Ute Mischke

Cyanobacteria and Cyanotoxins: The Influence of Nitrogen versus Phosphorus

The importance of nitrogen (N) versus phosphorus (P) in explaining total cyanobacterial biovolume, the biovolume of specific cyanobacterial taxa, and the incidence of cyanotoxins was determined for 102 north German lakes, using methods to separate the effects of joint variation in N and P concentration from those of differential variation in N versus P. While the positive relationship between total cyanobacteria biovolume and P concentration disappeared at high P concentrations, cyanobacteria biovolume increased continually with N concentration, indicating potential N limitation in highly P enriched lakes. The biovolumes of all cyanobacterial taxa were higher in lakes with above average joint NP concentrations, although the relative biovolumes of some Nostocales were higher in less enriched lakes. Taxa were found to have diverse responses to differential N versus P concentration, and the differences between taxa were not consistent with the hypothesis that potentially N2-fixing Nostocales taxa would be favoured in low N relative to P conditions. In particular Aphanizomenon gracile and the subtropical invasive species Cylindrospermopsis raciborskii often reached their highest biovolumes in lakes with high nitrogen relative to phosphorus concentration. Concentrations of all cyanotoxin groups increased with increasing TP and TN, congruent with the biovolumes of their likely producers. Microcystin concentration was strongly correlated with the biovolume of Planktothrix agardhii but concentrations of anatoxin, cylindrospermopsin and paralytic shellfish poison were not strongly related to any individual taxa. Cyanobacteria should not be treated as a single group when considering the potential effects of changes in nutrient loading on phytoplankton community structure and neither should the N2-fixing Nostocales. This is of particular importance when considering the occurrence of cyanotoxins, as the two most abundant potentially toxin producing Nostocales in our study were found in lakes with high N relative to P enrichment.

Phytoplankton assemblages and steady state in deep and shallow eutrophic lakes - an approach to differentiate the habitat properties of Oscillatoriales

Ecological conditions and phytoplankton succession in two shallow hypertrophic lakes (Langer See and Melangsee) and a dimictic, eutrophic lake (Scharmützelsee) in a lake chain in Eastern Germany were analyzed from 1999 to 2001 in order to find situations of phytoplankton steady state assemblages and variables controlling the phytoplankton composition according to Reynolds et al. (2002). Long term background data from 1993 to 2001 suggest steady state conditions in shallow lakes, whereas the deep lake exhibited irregular fluctuations between various phytoplankton stages. Since the phytoplankton composition in the shallow lakes was similar in all the 3 years, it was highly predictable. Steady state conditions dominated by different species of Oscillatoriales were detected during the summer period 1999 and 2000 in Langer See and in Melangsee (see Mischke & Nixdorf, this volume). This dominant assemblage found in both lakes (group S1 acc. to Reynolds et al., 2002): Planktothrix agardhii (Gom.) Anagn. et Kom., Limnothrix redekei (Van Goor) Meffert, Pseudanabaena (Lauterb.) is typical in turbid mixed layers with highly light deficient conditions, but it is also regularly dominant in the dimictic lake Scharmützelsee as observed in 1999 and 2001 (Pseudanabaena limnetica (Lemm.) Kom. The Nostocales Cylindrospermopsis raciborskii (Wolz.) Seenayya et Subba Raju and Aphanizomenon gracile (Lemmerm.) Lemmerm. were important in the shallow lakes as well as in lake Scharmützelsee. Nevertheless, the occurrence of filamentous cyanobacteria in the dimictic lake was not regular and an unpredictable change in phytoplankton development was observed in 2000. It is discussed, whether this phenomenon of regular succession in shallow hypertrophic lakes is caused by adaptation to a resilient and an extreme environment or by the pool of species that can live or survive in that environment. This was checked through comparison of the depth of the mixed layer, the mean daily irradiance within this layer and the nutrient resources. Although the nutrient resources in both types of lake are near threshold levels, indicating growth inhibition by dissolved nutrients (DIP, DIN, TIC, DSi), the under water light supply seems to be the key factor favoring the dominance of filamentous cyanobacteria belonging to the functional group S1.

