Experimental study of the effect of nutrients on phytoplankton of the shallow highly eutrophic Lake Nero

Experimental studies of the effect of phosphorus, nitrogen, and silicon on the phytoplankton of the shallow highly eutrophic Lake Nero have been carried out. It is shown that the enrichment of lake water with nitrogen leads to an increased concentration of chlorophyll a and the growth of the biomass of dominant algae species. Phosphorus limits the abundance of a certain species of algae; the limiting role of silicon in the development of spring phytoplankton has not been experimentaly confirmed.     

Long-term change in water transparency before and after the loss of eelgrass beds in an estuarine lagoon,Lake Nakaumi,Japan

Long-term changes in Secchi disk transparency in Lake Nakaumi, Japan, from 1932 to the present, which includes the periods before and after the loss of eelgrass (Zostera marina L.) beds, were compiled from previous reports. During the first (July 1932–February 1934) and the second (January 1949–November 1950) periods, the mean transparency was greater than 3 m at all observed stations. Data during the third period (July 1954–March 1956) are only available for the station at the center of the lake. The mean transparency during the third period was significantly lower than that of the second period but higher than the mean during the fourth period. This observation suggests that the decrease of transparency occurred during the third period. Because the decline of eelgrass beds in Lake Nakaumi occurred in the mid-1950s, the decrease of transparency most likely resulted from the shift in primary producers from submerged macrophytes to phytoplankton. Although the maximum transparency sometimes exceeded 3 m, mean transparencies during the fourth period (May 1975–December 2003) were less than 2 m at all stations, significantly lower than those during the first and second periods. The shift in the chief primary producer, from benthic macrophytes to phytoplankton, caused a subsequent shift in secondary producers. The opportunistic filter-feeding bivalve Musculista senhousia, regarded as a biofouling species of local fisheries, increased in Lake Nakaumi. The long-term monitoring data of transparency suggested that restoration of submerged aquatic vegetation may be indispensable for the remediation of the lake environment in this shallow eutrophic lagoon.     

Composition changes of phototrophic microbial communities along the salinity gradient in the solar saltern evaporation ponds of Eilat,Israel

There are several conflicting hypothesis that deal with the influence of flooding in the natural river–floodplain systems. According to the Flood Pulse Concept, the flood pulses are not considered to be a disturbance, while some recent studies have proven that floods can be a disturbance factor of phytoplankton development. In order to test whether flooding acts as a disturbance factor in the shallow Danubian floodplain lake (Lake Sakadaš), phytoplankton dynamics was investigated during two different hydrological years—extremely dry (2003) without flooding and usually flooded (2004). A total of 18 phytoplankton functional groups were established. The sequence of phytoplankton seasonality can be summarized P/D → E (W1, W2) → C/P (only in potamophase) → S2/H1/S_N/S1 → W1/W2 → P/D. The canonical correspondence analysis (CCA) demonstrated that the water level was a significant environmental variable in 2004. Due to the higher total biomass of Bacillariophyceae established under potamophase conditions, floodings in the early spring seem to be a stimulating factor for phytoplankton development. On the other hand, the flood pulses in May and June had dilution effects on nutrients, so that a significantly lower phytoplankton biomass was established indicating that flooding pulses can be regarded as a disturbance event. Such conditions supported diatom development (D, P, C species) and prolonged its dominance in the total phytoplankton biomass. A long-lasting Cyanoprokaryota bloom (various filamentous species—S1, S2, S_N and H1 representatives) with very high biomass characterized the limnophase (dry conditions) in summer and autumn of both years. In-lake variables (lake morphology, internal loadings of nutrients from sediments, light conditions) seem to be important for the appearance of Cyanoprokaryota bloom. The equilibrium phase was found during the Cyanoprokaryota bloom only in the extremely dry year. This study showed that depending on the time scale occurrence, flood pulses can be a stimulating or a disturbance factor for phytoplankton development in Lake Sakadaš. Handling editor: J. Padisak     

Distribution and special traits of naturalization of new and rare zooplankton species in waterbodies of the Upper Volga basin

