Quantitative responses of lake phytoplankton to eutrophication in Northern Europe

Based on the currently largest available dataset of phytoplankton in lakes in northern Europe, we quantified the responses of three major phytoplankton classes to eutrophication. Responses were quantified by modelling the proportional biovolumes of a given group along the eutrophication gradient, using generalized additive models. Chlorophyll-a (Chl-a) was chosen as a proxy for eutrophication because all classes showed more consistent responses to Chl-a than to total phosphorus. Chrysophytes often dominate in (ultra-) oligotrophic lakes, and showed a clear decrease along the eutrophication gradient. Pennate diatoms were found to be most abundant at moderate eutrophication level (spring-samples). Cyanobacteria often dominate under eutrophic conditions, especially in clearwater lakes at Chl-a levels >10 μg l⁻¹ (late summer samples). We compare the relationships among types of lakes, based on the lake typology of the northern geographic intercalibration group, and among countries sharing common lake types. Significant differences were found especially between humic and clearwater lakes, and between low- and moderately alkaline lakes, but we could not identify significant differences between shallow and deep lakes. Country-specific differences in response curves were especially pronounced between lakes in Norway and Finland, while Swedish lakes showed an intermediate pattern, indicating that country-specific differences reflect large-scale geographic and climatic differences in the study area.     

The relation of biotic and abiotic interactions to eutrophication in Tjeukemeer,The Netherlands

During the summer months of 1974–1985 chlorophyll-a and total P concentration, biomass of Daphnia hyalina, smelt Osmerus eperlanus, bream Abramis brema and pikeperch Stizostedion lucioperca, water temperature and water intake from lake IJsselmeer were monitored in Tjeukemeer. During this period there were manipulations with the bream and pikeperch stocks as a consequence of the termination of a gill-net fishery in 1977, and larval smelt immigrated each year from the large lake IJsselmeer and contributed largely to the yearly smelt recruitment.The correlation matrix of the nine variables mentioned above showed a positive correlation between bream and chlorophyll-a, but surprisingly a negative one between smelt and chlorophyll-a. The latter can only be explained when smelt is the dependent variable. In a multi-linear regression there was a negative effect of temperature, chlorophyll a and pikeperch on smelt and a positive effect of water intake. Daphnia hyalina was negatively influenced by the biomass of smelt and the water intake of lake IJsselmeer. The positive relation of Daphnia hyalina and chlorophyll-a was probably related to better survival chances of D. hyalina in an Oscillatoria-rich environment when smelt is the most important predator. An increasing biomass of bream coincided with higher total-P levels and probably contributed to higher chlorophyll-a levels.     

Unimodal patterns of microbial communities with eutrophication in Mediterranean shallow lakes

The role of the microbial communities in the classical planktonic food web and its response to eutrophication in shallow lakes is still contradictory. Mediterranean shallow lakes with different eutrophication levels were sampled to study the influence of eutrophication on the microbial food web (MFW) and their contribution to the planktonic food web. Percentage of ciliate biomass in the metazooplankton (MZP) showed a U-shaped trend with eutrophication, with maximum at both ends of the chlorophyll-a (Chla) gradient. The MZP to phytoplankton ratio demonstrated a unimodal pattern with minimum values at the two ends of the Chla gradient and maximum values in the Chla range 5-10 μg l^?1. In contrast, the MFW to phytoplankton ratio reached its minimum in the central part of the Chla gradient and maximum values at the extremes of the gradient. These patterns support the hypothesis that the relative importance of bacteria and ciliates is lowest in mesotrophic shallow lakes, and highest in oligotrophic and hypereutrophic systems. These results stress the importance of protozoan in the trophic web, and indicate it is essential to include this group, especially ciliates, when quantifying zooplankton in warm shallow lakes.     

