Stress and disturbance in the phytoplankton community of a shallow,hypertrophic lake

The validity of Connell's intermediate disturbance hypothesis in phytoplankton communities was tested on data from a hypertrophic, shallow lake, Hjarbæk Fjord, Denmark.The present data from Hjarbæk Fjord demonstrate the difficulties in distinguishing stress from disturbance in a phytoplankton community, and show that great changes in the phytoplankton community can take place within few days.A collapse of blue-green algae in late June 1986 caused remineralization of nutrients and resulted in a rapid increase of fast-growing small chlorococcal green algae and phytoplankton species diversity, without any external disturbances acting on the lake. External disturbances in the form of wind action and brackish water intrusion occurred several days after the onset of these events. Carbon depletion and pH 11.0 were severe stress factors on the phytoplankton community. They were induced by calm, warm weather, but eventually acted as a kind of disturbance to the normally well circulated lake.     

In situ measurements confirm the seasonal dominance of benthic algae over phytoplankton in nearshore primary production of a large lake

1. Despite the recognition of its importance, benthic primary production is seldom reported, especially for large lakes. We measured in situ benthic net primary production by monitoring flux in dissolved inorganic carbon (DIC) concentration in benthic incubation chambers, based on continuous measurements of CO_2(aq) flux, alkalinity, and the temperature‐dependent dissociation constants of carbonic acid (K₁ and K₂). This methodology has the advantages of monitoring net primary production directly as change in carbon, maintaining continuous water recirculation, and having sufficient precision to detect change in DIC over short (i.e. 15 min) incubations, even in alkaline waters. 2. Benthic primary production on Cladophora‐dominated rocky substrata in western Lake Ontario was measured biweekly. Maximum biomass‐specific net photosynthetic rates were highest in the spring (2.39 mgC g Dry Mass^?1 h^?1), decreased to negative rates by early summer (?0.76 mgC g DM^?1 h^?1), and exhibited a regrowth in late summer (1.98 mgC g DM^?1 h^?1). 3. A Cladophora growth model (CGM), previously validated to predict Cladophora biomass accrual in Lake Ontario, successfully simulated the seasonality and magnitude of biomass‐specific primary production during the first cohort of Cladophora growth. Averaged over this growing season (May–Aug), mean areal net benthic production at the estimated depth of peak biomass (2 m) was 405 mg C m^?2 d^?1. 4. We measured planktonic primary production in proximity to the benthic study and constructed a depth‐resolved model of planktonic production. Using the CGM, benthic primary production was compared with planktonic primary production for the period May–Aug. Net benthic production from the shoreline to the 12 m contour (1–2 km offshore) equalled planktonic production. Closer to shore, benthic primary production exceeded planktonic primary production. Failure to account for benthic primary production, at least during abundant Cladophora growth, will lead to large underestimates in carbon and nutrient flows in the nearshore zone of this Great Lake.     

Factors influencing taxonomic composition and abundance of macrozoobenthos in extralittoral zone of shallow eutrophic lakes

Zoobenthos is an essential part of shallow lake ecosystems, exerting a considerable impact upon their functioning. We studied 13 eutrophic, shallow, polymictic lakes from Northern Poland to find out which environmental factors influence taxonomic composition, abundance and biodiversity of their zoobenthos. The Canonical Correspondence Analysis allowed to distinguish three lake types: (1) macrophyte-dominated lakes, with high plant cover and well illuminated bottom, inhabited by abundant, diverse benthic taxa; (2) deeper phytoplankton-dominated lakes, with shaded bottom, high sediment oxygen demand (SOD) and rather sparse zobenthos community, dominated by Chironomus and Chaoborus larvae; (3) shallower phytoplankton-dominated lakes, with intermediate amount of light at the bottom and lower SOD values and comparatively diverse zoobenthos, but with lower number of taxa than in the first group. Apart from plant presence, distinguishing between macrophyte-dominated lakes and the other types, the most important variable in the CCA was amount of light reaching the bottom. Probably the impact of light on the bottom fauna was indirect: light stimulated development of macrophytes or phytobenthos (depending on its intensity) and thus improved food and oxygen conditions. Zoobenthos was also affected by oxygen conditions (mainly SOD), presumably by short-time oxygen depletions occurring in the deep phytoplankton-dominated lakes and preventing survival of some benthic taxa.     