More than ten new zooplankton species were registered in reservoirs of the Upper Volga during 2002–2005. Since 2003, the cladoceran Diaphanosoma orghidani Negrea, had intensively settled in different biotopes of Ivankovo, Uglich, Rybinsk, and Sheksna reservoirs and is the most abundant (16 000 ind./m³) in the latter one. The rotifer Asplanchna henrietta Langh. was found for the first time in Rybinsk Reservoir in 1985, and by 2004, this species had also occurred in Ivankovo, Uglich reservoirs, Lake Nero, and some small waterbodies of the basin. In Lake Nero its density reaches especially high values (>1 million ind./m³). Causes, mechanisms, and ecological consequences of the expansion of new zooplankton species are discussed.     

Ecology of Dictyosphaerium pulchellum Wood (Chlorophyta, Chlorococcales) in a shallow, acid, forest lake

This study relates to the ecology of Dictyosphaerium pulchellum Wood in Delamere Lake in Cheshire, UK. Dictyosphaerium pulchellum is a cosmopolitan, green colonial phytoplankton species that occasionally forms dense, monospecific populations in lakes. Delamere Lake is a small, shallow, acid lake (mean pH, 4.5) with very high phytoplankton biomass (annual mean chlorophyll a, 290 μg l⁻¹) and devoid of any significant cladoceran population, the efficient grazers of phytoplankton. A predominantly unicellular form of D. pulchellum was the dominant species in Lake Delamere, and it comprised on average ca. 80% (maximum >99%) of the lake phytoplankton biovolume. Laboratory and lake experiments were conducted on this species showed that its pH tolerance varied between 2.4 and 10.7, and its optimum tolerance range between 3.3 and 8.5 depending on other environmental variables. Low pH was not responsible for the unicellular habit of this alga, but a very high nutrient regime could be an important factor. Bioassays revealed that in Delamere Lake this species was limited by nitrogen, but nitrogen did not hamper high growth in the lake. Dictyosphaerium pulchellum can persist at low light levels, tolerate CO₂-deficiency and can grow in polyhumic water with water colour around 300 mg Pt l⁻¹, but probably not in darker waters. The dominance of D. pulchellum in Delamere Lake is apparently due to a combination of several factors: its ability to tolerate both low pH and high turbidity, exploit high nutrient conditions, absence of effective grazing pressure by zooplankton and being a superior competitor.     

Role of atelomixis in replacement of phytoplankton assemblages in Dom Helvécio Lake, South-East Brazil

Scale and frequency of changes in a lake’s physical structure, light dynamics, and availability of nutrients are closely related to phytoplankton ecology. Since phytoplankton assemblages were first described, phytoplankton ecologists concluded that these assemblages provide insight into phytoplankton responses to environmental changes. Objectives of this study were to investigate ecology of phytoplankton during a complete hydrological cycle in the deepest natural lake in Brazil, Dom Helvécio, and to sort species into the list of assemblages, checking its accordance with environmental changes in a tropical system within the middle Rio Doce Lake district, South-East Brazil. Canonical Correspondence Analysis, t-test, Mann–Whitney U-test, and Kruskal–Wallis test were used to analyze climatological, environmental, and plankton data, which were obtained monthly in 2002. A new phytoplankton assemblage, N_A (atelomixis-dependent desmids), is suggested because atelomixis (robust movement of water occurring once a day) contributed to replacement of species in Dom Helvécio Lake. Stability of stratification, water chemistry, and composition of phytoplankton assemblages characterized two periods. The first period occurred in six rainy months (Jan–Mar and Oct–Dec) when the lake was stratified and phytoplankton was dominated by two assemblages: N_A and F. The second period occurred in six dry months (Apr–Sep) when the lake was nonstratified and phytoplankton was dominated by four assemblages: S2, X1, A, and L_O. Results suggest that phytoplankton in Dom Helvécio Lake was shaped by seasonal and daily changes of water temperature, even with its lower amplitude of variation within 2002 (El Niño year). These changes promoted water column stratification or mixing, reduced light, and increased nutrient availability. Temperature, therefore, is similarly important to phytoplankton ecology in tropical regions as it is in temperate ones. Sorting phytoplankton species into assemblages matched well with environmental changes and periods identified so it is also suggested that this can be further used as an appropriate tool to manage water quality when evaluating tropical lakes.     