Comparision of historical and recent data on hydrochemistry and phytoplankton in the Rijnland area (The Netherlands)

In two canals and two lakes in the western part of the Netherlands a comparision is made between data on water chemistry and phytoplankton from 1941/1942 with recent data. Orthophosphate in particular, but also inorganic nitrogen, has increased tremendously, especially in the Gouwe canal, where Rhinewater enters the area. The inorganic N/P mass ratio decreased in the last 45 years, indicating that the limiting nutrient has changed from phosphate in 1941 to nitrogen in 1987.The average seston volume, measured by filtering 1001 water through a plankton net (50 µm), has doubled. In the early 1940s the blue-green alga Microcystis aeruginosa regularly formed waterblooms, as it does now. The plankton composition seems to have become impoverished in the last decennia, since several taxa have disappeared. Others are strongly reduced in number. The saprobic index has not changed.Based on the relations between chlorophyll-a and biological oxygen demand (BOD) and between transparency and seston volume, BOD, dry weight and ash free dry weight in 1987 chlorophyll-a and transparency in 1941 are estimated. The average chlorophyll-a concentration in the lakes has doubled or tripled in the last 45 years and the mean transparency in the Gouwe canal declined from 75 to 50 cm. Submerged higher plants at some sites have disappeared in the last decennia. The results can be used to develop ecological objectives for combatting eutrophication in canals and lakes and possibly in the river Rhine.     

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

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

Is reduction of the benthivorous fish an important cause of high transparency following biomanipulation in shallow lakes?

Experimental reduction of the fish stock in two shallow lakes in The Netherlands shows that such a biomanipulation can lead to a substantial increase in transparency, which is caused not only by a decrease in algal biomass, but also by a decrease in resuspended sediment and detritus. A model was developed to describe transparency in relation to chlorophyll-a and inorganic, suspended solids (resuspended sediment). With the use of this model it is shown that more than 50% of the turbidity in these shallow lakes before biomanipulation was determined by the sediment resuspension, mainly caused by benthivorous fish. Another analysis reveals that the concentration of inorganic suspended solids and the biomass of benthivorous fish are positively correlated, and that even in the absence of algae a benthivorous fish biomass of 600 kg ha⁻¹ can reduce the Secchi depth to 0.4 m in shallow lakes. In addition, it is argued that algal biomass is also indirectly reduced by removal of benthivorous fish. Reduction of benthivorous fish is necessary to get macrophytes and macrophytes seem to be necessary to keep the algal biomass low in nutrient-rich shallow lakes. It is concluded that the impact of benthivorous fish on the turbidity can be large, especially in shallow lakes.     

Lake eutrophication associated with geographic location,lake morphology and climate in China

Lake eutrophication is influenced by both anthropogenic and natural factors. Few studies have examined relationships between eutrophication parameters and natural factors at a large spatial scale. This study explored these relationships using data from 103 lakes across China. Eutrophication parameters including total nitrogen (TN), total phosphorus (TP), TN:TP ratio, chemical oxygen demand (COD_Mn), chlorophyll-a (Chl-a), Secchi depth (SD), and trophic state index (TSI) were collected for the period 2001–2005. Sixteen natural factors included three of geographic location, five of lake morphology, and eight of climate variables. Pearson correlation analysis showed that TP and TSI were negatively related to elevation, lake depth, and lake volume, and positively related to longitude. All eutrophication parameters, except for COD_Mn and Chl-a, showed no significant correlation with climate variables. Multiple regression analyses indicated that natural factors together accounted for 13–58% of the variance in eutrophication parameters. When the 103 study lakes were classified into different groups based on longitude and elevation, regression analyses demonstrated that natural factors explained more variance in TN, TP, COD_Mn, Chl-a, and TSI in western lakes than in eastern lakes. Lake depth, volume, elevation, and mean annual precipitation were the main predictors of eutrophication parameters for different lake groups. Although anthropogenic impacts such as point- and nonpoint-source pollution are considered as the main determinants of lake eutrophication, our results suggest that some natural factors that reflect lake buffer capacity to nutrient inputs can also play important roles in explaining the eutrophication status of Chinese lakes.     