Chrysophytes and chlamydomonads: pioneer colonists in extremely acidic mining lakes (pH <3) in Lusatia (Germany)

Twenty-three extremely acidic (pH between 2.5 and 3.5) mining lakes in Lusatia (Germany) were analysed in order to classify their hydrochemistries and to assist the understanding of phytoplankton colonization of these extreme environments. Neither morphometric nor physical parameters influence phytoplankton composition but determine the extent to which the nutrient supply supports the mass development of Chrysophyceae and Chlorophyceae in certain layers of the water (hypo- or epilimnetic chlorophyll maxima and short mass developments). Conventional trophic classification is not readily applicable to these lakes but a chemical classification on the basis of hydrogen, total iron and acidity is proposed. Species of Ochromonas and Chlamydomonas dominate the phytoplankton in fourteen of the most acid lakes; dinoflagellates occurre additionally in four; a more diverse algal assemblage with diatoms and cryptophytes is found in lakes with moderately acidic (pH 5.7–7.0) or alkaline conditions (pH 7.0–9.4). The lake chemistry is the main determinant for the planktonic composition of the water bodies whereas the trophic state mainly determines the level of algal biomass.

Chlorophyll reference conditions for European lake types used for intercalibration of ecological status

The Water Framework Directive (WFD), requires European Member States to assess the “ecological status” of surface waters. As part of this, many European countries have developed an ecological quality classification scheme for chlorophyll concentrations as a measure of phytoplankton abundance. The assessment of ecological quality must be based on the degree of divergence of a water body from an appropriate baseline, or ‘reference condition’. It is, therefore, necessary to determine chlorophyll reference conditions for all European lake types. This involves examining how chlorophyll concentrations vary by lake type, in the absence of any nutrient pressures from agriculture or wastewater. For this purpose, a dataset of 540 European lakes considered to be in a relatively undisturbed reference condition has been assembled, including data on chlorophyll concentration, altitude, mean depth, alkalinity, humic content, surface area and geographical region. Chlorophyll was found to vary with lake type and geographical region, and to be naturally highest in low-altitude, very shallow, high alkalinity and humic lake types and naturally lowest in clear, deep, low alkalinity lakes. The results suggest that light and mineral availability are important drivers of chlorophyll concentrations in undisturbed lakes. Descriptive statistics (median and percentiles) of chlorophyll concentrations were calculated from populations of lakes in this reference lake dataset and used to derive lake-type specific reference chlorophyll concentrations. These reference conditions can be applied, through a comparison with observed chlorophyll concentrations at a site, in the assessments of ecological status and provide a consistent baseline to adopt for European countries.

Sustaining recreational quality of European lakes: minimizing the health risks from algal blooms through phosphorus control.

1. A safe, clean water supply is critical for sustaining many important ecosystem services provided by freshwaters. The development of cyanobacterial blooms in lakes and reservoirs has a major impact on the provision of these services, particularly limiting their use for recreation and water supply for drinking and spray irrigation. Nutrient enrichment is thought to be the most important pressure responsible for the widespread increase in cyanobacterial blooms in recent decades. Quantifying how nutrients limit cyanobacterial abundance in lakes is, therefore, a key need for setting robust targets for the management of freshwaters. 2. Using a dataset from over 800 European lakes, we highlight the use of quantile regression modelling for understanding the maximum potential capacity of cyanobacteria in relation to total phosphorus (TP) and the use of a range of quantile responses, alongside World Health Organisation (WHO) health alert thresholds for recreational waters, for setting robust phosphorus targets for lake management in relation to water use. 3. The analysis shows that cyanobacteria exhibit a non-linear response to phosphorus with the sharpest increase in cyanobacterial abundance occurring in the TP range from about 20 µg L-1 up to about 100 µg L-1. 4. The likelihood of exceeding the World Health Organisation (WHO) ‘low health alert’ threshold increases from about 5% exceedance at 16 µg L-1 to 40% exceedance at 54 µg L-1. About 50% of the studied lakes remain below this WHO health alert threshold, irrespective of high summer TP concentrations, highlighting the importance of other factors affecting cyanobacteria population growth and loss processes, such as high flushing rate. 5. Synthesis and applications. Developing a more quantitative understanding of the effect of nutrients on cyanobacterial abundance in freshwater lakes provides important knowledge for restoring and sustaining a safe, clean water supply for multiple uses. Our models can be used to set nutrient targets to sustain recreational services and provide different levels of precaution that can be chosen dependent on the importance of the service provision.