Size fractionated phytoplankton production in two humic shallow lakes with contrasting coverage of free floating plants

We conducted a 1-year survey in two humic shallow lakes from the floodplain of the Lower Paraná River, Laguna Grande Lake (LGL) and a relictual oxbow lake (ROL). We aimed to test two hypotheses: (1) the efficiency in light use of picoplankton (0.2–3 μm) is greater as light restriction increases and (2) the contribution of picoplankton to the total productivity is higher when the total photosynthetic biomass is lower. We performed P–E curves for picoplankton and nano- and microplankton (>3 μm) using the ¹⁴C assimilation technique. The light environments of the water bodies differed mainly owing to the development of free floating plants on the surface of the ROL and the dominance of phytoplankton in LGL. Primary productivity patterns in LGL were seasonality driven whilst in the ROL they were related to the coverage of floating macrophytes, which promoted light limitation and a lower productivity. In LGL, nano- and microplankton were in general more productive and the relative contribution of picoplankton to the total phytoplankton production decreased with the increase in total photosynthetic biomass. Hence, our study extends previously observed patterns to subtropical shallow lakes, where seasonality and free floating plants may influence the dynamics of phytoplankton production.     

A five-year study of autotrophic winter picoplankton in Lake Balaton,Hungary

Picoeukaryotes dominate the phytoplankton of Lake Balaton—the largest shallow lake in Central Europe—in the winter period. We examined the annual dynamics of picoplankton abundance and composition in the lake in order to establish if the picoeukaryotes merely survive the harsher winter conditions or they are able to grow in the ice-covered lake when the entire phytoplankton is limited by low light and temperature. Lake Balaton has an annual temperature range of 1–29°C, and it is usually frozen between December and February for 30–60 days. In the spring-autumn period phycocyanin and phycoerythrin rich Cyanobacteria are the dominant picoplankters, and picoeukaryotes are negligible. Our five-year study shows the presence of three types of picophytoplankton assemblages in Lake Balaton: (1) Phycoerythrin-rich Cyanobacteria—the dominant summer picoplankters in the mesotrophic lake area; (2) Phycocyanin-rich Cyanobacteria—the most abundant summer picoplankters in the eutrophic lake area and; (3) Picoeukaryotes—the dominant winter picoplankters in the whole lake. The observed winter abundance of picoeukaryotes was high (up to 3 × 10⁵ cells ml⁻¹), their highest biomass (520 μg l⁻¹) exceeded the maximum summer biomass of picocyanobacteria (500 μg l⁻¹). Our results indicate that the winter predominance of picoeukaryotes is a regular phenomenon in Lake Balaton, irrespective of the absence or presence of the ice cover. Picoeukaryotes are able to grow at as low as 1–2°C water temperature, while the total phytoplankton biomass show the lowest annual values in the winter period. In agreement with earlier findings, the contribution of picocyanobacteria to the total phytoplankton biomass in Lake Balaton is inversely related to the total phytoplankton biomass, whereas no such relationship was observable in the case of picoeukaryotes.     

Reconstruction of the Time Series of the Underwater Light Climate in a Shallow Turbid Lake

Lake ülemiste is a shallow, eutrophic lake which has served the city of Tallinn as a water reservoir for many centuries. Its light climate was studied by combining a routinely measured data set with a modelling approach. For 26 years (1978–2004), data was collected on such optically active substances (OAS) and water parameters as water colour, turbidity and phytoplankton biomass. Simple modelling enabled the quantification of long-term time-series data and the subsequent calculation of the diffuse attenuation coefficient, euphotic depth and average light of the mixed layer. Several changes in the hydrological cycle have taken place during the period under study, among which are an increase in the water level of about 0.5 m and a decrease in the external water load from 108 million m³ year⁻¹ to about 25 million m³ year⁻¹. At the same time euphotic depth has shown a distinct trend towards increasing since the early 1990s. The euphotic depth also showed an increase (from 1.1 to 1.4 m) due to an improvement in underwater light conditions – mainly in the spring (April and May) and autumn (October and November) because of the lower amount of dissolved organic matter in the lake. The average light availability in the mixed layer has increased, but this has not affected the phytoplankton biomass as the latter is not light-limited during the summer period.     