Assessments of primary and bacterial production in three large mountain lakes in Alberta,Western Canada

Preliminary studies on production by phytoplankton and bacteria in three large mountain lakes in Alberta, Canada (two in Waterton Lakes National Park and one in Jasper National Park) were concluded mainly through the use of the ¹⁴C technique. The main experiments were conducted in August, 1974, and some were repeated in August, 1975. Net primary production rates varied little from 1974 to 1975, even though there were drastic changes in the phytoplankton composition. Production in the largest lake (max. depth 135 m; mean phytoplankton production 206.5 mgC.m⁻².d⁻¹) was approximately twice that for each of the two smaller lakes (max. depths 19 and 27 m: average phytoplankton production 109 mgC.m⁻².d⁻¹). Bacterial production estimates averaged 3.8 times those for the phytoplankton production, after a proportionalety large error in the dark-uptake technique was subtracted. High production rates in the largest lake are probably due to enrichment. Bacterial production rates are comparable to those in smaller oligotrophic lakes in Europe.     

Modeling the foodweb in coastal areas: a case study of Ringkøbing Fjord,Denmark

This work utilizes the CoastWeb model, a foodweb model for coastal areas that also includes a mass-balance model (CoastMab) for phosphorus and many abiotic/biotic interactions, to study the development in Ringkøbing Fjord, Denmark, from 1985 to 2004. This shallow coastal lagoon has an area of 300 km² and a mean depth of 1.9 m. The water exchange between the lagoon and the North Sea is regulated by a sluice. In 1996 there was a major regime shift in this lagoon with drastic reductions in chlorophyll-a concentrations, significant increases in water clarity (Secchi depth) and major changes in the number and biomass of clams as well as in macrophyte cover. Regime shifts is a “hot” topic in aquatic ecology and in this work the CoastWeb model is used as a tool to understand and quantify the causes behind this regime shift. The CoastWeb model is general and can also be used for other coastal areas. The basic model calculates monthly production values and changes in biomasses of ten functional groups of organisms (phytoplankton, bacterioplankton, herbivorous, and predatory zooplankton, benthic algae, macrophytes, jellyfish, zoobenthos and prey and predatory fish) and in Ringkøbing Fjord, also for clams (Mya arenaria). In spite of its complexity, the model is relatively simple to use, since all driving variables may be readily accessed from maps or monitoring programs. The model includes much abiotic/biotic feedback and it can also be used to address other causes for regime shifts other than the changes in salinity and nutrient inflow, which have caused the changes in Ringkøbing Fjord. The model has previously been tested for more than 20 smaller coastal areas and was shown to predict variations in foodweb characteristics very well. The focus of this paper is on temporal variations within one well-studied coastal area. The paper compares modeled values to empirical data for Ringkøbing Fjord and discusses fundamental ecosystem features such as regime shifts and compensatory effects in a way that is not practically feasible without the use of quantitative models.     

Diatom–environmental relationships and nutrient transfer functions from contrasting shallow and deep limestone lakes in Ontario, Canada

Diatom assemblages were analysed in the surface sediments of 44 alkaline lakes in south-western Ontario, Canada, and combined with a pre-existing 58 south-eastern Ontario lake set: (1) to determine if shallow, polymictic Ontario lakes contain different diatom assemblages from deeper, dimictic lakes, and if so, which environmental variables most influence assemblages; (2) to improve the existing transfer functions; (3) to construct and compare transfer functions separately for dimictic, deep lakes and for polymictic, shallow lakes. Polymictic and dimictic lakes covered a similar nutrient range (spring total phosphorus (TP)=4–54 g/l, spring total nitrogen (TN)=200–927 g/l; n=101) and spring pH levels (7.6–9; n=101). However, polymictic lakes were shallower (median mean depth = 2.9 m vs. 7.3 m in dimictic lakes). Benthic diatoms (average 60% relative abundance) dominated the polymictic lakes, whereas planktonic diatoms (average 60%) dominated dimictic lakes. A Canonical Correspondence Analyses with forward selection (p < 0.05, 999 Monte Carlo permutations) identified TP, alkalinity, watershed to volume ratios and lake depth as the most important measured environmental variables influencing diatom distribution in both polymictic and dimictic lakes. Additionally, pH was identified as an important variable in polymictic lakes, whereas TN was also forward selected in the dimictic lakes. Adding more lakes to the original southern Ontario calibration set improved the TN transfer function (r² _jack=0.42, root mean squared error of prediction (RMSEP)_jack=0.11 [log g TN/l]), although there was a high systematic error in the revised model (r² _residual = 0.48). However, the strongest TP model was derived from the polymictic lakes (r² _boot =0.44, RMSEP_boot=0.20 [log g TP/l]), which was the smallest lake set (n=30) with the lowest number of diatom species. The stronger TP model from the polymictic lakes may be partly due to the relatively low macrophyte cover in our polymictic lakes, which may lead to stronger benthic–pelagic coupling than in lakes with large macrophyte populations. Additionally, our study suggests that the Chrysophyceae cyst:diatom frustule ratio may be useful for indicating trends in TP levels of 35 g/l in alkaline lakes that are dimictic, but is not necessarily indicative of trophic state changes in shallow, polymictic lakes. Our study demonstrates that it may be important to construct separate diatom-based nutrient transfer functions for polymictic and dimictic lakes.     