Ecological status assessment of European lakes: a comparison of metrics for phytoplankton, macrophytes, benthic invertebrates and fish

Data on phytoplankton, macrophytes, benthic invertebrates and fish from more than 2000 lakes in 22 European countries were used to develop and test metrics for assessing the ecological status of European lakes as required by the Water Framework Directive. The strongest and most sensitive of the 11 metrics responding to eutrophication pressure were phytoplankton chlorophyll a, a taxonomic composition trophic index and a functional traits index, the macrophyte intercalibration taxonomic composition metric and a Nordic lake fish index. Intermediate response was found for a cyanobacterial bloom intensity index (Cyano), the Ellenberg macrophyte index and a multimetric index for benthic invertebrates. The latter also responded to hydromorphological pressure. The metrics provide information on primary and secondary impacts of eutrophication in the pelagic and the littoral zone of lakes. Several of these metrics were used as common metrics in the intercalibration of national assessment systems or have been incorporated directly into the national systems. New biological metrics have been developed to assess hydromorphological pressures, based on aquatic macrophyte responses to water level fluctuations, and on macroinvertebrate responses to morphological modifications of lake shorelines. These metrics thus enable the quantification of biological impacts of hydromorphological pressures in lakes.

Strength and uncertainty of phytoplankton metrics for assessing eutrophication impacts in lakes

Phytoplankton constitutes a diverse array of short-lived organisms which derive their nutrients from the water column of lakes. These features make this community the most direct and earliest indicator of the impacts of changing nutrient conditions on lake ecosystems. It also makes them particularly suitable for measuring the success of restoration measures following reductions in nutrient loads. This paper integrates a large volume of work on a number of measures, or metrics, developed for using phytoplankton to assess the ecological status of European lakes, as required for the Water Framework Directive. It assesses the indicator strength of these metrics, specifically in relation to representing the impacts of eutrophication. It also examines how these measures vary naturally at different locations within a lake, as well as between lakes, and how much variability is associated with different replicate samples, different months within a year and between years. On the basis of this analysis, three of the strongest metrics (chlorophyll-a, phytoplankton trophic index (PTI), and cyanobacterial biovolume) are recommended for use as robust measures for assessing the ecological quality of lakes in relation to nutrient-enrichment pressures and a minimum recommended sampling frequency is provided for these three metrics.

Equilibrium phase conditions in shallow German lakes: How Cyanoprokaryota species establish a steady state phase in late summer