Cryptic speciation in the cosmopolitan <Emphasis Type="Italic">Epiphanes senta</Emphasis> complex (Monogononta,Rotifera) with the description of new species

The phytoplankton species composition and seasonal succession were examined in Lake Kastoria during the period November 1998–October 1999. A total of 67 species and 19 functional groups were identified. Only 4 out of the 67 species, all Cyanobacteria, were dominant (Limnothrix redekei, Microcystis aeruginosa, Cylindrospermopsis raciborskii and Aphanizomenon gracile). Diatoms were rare, not only in terms of species number, but also in terms of biomass (contributing <5% to the total phytoplankton biomass) in relation to the rather low silicon concentrations throughout the year. The functional groups S1, S_N, M and H1 were found dominant in the lake. The species A. gracile (functional group H1) behaved like the species Cylindrospermopsis raciborskii (functional group S_N) which is tolerant to mixing and poor light conditions. The phytoplankton seasonal succession showed similar patterns in all six sampling stations, both at the surface and the bottom water layer, with minor differences during Microcystis aeruginosa dominance. Two steady-state phases were identified within a year lasting for 4 months under relatively stable physical conditions. In these steady-states, the Limnothrix redekei persistent dominance under low light availability and low inorganic nitrogen has been explained by its specific ability such as buoyancy regulation to exploit resources in the water column. Moreover, high population densities over the winter and before the development of daphnids may contribute to the steady-state dominance of Limnothrix. Different niches separated vertically in the water column is one of the explanations for the Limnothrix–Microcystis steady-state when a replacement between the two species was observed in different water layers and areas of the lake. Long lasting steady-states of Cyanobacteria observed in Lake Kastoria and in other Mediterranean and tropical freshwaters may indicate influence of warm climate properties on phytoplankton dynamics. Electronic Supplementary Material Supplementary material is available to authorized users in the online version of this article at http://dx.doi.org/10.1007/s10750-006-0360-4 Handling editor: K. Martens     

Dynamics and limitations of phytoplankton biomass along a gradient in Mwanza Gulf,southern Lake Victoria (Tanzania)

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Relative importance of periphyton and phytoplankton in turbid and clear vegetated shallow lakes from the Pampa Plain (Argentina): a comparative experimental study

We analyzed experimentally the relative contribution of phytoplankton and periphyton in two shallow lakes from the Pampa Plain (Argentina) that represent opposite scenarios according to the alternative states hypothesis for shallow lakes: a clear lake with submerged macrophytes, and a turbid lake with high phytoplankton biomass. To study the temporal changes of both microalgal communities under such contrasting conditions, we placed enclosures in the littoral zone of each lake, including natural phytoplankton and artificial substrata, half previously colonized by periphyton until a mature stage and half clean to analyze periphyton colonization. In the clear vegetated shallow lake, periphyton chlorophyll a concentrations were 3–6 times higher than those of the phytoplankton community. In contrast, phytoplankton chlorophyll a concentrations were 76–1,325 times higher than those of periphyton in the turbid lake. Here, under light limitation conditions, the colonization of the periphyton was significantly lower than in the clear lake. Our results indicate that in turbid shallow lakes, the light limitation caused by phytoplankton determines a low periphyton biomass dominated by heterotrophic components. In clear vegetated shallow lakes, where nitrogen limitation probably occurs, periphyton may develop higher biomass, most likely due to their higher efficiency in nutrient recycling.     

