peaks (40–50 days colonization) were achieved at Gusheng where nutrient level was higher, comparing with that of Xiahe. A 10-day incubation period is a dividing line from an initial successional stage to the following stage during epilithon succession. The epilithic biomass and biodiversity exhibited distinctly spatial-temporal characteristics, showing higher values at the site being affected by human activities than that being under the natural condition. The spatial characteristic was associated with nitrate enrichment (the most important), and the positively biological responses of the epilithon (higher biomass and diversity) to nitrate enrichment highlight the occurrence of non-point-source pollution on the west bank of Erhai Lake. The temporal characteristic was associated with water temperature (the second important), exhibiting a high biomass accumulation in winter compared with that in summer. This may be caused by other biotic factors not measured in this study, therefore, a detailed investigation regarding the interactions among biotic components in Erhai Lake is essentially needed.     

Seasonal Dynamics of Periphyton in a Large Tropical Lake

Tropical aquatic systems are generally assumed to have little seasonality in productivity patterns. However, this study indicated that there was substantial seasonal variation in epilithic productivity and biomass in tropical Lake Tanganyika, due primarily to seasonal patterns in lake hydrodynamics that influence nutrient availability. Although they support much of the lake’s biological diversity, epilithic algae made a minor contribution to the total energy budget in Lake Tanganyika. A comparison among large, oligotrophic lakes revealed no significant latitudinal trends in periphyton productivity or biomass. However, Lake Tanganyika has relatively low benthic algal biomass and is therefore more efficient at photosynthesis than the temperate lakes. The influence of wave action and consumer density and diversity may be important in moderating productivity of the epilithic community.     

Horizontal variation of biomass and C:N:P ratios of benthic algae in lakes

The horizontal variation of chlorophyll a (chl a) and C:N:P (carbon:nitrogen:phosphorus) ratio was estimated for benthic algal communities attached to living substrates (mussels and macrophytes) and to rocks and stones in three lakes of different trophy. Samples were taken in a nested hierarchical design with replicates separated by several cm, dm, 10 m, and km. The observed horizontal variation of chl a, C:N, and C:P ratios did not differ for horizontal scales, substrates, or lakes. Although the investigated lakes were quite unlike regarding nutrient status, light regime, or morphology, the patchiness was similar in all lakes. Moreover, patchiness was also similar on stones, macrophytes and mussels, although those substrates differed in longevity and surface structure. Similar patchiness regardless of scale, substrate, or sampled lake, implies the possibility of using an optimal sampling design calculated for one lake and substrate also in other lakes and on other substrates.     

Nutrient limitation of epilithic and epixylic biofilms in ten North American streams

1. Nutrient diffusing substrata were used to determine the effect of added inorganic nitrogen (N) and phosphorus (P) on the development of epilithic and epixylic biofilms in 10 North American streams. Four treatments of diffusing substrata were used: Control (agar only), N addition (0.5 m NaNO₃), P addition (0.5 m KH₂PO₄), and N + P combined (0.5 m NaNO₃ + 0.5 m KH₂PO₄). Agar surfaces were covered with glass fibre filters (for epilithon) or discs of untreated white oak wood veneer (for epixylon). 2. We found that if algae showed significant response to nutrient addition, N limitation (either N alone or N with P) was the most frequent response both on GF/F filters and on wood. Despite the low dissolved nutrient concentrations in our study streams, more than a third of the streams did not show any response to N or P addition. In fact, P was never the sole limiting nutrient for algal biofilms in this study. 3. Nutrient addition influenced algal colonisation of inorganic versus organic substrata in different ways. The presence of other biofilm constituents (e.g. fungi or bacteria) may influence whether algal biomass on wood increased in response to nutrient addition. Algae on organic and inorganic substrata responded similarly to nutrient addition in only one stream. 4. Fungal biomass on wood was nutrient limited in six of 10 study streams. N limitation of fungal biomass (with or without secondary P limitation) was most frequent, but P limitation did occur in two streams. 5. Our results show that biomass responses to nutrient addition by the heterotrophic and autotrophic components of the epixylic biofilm were different, though both experienced the same stream nutrient conditions. For algae and fungi growing on wood, limiting nutrients were rarely similar. Only three of nine streams showed the same biomass response to nutrient addition, including two that showed no significant change in biomass despite added nutrients.     