In 2000, a field study in two shallow, polytrophic lakes (Langer See and Melangsee) in eastern Germany revealed an equilibrium state assemblage of Cyanoprokaryota in late summer. During 4 successive weeks in Langer See Planktothrix agardhii (Gom.) Anagn. et Kom., Aphanizomenon gracile (Lemmerm.) Lemmerm. and Pseudanabaena limnetica Lemmerm. were more than 80% of the standing biomass of phytoplankton, and their cumulative biovolume was around 33 mm3 l−1 ((±3.2 SD). In Melangsee, the very small Limnothrix species L. amphigranulata (Van Goor) Meffert was the most common species, accompanied by Pseudanabaena limnetica and Planktothrix agardhii. For 3 weeks, their cumulative biovolume was about 23 mm3 l−1 ((±3.4 SD), which represented 75 – 82% of total biovolume. The dominant species all belong to the functional group S1 defined by Reynolds (1997), except for A. gracile, which we suggest to be included in group SN. In both lakes mean light intensities ranged between 2.2 and 8.3 E m−2 d−1. Overall species spectra were very similar in both lakes, but dominance by Limnothrix and by Planktothrix in the respective lakes is observed repeatedly. The success of these species is discussed in the context of the habitat properties in August/September. Summer mixing events represented no disturbances in the sense of Connell (1978), since they do not interrupt the species dominance. More frequent mixing events and higher concentrations of dissolved nitrogen occurred in Langer See than in the more shallow, but wind protected Melangsee. In Langer See light deficient conditions were intensified by an increasing biomass of P. agardhii, and this species probably benefited from nutrient input by more frequent resuspension. The light deficiency also affected the diversity, expressed as Shannon–Wiener Index (H), which was reduced more in lake Langer See (H = 0.51) than in Melangsee (0.74) during steady state periods. Recognizing the important effects of mixing, we suggest an additional variable to describe habitat properties: the number of full mixing days as a proportion of total days of observation should help to discriminate between shallow habitats with intermittent mixing events, and those with more regularly mixing in summer period.

A phytoplankton trophic index to assess the status of lakes for the Water Framework Directive

Despite improvements in wastewater treatment systems, the impact of anthropogenic nutrient sources remains a key issue for the management of European lakes. The Water Framework Directive (WFD) provides a mechanism through which progress can be made on this issue. The Directive requires a classification of the ecological status of phytoplankton, which includes an assessment of taxonomic composition. In this paper, we present a composition metric, the plankton trophic index, that was developed in the WISER EU FP7 project and demonstrate how it has been used to compare national phytoplankton classification systems in Northern and Central Europe. The metric was derived from summer phytoplankton data summarised by genus from 1,795 lakes, covering 20 European countries. We show that it is significantly related to total phosphorus concentrations, but that it is also sensitive to alkalinity, lake size and climatic variables. Through the use of country-specific reference values for the index, we demonstrate that it is significantly related to other national phytoplankton assessment systems and illustrate for a single European (intercalibration) lake type how it was used to intercalibrate WFD boundaries from different countries.

Defining ecologically relevant water quality targets for lakes in Europe

Summary 1. The implementation of the Water Framework Directive requires EU member states to establish and harmonize ecological status class boundaries for biological quality elements. In this paper, we describe an approach for defining ecological class boundaries that delineates shifts in lake ecosystem functioning and, therefore, provides ecologically meaningful targets for water policy in Europe. 2. We collected an extensive data set of 810 lake-years from nine Central European countries, and we used phytoplankton chlorophyll a, a metric widely used to measure the impact of eutrophication in lakes. Our approach establishes chlorophyll a target values in relation to three significant ecological effects of eutrophication: the decline of aquatic macrophytes, the dominance of potentially harmful cyanobacteria and the major functional switch from a clear water to a turbid state. 3. Ranges of threshold chlorophyll a concentrations are given for the two most common lake types in lowland Central Europe: for moderately deep lakes (mean depth 3–15 m), the greatest ecological shifts occur in the range 10–12 l gL � 1 chlorophyll a, and for shallow lakes (<3 m mean depth), in the range 21–23 l gL � 1 chlorophyll a. 4. Synthesis and applications. Our study provides class boundaries for determining the ecological status of lakes, which have robust ecological consequences for lake functioning and which, therefore, provide strong and objective targets for sustainable water management in Europe. The results have been endorsed by all participant member states and adopted in the European Commission legislation, marking the first attempt in international water policy to move from physico-chemical quality standards to harmonized ecologically based quality targets.