Light-harvesting properties of zinc complex of chlorophyll-<Emphasis Type="Italic">a</Emphasis> from <Emphasis Type="Italic">spirulina</Emphasis> in surfactant micellar media

Zn chlorophyll-a was prepareted from Mg chlorophyll-a from spirulina and the optical properties of the ground state and the photoexited state of Zn chlorophyll-a in aqueous surfactant micellar media were studied using UV-vis absorption, fluorescence emission spectra, electrochemical and fluorescence lifetime measurements. In comparison of the UV-vis absorption and fluorescence emission spectra of Zn chlorophyll-a and Mg chlorophyll-a, the blue-shift in the absorption bands and emission peak of Zn chlorophyll-a was observed. The energies of the first excited singlet state of Zn chlorophyll-a was 1.87eV. The first oxidation and reduction potentials of the photoexcited singlet state of Zn chlorophyll-a were −0.67 and 0.60V, respectively. Fluorescence lifetime of Zn chlorophyll-a was 9.0 ns in CTAB micellar solution. The fluorescence lifetime of Zn chlorophyll-a is shorter than that of Mg chlorophyll-a (9.8 ns). The photositability of Zn chlorophyll-a was superior to that of Mg chlorophyll-a in various pH conditions. Published online December 2004     

Phytoplankton biomass and chlorophyll-a in some shallow lakes in central Europe

Based on 388 parallel data of phytoplankton biomass and chlorophyll-a of two shallow lakes and two ponds, the following results were obtained:

1. Relative chlorophyll-a content of phytoplankton biomass varied between 0.08–1.88%; chlorophyll-a concentration showed great differences among lakes.
2. Significant correlations (r = 0.68–0.92) were established between phytoplankton biomass and chlorophyll-a concentration. The regression in the artificial ponds was non-linear.
3. In parallel with the increase of average cell volume, a decrease in relative chlorophyll-a content was observed. A significant correlation (r = + 0.83) between the two variables was found. Relative chlorophyll-a content of phytoplankton is a log-function of average cell volume.

     

Impacts of environmental scenarios on chlorophyll-a in the management of shallow, eutrophic lakes following biomanipulation: An application of a numerical model

Accurate prediction of species changes in lake ecosystems following biomanipulation measures is of paramount importance in view of water quality management. The temporal variation of phytoplankton biomass as chlorophyll-a and transparency as Secchi depth measurements are studied in the Lake Bleiswijkse Zoom, The Netherlands, with a comprehensive structural dynamic model. In the formulation of the biological model, phytoplankton as several species, zooplankton, detritus, planktivores and benthivores, and piscivores are considered to be major contributing state variables for the model. The primary goal of this paper is to describe the possible impacts of several environmental scenarios on chlorophyll-a biomass qualitatively as it would help lake and environmental managers and relevant authorities elucidate the processes of eutrophication and biomanipulation in a broad way. Some of the scenarios that have been studied by this model are: (1) The effect of fixed stoichiometry in terms of internal nitrogen and phosphorus that are tied up within algal cells; (2) the effects of external phosphorus limitation; (3) light limitation and external nitrogen limitation on algal growth; (4) probable consequences that have taken place within the chlorophyll-a biomass due to change in biomasses of various aquatic organisms; and (5) possible changes of chlorophyll-a biomass due to higher temperatures caused by global warming.     

The physiological state of the phytoplankton community of three types of lakes as estimated by its adenylate energy charge

Variations in physiological state and biomass of the phytoplankton community were examined in three different types of lakes, namely Lake Barato, Lake Akan, and Lake Shikaribetu. When the physiological state of the phytoplankton community was estimated by its adenylate energy charge (AEC), low biomass and low physiological state co-appeared gradually in the metalimnion and hypolimnion during stratification. The physiological state of the phytoplankton as estimated by its AEC value did not always correspond to its biomass, estimated by chlorophyll-a and ATP in these three lakes. A high physiological state of the community was usually observed in the euphotic zone, but the low AEC value observed in the euphotic zone of Lake Barato was not identified in the euphotic zones of the other lakes. Thus, the relationship between the value of AEC, and biomass of phytoplankton is a complex variable, which is further discussed in this paper.     