The impact of substrate and lake trophy on the biomass and nutrient status of benthic algae

Algal biomass, C:N:P (carbon:nitrogen:phosphorus) ratios and APA (biomass specific alkaline phosphatase activity) were measured in benthic algal communities on living substrates (mussels and macrophytes) and on rocks and stones (epilithon) in three lakes of different trophy. Benthic algal communities on living substrates had lower C:N:P ratios than epilithon, whereas algal biomass was highest on rocks and stones. Benthic algal biomass increased with the trophic level of a lake despite an increase of C:N:P ratios in the benthic community. The differences in C:N:P ratios and algal biomass between lakes of different trophy were higher on inert substrates than on macrophytes and mussels, probably because algae on living substrates could compensate a poor nutrient supply from lake water with substrate nutrients. However, the substrate was not, as expected, the most important nutrient supply in the oligotrophic lake, but in the eutrophic lake. Therefore, differences between inert and living substrates in a single lake were highest in the eutrophic lake. APA values of the oligotrophic lake were very high especially for benthic algae on stones, indicating an ability of the community to take up nutrients from organic sources. In conclusion, living substrates were an important nutrient source for benthic algae and the importance of this nutrient supply did not decrease with increasing lake trophy.     

Seasonal changes in the biomass,distribution, and patchiness of zooplankton and fish in four lakes in the Sierra Nevada,California

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The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

Changes in community composition, carbon and nitrogen stable isotope signatures and feeding strategy in epilithic aquatic nematodes along a depth gradient

Periphyton is an important component within the littoral zones of lakes, but it is known to vary dramatically on small (cm–m) and large (km) spatial scales, showing differences in composition and abundance. Until relatively recently, changes in periphyton composition along depth gradients have not been studied sufficiently and the response of small meiobenthic invertebrates inhabiting the periphyton to vertically changing environmental conditions such as light are poorly understood. To investigate the changing community composition of epilithic (on stones) nematodes along a depth gradient, we conducted a field study at Lake Erken, Sweden, with the specific objective to investigate whether changes in periphyton composition (algae, bacteria) are reflected in changing nematode feeding types and what the consequences are for nematodes and their resource consumption. We analysed the abundance, species composition and feeding type distribution of epilithic nematodes along 11 depth levels, from 5 to 300?cm water depth. Our study resulted in the first measurements of carbon (¹³C) and nitrogen (¹⁵N) stable isotopes in free living nematodes in lakes. Nematode community composition and feeding type distribution exhibited dramatic changes along the depth gradient. Nematode feeding types changed from a dominance of algae-feeding species in the shallow littoral zone to one of bacteria-feeding species in the deep littoral zone. The ¹³C stable isotope signatures of nematodes and the small (<20?μm) periphyton fraction changed with increasing depth, with nematodes shown to feed on this small periphytic component. Nematodes were identified as primary consumers by means of trophic level calculations based on ¹⁵N stable isotopes.     

NUTRIENT LIMITATION OF BENTHIC ALGAE IN LAKE MICHIGAN: THE ROLE OF SILICA1

In the Laurentian Great Lakes, phytoplankton growth and biomass are secondarily limited by silica (Si), as a result of phosphorus (P) enrichment. Even modest levels of P enrichment can induce secondary Silimitation, which, in turn, promotes a shift from the native diatom phytoplankton flora to chlorophyte and cyanobacteria species. However, very little is known about the nutritional status of benthic populations and their response to nutrient enrichment. Two experiments were performed in the littoral zone of Lake Michigan where nutrients were delivered to in situ benthic algal (episammic and epilithic) assemblages using nutrient‐diffusing substrata. In order to test the hypothesis that benthic algae in Lake Michigan are Si limited, a 2 × 3 factorial experiment was used to deliver all combinations of Si, N, and P to resident assemblages growing on artificial substrata composed of natural (Si rich) versus calcium carbonate (Si poor) sand. A second experiment utilized a serial enrichment to evaluate the role of Si in mediating changes in taxonomic composition. These findings indicate that benthic algae in Lake Michigan exhibit signs of secondary Si limitation, and that their response to enrichment is similar to the phytoplankton. Moreover, natural sand substrata may provide a source of Si to resident benthic algae.     