Spatial and temporal diversity of small shallow waters in river Lužnice floodplain

To propose a concept of their mutual diversity, twenty-nine permanent shallow floodplain pools and oxbows in the river Lužnice floodplain were analysed for area, depth, shape, flooding, and shading by terrestrial vegetation, and sampled in all seasons for their water chemistry, phytoplankton composition and biomass, and zooplankton composition. The sites are regularly flooded, eutrophic, and often shaded by surrounding vegetation. Cryptophyceae, Chrysophyceae and Euglenophyceae dominated the phytoplankton, while Cyanophytes were rare. Within the rich zooplankton assemblage (63 species), cladocerans and rotifers dominated. Correlation matrices and multivariate analyses indicated that shaded and relatively deeper sites had lower oxygen saturation and higher concentrations of PO₄–P and NH₄–N. Shade and relative depth correlated negatively with phytoplankton biomass and number of phytoplankton taxa, and positively with Cryptophytes and large cladocerans—thus indicating poor mixing, poor light availability and low fish pressure on herbivores. Decomposition of leaf litter increased oxygen consumption, while shade from terrestrial vegetation restricted photosynthesis and decreased oxygen production. Larger sites were more species-rich in phytoplankton and supported Euglenophyceae, green algae and rotifers.     

Rarity,ecological memory,rate of floral change in phytoplankton—and the mystery of the Red Cock

In this article, we attempt to estimate the contemporary phytoplankton species pool of a particular lake, by assessing the rate of floral change over a period of 15 years. Phytoplankton time series data from Lake Stechlin, an oligo-mesotrophic lake in the Baltic Lake District (Germany) were used. Of the 254 algal species recorded during the 15-year of studies with roughly biweekly sampling, 212 species were planktonic. In the individual plankton years, the recorded total number of species changed between 97 and 122, of which the number of dominants (>1% contribution to the annual average of total biomass) was only 10–19. The 15-year cumulative number of species exhibited an almost linear increase after an initial saturation phase. This increase was attributed to two reasons: increase of sample size and immigration of species new to the flora. Based on a probabilistic model developed in this study, we estimated the number of co-existing planktonic species of the lake as some 180, and the rate of floral change as 1–2 species per year. Of these co-existing species, only few maintain the matter–energy processing ecosystem functions in any particular plankton year. Selection of these dominants is probably driven by mesoclimatic cycles, coupled with human-induced forcing, like eutrophication. All others are hiding as an ecological memory, in the sense of the capacity or experiences of past states to influence present or future responses of the community. Data analyses suggest that selection of the ‘memory species’ that show temporary abundance increases over shorter (several years) periods are largely dependent upon the dominants. These results show that interspecific interactions and the particular autecological features of the dominants, together with their effects on the whole ecosystem, act as a major organizing force. Some phytoplankton species, like Planktothrix rubescens, are efficient ecosystem engineers with cascading effects of both a top-down and bottom-up nature. Historical scientific data on Planktothrix blooms in Lake Stechlin suggest cyclic patterns in long-term development of phytoplankton which, as the legend of the Red Cock suggests, dates back much further than scientific archives.     

Life strategies, dominance patterns and mechanisms promoting species coexistence in phytoplankton communities along complex environmental gradients

This paper analyses the life strategies, the dominance patterns and the diversity in phytoplankton communities in large and deep lakes. The study was carried out on the largest Italian Lake (Lake Garda) from 1995 to 2000. Different statistical analyses were applied. For phytoplankton the time variable represents a complex environmental gradient driving annual succession; this gradient was made explicit by the application of PCA analyses to the environmental data. The use of Non Metric Multi Dimensional Scaling applied to Bray-Curtis dissimilarity matrices revealed an ordered and cyclic development of phytoplankton every year; the Bray-Curtis index, calculated between pairs of chronologically contiguous samples, was also used as a measure of the community change rate (β_t) over the temporal succession. A significant relationship between β_t and the complex environmental gradient was assessed. Finally, for every phytoplankton species, the optimum conditions for growth and the realised niches were determined. The positioning of the species on the complex environmental gradient, and the contemporaneous application of cluster analysis based on the different specific environmental optima, highlighted primarily the existence of two groups at the extreme of the complex environmental gradient. The first group included the large late winter/spring diatoms, which developed during high water turbulence and strong physical control, high nutrient concentrations, low light conditions and reduced competition. The second group was composed by many heterogeneous summer species characterised by the ability to contrast losses by grazing and sinking in stratified and stable conditions, and the ability of tolerating nutrient deficiency. A third group of species developed during environmental conditions in the middle of the two previous extremes. These included the three master species Mougeotia sp., Fragilaria crotonensis and Planktothrix rubescens/agardhii. The endogenous and exogenous mechanisms promoting species coexistence are discussed, along with the applicability of competitive and equilibrium/non-equilibrium theories to phytoplankton dynamics.     