The zooplankton and other limnological features of a humic coastal lagoon (Lagoa Comprida,Macé, R.J.) in Brazil

Comprida lagoon is a shallow lagoon separated from the sea by a sand barrier. It has a brown-coloured freshwater with a high concentration of humic compounds. Its zooplankton community and limnological features were studied from March 1992 to December 1995. The lagoon was characterized by low transparency, acid water and relative constant physical–chemical features, except during sporadic marine entrances. The zooplankton, composed of holoplanktonic and meroplanktonic forms, consisted of 60 taxa. Eleven were permanent elements of the community: e.g. Bosminopsis deitersi, nauplii and copepodites of `Diaptomus' azureus. B. deitersi correlated positively with the lowest pH values and with the highest total dissolved nitrogen. Only four taxa correlated with chlorophyll-a concentration. Pigmentation in `D.' azureus is suggested to be as an energy reserve in a system where phytoplankton is probably light-limited.     

Pigment Diversity in Freshwater Phytoplankton. I. A Comparison of Spectrophotometric and Paper Chromatographic Methods

Spectrophotometric and paper chromatographic analyses of the pigments in the phytoplankton were made from early spring till the end of summer in two small Dutch freshwater lakes. It was found that pigment diversity cannot be adequately estimated by MARGALEF'S pigment ratio nor by polychromatic spectrophotometric methods. The pigments detected with the paper chromatographic method were: chlorophyll-a, chlorophyll-b, chlorophyll-c, phaeophytin-a (traces), phaeophorbide-a, Mg-containing chlorophyll-derivatives, carotene, lutein, violaxanthin, neoxanthin (traces), fucoxanthin, diadinoxanthin, diatoxanthin (traces), peridinin and keto-carotenoids (traces). It is suggested to distinguish between a richness-component and an evenness-component of pigment diversity.     

Larger zooplankton in Danish lakes after cold winters: are winter fish kills of importance?

Winter fish kills can be intense under ice in shallow lakes, and have cascading effects on the food web and ultimately on lake water clarity. In maritime Western Europe, winters are usually mild, but occasional colder periods may also have strong effects on lake fish communities. Global warming may have disproportionate effects by delaying freezing and shortening the period of ice coverage. We studied differences in zooplankton (cladocerans, copepods, and rotifers): phytoplankton biomass, zooplankton community structure, and individual body size among 37 Danish lakes of various depths, chemical characteristics, and trophy, by comparing four winters of different severity (mean winter temperatures ranging from −1.19°C in 1996 to +2.9°C in 1995). We found that crustacean mean body sizes were significantly larger in the summer following a severely cold winter. The zooplankton communities in the summer after a cold winter had a significantly larger proportion of larger-bodied species and taxa. Phytoplankton biomass, expressed as chlorophyll-a (chl-a), was lower and zooplankton herbivory (chl-a:TP index), higher, in the summer after the severely cold winter of 1995/1996. All these effects were stronger in shallow lakes than in deep lakes. Changes in zooplankton during summer 1996, compared with other years, were likely caused by fish kills under ice during the preceding severe winter of 1995–1996. Fish kills due to under ice oxygen depletion would be expected to occur earlier and be more complete in the shorter water columns of shallow lakes. With climate change, severe winters are predicted to become less frequent and the winters to be milder and shorter. In general, this is likely to lead to higher winter survival of fish, lower zooplankton grazing of phytoplankton the following summer and more turbid waters, particularly in shallow eutrophic lakes.     

The trophic state ‘chain of relationships’ in ponds: does size matter?