Scales of horizontal patchiness in chlorophyll a, chemical and physical properties of landfast sea ice in the Gulf of Finland (Baltic Sea)

Horizontal variation of first-year landfast sea ice properties was studied in the Gulf of Finland, the Baltic Sea. Several scales of variation were considered; a number of arrays with core spacings of 0.2, 2 and 20 m were sampled at different stages of the ice season for small-scale patchiness. Spacing between these arrays was from hundreds of meters to kilometers to study mesoscale variability, and once an onshore–offshore 40-km transect was sampled to study regional scale variability. Measured variables included salinity, stable oxygen isotopes (¹⁸O), chlorophyll a (chl-a), nutrients and dissolved organic carbon. On a large scale, a combination of variations in the under-ice water salinity (ice porosity), nutrient supply and the stage of ice development control the build-up of ice algal biomass. At scales of hundreds of meters to kilometers, there was significant variability in several parameters (salinity, chl-a, snow depth and ice thickness). Analyses of the data from the arrays did not show evidence of significant patchiness at scales <20 m for algal biomass. The results imply that the sampling effort in Baltic Sea ice studies should be concentrated on scales of hundreds of meters to kilometers. Using the variations observed in the study area, the estimate for depth-integrated algal biomass in landfast sea ice in the Gulf of Finland (March 2003) is 5.5±4.4 mg chl-a m^–2.     

Direct and indirect effects of UV radiation on benthic communities: epilithic food quality and invertebrate growth in four montane lakes

The direct harmful effects of ultraviolet radiation (UVR) on benthic and planktonic organisms have been well studied in aquatic systems. Less clear, however, is how UVR might affect aquatic communities through its effects on trophic interactions. The focus of this study was twofold: first, to examine the direct effect of UVR on benthic invertebrates and epilithon, the rock-dwelling matrix of algae, bacteria, viruses, fungi and detritus, and second, to examine the indirect effect of UVR-mediated shifts in epilithic food quality on epilithic consumers. Food quality was assessed by measuring carbon to nutrient ratios and the concentration of polyunsaturated fatty acids (PUFA) in the epilithic matrix; the effect of its change on epilithic consumers was measured using a feeding experiment. The study was conducted in four montane lakes, where downwelling UVR can be intense. Of these lakes, the benthic community of only one was strongly affected by UVR. In this lake, exposure to UVR decreased epilithic accrual and invertebrate colonization, and, contrary to our expectations, increased food quality in the shallows through decreased carbon to phosphorus ratios and increased PUFA concentrations. In another of the four study lakes, the feeding experiment showed no significant difference in growth rates between invertebrates fed UVR-exposed and UVR-shielded epilithon, or invertebrates directly exposed to or shielded from UVR. This study demonstrates that although UVR can play an important role in structuring the trophic dynamics of benthic communities, its effects will not be constant across systems, or important in all environments.     

Structure and dynamics of the periphytic algae of Jinyang Lake in Shanxi Province, North China

The structure and dynamics of periphytic algae in Jinyang Lake, the largest man-made lake in Taiyuan, Shanxi, have been studied by examining the periphytic algae samples collected from four sampling stations, during March 2006 to February 2007. In total, 227 species of periphytic algae (including varieties), belonging to three divisions, 27 families, 62 genera, have been identified. Diatoms were the dominant periphytic algae group. Taken the Shannon–Weaver, Margalef and Simpson diversity index together, it showed obvious temporal and spatial dynamics among different seasons and different sampling stations. The highest periphytic community structure evenness appeared at sampling station I in spring, summer and autumn, while at sampling station III in winter. The average biomass of periphytic algae varied from seasons. Our data showed that major factors affecting the periphytic algae in Jinyang Lake was water temperature, nutrient levels as well as human activities. In addition, the main effect caused by thermal power plant has been discussed.     