Abundance and primary production of filamentous green algae Zygogonium ericetorum in an extremely acid (pH 2.9) mining lake and its impact on alkalinity generation

1. In extremely acid mining lakes, benthic filamentous green algae (Zygnemataceae, Chlorophyta) thrive as effective competitors for limited carbon (C). These algae could supply C for microbial‐mediated benthic alkalinity generation. However, biomass, productivity and impact of the acidobiontic filamentous green algae at pH ≤3 have not previously been determined. 2. Periphytic filamentous green algae was mapped by harvesting their biomass from 85 1 × 1 m quadrats in mining lake Grünewalder Lauch. Zygogonium ericetorum colonised water depths between 1.6 and 10.5 m covering 88% of total area. Biomass peaked at 5–6 m depth. Total Zygogonium biomass amounted to 72.2 t dry weight for the whole lake (0.94 km²), which corresponds to 16.1 t C and the accumulation of primary production from 2.2 years. 3. Growth of Zygogonium is moderately N, C and extremely P deficient, and seriously stressed by high rates of Fe deposition during summer. Consequently, net primary production (NPP) of Zygogonium, calculated from measured photosynthesis versus irradiance characteristics and calculated underwater irradiance (0.13 g C m^?2 year^?1) and in situ oxygen measurements (7.8 g C m^?2 year^?1), corresponds to only 0.3% and 18.1% of pelagic NPP. 4. Neither pelagic nor benthic Zygogonium primary production can supply enough C for efficient acidity removal. However, at rates of benthic NPP in summer of 21.4 mg C m^?2 day^?1, Zygogonium contributed 26% of the C equivalents to remove acidity associated with ferric iron, contributing at least seasonally to efficient alkalinity generation.     

Empirical and dynamical models of production and biomass of benthic algae in lakes

This work presents new empirical and dynamical models for benthic algae in lakes. The models were developed within the framework of a more comprehensive lake ecosystem model, LakeWeb, which also accounts for phytoplankton, bacterioplankton, two types of zooplankton (herbivorous and predatory), macrophytes, prey fish and predatory fish. The new dynamic model provides seasonal variations (the calculation time is 1 week). It is meant to account for all factors regulating the production and biomass of benthic algae for lakes in general. This work also presents and uses a new data-base established by us from published sources. Many of the lakes included in this study are situated in the former Soviet Union. They were investigated during the Soviet period and the data and results have up until now been largely unknown in the West. We present empirical models for benthic algae, and show that the biomass of benthic algae in whole lakes can be estimated from the ratio between the lake area above the Secchi depth to the total lake area and the primary production of phytoplankton. We also present several critical tests of the dynamical model. The dynamical and empirical models give corresponding results over a wide limnological domain. We provide algorithms for (1) the production rate of benthic algae (2) the elimination rate (related to the turnover time of benthic algae), (3) the rate of benthic algae consumption by zoobenthos, and (4) the rate of physical erosion of benthic algae. Our results indicate that the production of benthic algae is highly dependent on lake morphometry and sediment character, as well as water clarity, and less dependent on nutrient (phosphorus) concentrations in water and sediments. This work provides new quantitative support to such conclusions and also a useful model for predictions of production and biomass of benthic algae.     

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.     

EFFECTS OF LIGHT AND SUBSTRATE ON THE BENTHIC DIATOMS IN AN OLIGOTROPHIC LAKE: A COMPARISON BETWEEN NATURAL AND ARTIFICIAL SUBSTRATES1

Benthic diatoms form a particularly important community in oligotrophic lakes, but factors influencing their distribution are not well known. This study reports the depth distribution of living motile and total diatoms (living plus dead diatoms) on both natural (from sand to fine organic mud) and artificial substrates in an oligotrophic lake. On artificial substrates, motile diatom densities peaked in abundance (24–30 cells · mm^?2) between 0.6 and 1.9 m depth; on natural sediment surfaces, motile diatoms were generally more numerous and peaked in abundance (925 cells · mm^?2) at 1.3 m depth. Total diatom densities on artificial substrates were highest (1260 valves · mm^?2) at 0.6 m depth, with very low values below 3 m depth; on natural sediment surfaces, total diatom abundances were generally much higher (21600 valves · mm^?2) at 3 m depth and declined gradually with depth. Significant relationships were found between light and diatom densities on the artificial substrate. Ordination analysis indicated that substrate type significantly correlated with the variation of diatom composition on artificial and natural substrates. Our results suggest that in oligotrophic lakes, light influences benthic diatom abundance, whereas substrate type has more influence on benthic diatom composition.