In contrast to shallow lakes, factors affecting trophic state in ponds are not well described, and may include unique effects related to pond size. We examined the interdependence of ambient N and P, phytoplankton biomass, light penetration and stratification in 13 ponds of varying area and mean depth in Chester County, PA (USA) during March, May and July of 2002. Seven of the ponds had primarily residential watersheds, and six were farm ponds. The ponds were all eutrophic or hypereutrophic based on Carlsons Trophic State Index, but varied widely in nutrient content (from 20 to 179 g total P/L and from 683 to 3895 g total N/L) and size (surface areas from 0.1 to 1.7 ha, mean depths from 0.6 to 1.8 m). Although total N (TN) typically declined while total P (TP) rose from March to July, the ratio TN/TP remained sufficiently high that algal growth was probably limited by P during the study period. Phytoplankton biomass (as chlorophyll-a) was positively correlated with TP and negatively correlated with Secchi depth on all three sampling dates. Despite their shallow nature, the ponds were frequently stratified with near-anoxic hypolimnia later in the study period; mixing depth was greater in ponds with greater light penetration. Residual variation in chlorophyll-a unexplained by TP, and in Secchi depth unexplained by chlorophyll-a, was related to either mean depth or surface area, suggesting that trophic state in ponds may be controlled differently than in much larger, shallow lakes.     

Phytoplankton productivity and chlorophyll-A concentration of Oguta Lake in Southeastern Nigeria

The phytoplankton productivity and chlorphyll-a concentration of Oguta Lake, the largest natural lake in south-eastern Nigeria, are presented (Dec. 1983. Nov. 1984). The gross productivity ranged from 1.3 to 3.77 g C.m^–2.day^–1 for the water column, dropped during the period of heavy rainfall and varied with depth. The chlorophyll-a concentration had monthly means ranging from 2.31 to 4.00 mg.m^–3, with a drop during the rains, but little depth variation. Both productivity and chlorophyll-a showed non-significant correlation with the physico-chemical features of the water. The values of the biological parameters showed the lake as mesotrophic. The values are compared with those of other African lakes.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

The Saline Lakes of Saskatchewan IV. Primary Production by Phytoplankton in Selected Saline Ecosystems

Primary production by phytoplankton, efficiency of photosynthesis, and chlorophyll-a concentrations were determined for seven saline lakes that varied widely in ionic concentration and composition. The investigations were done during the summer months of 1972 and 1973. Productivity ranged from 0.001 to 11.135 g C m⁻³ day⁻¹ and 0.053 to 7.968 g C m⁻² day⁻¹. Highest productivities were measured in two lakes that supported blooms of Aphanizomenon flos-aquae and Nodularia spumigena, respectively. Species of Cyanophyceae, Bacillariophyceae and Chlorophyceae dominated the phytoplankton of the study lakes. Active chlorophyll-a ranged from 0.01 to 116 mg m⁻³. Integral photosynthetic efficiency estimates were <1% except during phytoplankton blooms when they were considerably higher. The overall range of 0.03 to 3.8% is concordant with estimates for other lacustrine ecosystems. The extinction of light caused by photo-synthetic processes, or in situ efficiency, was <1% in the trophogenic zone for most lakes but, it was considerably higher during blooms. In situ efficiencies invariably increased with depth in ail lakes.     

P-load,phytoplankton, zooplankton and fish stock in Loosdrecht Lake and Tjeukemeer: confounding effects of predation and food availability

The fish community in the Loosdrecht lakes is dominated by bream, pikeperch and smelt and is characteristic of shallow eutrophic lakes in The Netherlands. The biomasses of the respective fish species amount to ca. 250, 25 and 10 kg ha^–1 and correspond to those in Tjeukemeer, another lake in The Netherlands. The average size of bream, however, is much smaller in the Loosdrecht lakes as a consequence of poorer feeding conditions. The zooplankton community in the Loosdrecht lakes is predominantly composed of relatively small species such as Daphnia cucullata, Bosmina coregoni and cyclopoid copepods, whereas in Tjeukemeer, Daphnia hyalina is permanently present in relatively high densities and the other species show a larger mean length. In the Loosdrecht lakes, the absence of D. hyalina and the smaller sizes of the other zooplankton species could be the consequence of a higher predation pressure, in combination with unfavourable feeding conditions for the zooplankton including the low density of green algae and the high density of filamentous cyanobacteria. A biomanipulation experiment in Lake Breukeleveen, one of the Loosdrecht lakes, indicated that feeding conditions were too unfavourable for large zooplankton to develop in spring, when the reduced fish biomass was not yet supplemented by natural recruitment and immigration.