Inverse relationship of epilithic algae and pelagic phosphorus in unproductive lakes: Roles of N2 fixers and light

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Seasonal variation in biomass and photosynthetic activity of epilithic algae on a rock at the upper littoral area in the north basin of Lake Biwa, Japan

Vertical distributions of biomass and the photosynthetic activity of epilithic algae were observed seasonally with regard to their physical and chemical parameters along a rocky shoreline of the upper littoral area in the north basin of Lake Biwa. Chlorophyll a amounts in epilithic algae showed a marked change with depth from spring to early summer. The algal biomass showed higher values on the lower part of a large rock in early spring, and gradually increased on the upper part as a result of the growth of Spirogyra sp., a filamentous Chlorophyceae. This growth seems to be associated with a rise in water temperature, with the biomass at the upper depths seeming to be affected by a hydraulic disturbance. The standing crop of epilithic algae showed a maximum value in June and declined to a minimum in August because of the corresponding proliferation or disappearance of Spirogyra as well as changes in the water level. The maximum value of the assimilation number calculated from the potential photosynthetic rate of epilithic algae was obtained at the lower depths in June when Spirogyra grew vigorously. Appreciably high assimilation numbers were also observed in summer and fall, although chlorophyll a amounts were low. The present study is the first in a series to clarify the contribution of epilithic algae on rocky areas to productivity throughout the entire littoral ecosystem.     

Production and chlorophyll concentration of epipelic and epilithic algae in fertilized and nonfertilized subarctic lakes

The production and chlorophyll concentration of epipelic and epilithic algae was measured during four years (1972–1975) in two shallow, Swedish subarctic lakes. One lake (Lake Hymenjaure) was fertilized with phosphorus or a combination of phosphorus and nitrogen while the other (Lake Stugsjön) served as a reference. The benthic algae in both lakes were dominated by Cyanophyceae of the same species during the whole investigation. The chlorophyll concentration of epipelic and epilithic algae was 100 and 20 mg·m^–2 respectively and fairly constant during the season. In 1974–1975 there was a significant increase in chlorophyll concentration of the benthic algae in Lake Hymenjaure, probably as a response to the poorer light climate in the lake due to a large phytoplankton development. The annual benthic production was 3.4–7.2 gC·m^–2 and it was not enhanced by the fertilization. Compared to the other primary producers (phytoplankton and macrophytes) the benthic algae constituted 70–83% of the total production in Lake Stugsjön. In Lake Hymenjaure, however, the importance of the benthic algae decreased from 50 to 22% of the total due to the great increase in phytoplankton production induced by the lake fertilization.     

Epilithon δ15N signatures indicate the origins of nitrogen loading and its seasonal dynamics in a volcanic Lake

The intensification of agricultural land use and urbanisation has increased nutrient loads in aquatic ecosystems. Nitrogen loads can alter ecosystem structure and functioning, resulting in increased algal productivity, algal blooms and eutrophication. The principal aim of the present paper is to extend the use of epilithic δ¹⁵N signatures to a lake ecosystem in order to evaluate the potential impact of anthropogenic nitrogen discharges (organic and inorganic) that can also reach coastal waters.Epilithic associations were collected from volcanic rocks in different seasons in shallow water along the entire perimeter of Lake Bracciano and analysed for their nitrogen stable isotope signatures. Furthermore, some stones were moved from an unpolluted site to a polluted one in order to verify the effect on the nitrogen signature of the epilithic association. The epilithon’s δ¹⁵N signatures provided strong evidence of the space-time variability of N inputs. The differing quality of nitrogen loads was reflected in high isotopic variation within the lake, especially at the beginning of summer (1.7‰ ≤ δ¹⁵N ≤ 13.3‰), while in winter, when anthropogenic pressure was lowest, the δ¹⁵N signature variation was less accentuated (3.1‰ ≤ δ¹⁵N ≤ 7.6‰). At all sampling times, spatial variability was found to be related to the various human activities along the lake shore (especially tourism and agriculture), while seasonal variation at all sampling sites was related to the intensity of anthropogenic pressures (higher in summer and lower in winter).Our results showed that epilithic algal associations and the physicochemical properties of the water did not influence the δ¹⁵N signature, which in contrast was strongly related to the site-specific effect of human activities around the lake. Thus, the distribution of δ¹⁵N across space and time can be used to direct nutrient reduction strategies in the region and can assist in monitoring the effectiveness of environmental protection measures.     

Phosphorus and nitrogen limitation of phytoplankton growth in eutrophic Lake Inba, Japan

Lake Inba is one of the most eutrophic lakes in Japan. In this study, field sampling and nutrient enrichment bioassays were conducted to determine the seasonal patterns of nutrient limitation for phytoplankton growth in this lake. Phytoplankton biomass increased significantly with the additions of phosphorus (P) on almost all sampling dates, indicating P limitation of phytoplankton growth from spring to autumn. However, nitrogen (N) limitation was also observed during summer (i.e., 19 August). On 10 August, a typhoon struck Lake Inba. After this event, dissolved inorganic nitrogen (DIN) and phosphorus concentrations increased, probably because of increased river discharge. At the same time, phytoplankton growth in the control treatment became relatively high, with the addition of neither P nor N stimulating the growth. However, 10 days after the typhoon, the phytoplankton growth rate in the control treatment decreased, with only the addition of N having a significant positive effect on phytoplankton growth. N limitation during summer is caused by the low concentrations of DIN, as well as changes in the N:P ratio due to allochthonous nutrient loads. These results indicate that a reduction of both P and N input is necessary to control phytoplankton blooms in Lake Inba.     

Nitrogen dynamics and microbial food web structure during a summer cyanobacterial bloom in a subtropical,shallow, well-mixed,eutrophic lake (Lake Taihu,China)

Nitrogen dynamics and microbial food web structure were characterized in subtropical, eutrophic, large (2,338 km²), shallow (1.9 m mean depth), and polymictic Lake Taihu (China) in Sept–Oct 2002 during a cyanobacterial bloom. Population growth and industrialization are factors in trophic status deterioration in Lake Taihu. Sites for investigation were selected along a transect from the Liangxihe River discharge into Meiliang Bay to the main lake. Water column nitrogen and microbial food web measurements were combined with sediment–water interface incubations to characterize and identify important processes related to system nitrogen dynamics. Results indicate a gradient from strong phosphorus limitation at the river discharge to nitrogen limitation or co-limitation in the main lake. Denitrification in Meiliang Bay may drive main lake nitrogen limitation by removing excess nitrogen before physical transport to the main lake. Five times higher nutrient mineralization rates in the water column versus sediments indicate that sediment nutrient transformations were not as important as water column processes for fueling primary production. However, sediments provide a site for denitrification, which, along with nitrogen fixation and other processes, can determine available nutrient ratios. Dissimilatory nitrate reduction to ammonium (DNRA) was important, relative to denitrification, only at the river discharge site, and nitrogen fixation was observed only in the main lake. Reflecting nitrogen cycling patterns, microbial food web structure shifted from autotrophic (phytoplankton dominated) at the river discharge to heterotrophic (bacteria dominated) in and near the main lake.     

The importance of nitrogen in Pyramid Lake (Nevada,USA), a saline,desert lake

The increase in human development in the downstream portion of the Pyramid Lake drainage basin has resulted in increased nutrient loading to the lake. Since this is a deep, terminal lake, concern over nutrient build up and change in trophic status exists. On the basis of lake chemistry which shows consistently high concentrations of total reactive-P (mean = 55 µg P l^–1) relative to dissolved inorganic-N (DIN) (mean = 15 µg N 1^–1), it has been hypothesized that Pyramid is N-limited. However, no systematic study of nutrient limitation had been undertaken. Nutrient enrichment bioassays conducted throughout an entire year clearly showed that additions of DIN resulted in a 350–600% stimulation of chlorophyll production. Phosphate, when added singly or in combination with DIN, had no effect. This positive response to N-addition was significant at all times of the year except, (1) immediately after complete lake mixing in February when a large pool of hypolimnetic nitrate was injected into the euphotic zone, and (2) during a fall bloom of the nitrogen fixing species Nodularia spumigena. The positive response to N-addition in the bioassay experiments was strong between March and November. However, the seston exhibited only a gradual depletion of nitrogen relative to carbon over this same period. PN:PC ratios suggested no N-deficiency in phytoplankton biomass in February, March and April, moderate N-deficiency in May, June and July and, severe N-deficiency from August until winter turnover. The appearance of nitrogen fixing blue-green algae in September supports the hypothesis of N-limitation in the summer-autumn. In evaluating the nutrient status of a lake, the concepts of nutrient stimulation versus nutrient deficiency versus nutrient limitation must clearly be defined.This paper is dedicated to G. Evelyn Hutchinson who first visited Pyramid Lake in 1933.