Artificial destratification of a small tropical reservoir: effects upon the phytoplankton

Seasonal changes in the phytoplankton community of a small tropical reservoir were monitored over a four year period comprising of an initial two seasonal cycles during which the water column stratified strongly for extended periods each year, and two further seasonal cycles after installation of a mechanical aeration system to induce artificial destratification. In the unmanaged reservoir, the concentration of chlorophyll a at 0.5 m reached maximum values (on one occasion > 90 mg m⁻³) when the water column was stratified and the epilimnion was very shallow (ca 2 m depth). The hypolimnion at this time was anoxic (less than 2% oxygen saturation) and had a high concentration of bacteriochlorophyll (100–200 mg m⁻³). The phytoplankton community of the unmanaged reservoir was generally dominated by cyanobacteria (Cylindrospermopsis raciborskii, Anabaena tenericaulis) during the warmer months of the year (November–March) (but replaced by chlorophyta, dinophyceae and euglenophyceae after periods of intense rain) and by bacillariophyceae (Synedra ulna var. chaseana, S. tenera) during the cooler, dry months. In the artificially destratified reservoir (8 h aeration day⁻¹), the phytoplankton community was largely dominated by diatoms except after depletion of the silica content of the water column which caused diatoms to be replaced by cyanobacteria (dominated by A. tenericaulis) and a range of chlorophytes. The changing pattern of stratification and circulation of the water column in the unmanaged reservoir caused repeated disruption of the established phytoplankton assemblage with peaks of high biomass associated with transient cyanobacterial blooms. Continuous aeration and the consequent increase in the ratio mixed: euphotic depth provided conditions suitable for dominance of the phytoplankton by diatoms, as long as silica was available, and resulted in average chlorophyll levels higher than in the unmanaged reservoir (120 ± 10 v. 64 ± 9 mg m⁻²). Hierarchical fusion analysis based on the biomass of species differentiated the phytoplankton samples into cluster groups that could be related primarily to stratification or mixing of the water column.     

Zooplankton–phytoplankton relationships in shallow subtropical versus temperate lakes Apopka (Florida,USA) and Trasimeno (Umbria,Italy)

This study compares and contrasts the dynamics of phytoplankton, zooplankton, and nutrients in two of the largest shallow lakes in the USA (Lake Apopka, Florida) and Europe (Lago Trasimeno, Umbria, Italy) and considers particularly the biomass ratio of zooplankton to phytoplankton (BZ:BP) in relation to nutrient levels and in the context of data from other subtropical and temperate lakes. Lake Apopka is hypereutrophic with higher concentrations of total phosphorus (TP), nitrogen (TN), and nearly an order of magnitude higher BP than Lago Trasimeno. However, combined data from the two lakes can be fit to a single log–log regression model that explains 72% of the variability in BP based on TP. In contrast, BZ has a significant positive log–log relationship with TP only for Lago Trasimeno, and is much lower than expected based on the TP concentrations observed in Lake Apopka. Lake Apopka has a fish assemblage that includes high densities of gizzard shad (Dorosoma cepedianum) and threadfin shad (D. petenense), similar to other eutrophic Florida lakes that also have extreme low BZ. The ratio BZ:BP is below 0.01 in Lake Apopka, 10-fold lower than in Trasimeno and among the lowest values reported in the literature. Although stress of high water temperature and a greater proportion of inedible cyanobacteria may be contributing factors, the collective results support an emerging view that fish predation limits the biomass of crustacean zooplankton in subtropical lakes. Handling editor: S. I. Dodson     

Dynamics and ecological significance of daily internal load of phosphorus in shallow Lake Balaton, Hungary

1. As supported by field data, turbidity recorded by light scattering sensors could reliably be converted into concentration of suspended particulate matter (SPM) and coefficient of vertical light attenuation (K_d) in Lake Balaton. 2. Autocorrelation analysis revealed that proper determination of SPM concentration and K_d required daily sampling. To approximate daily rate of resuspension, 15 min or more frequent measurements were needed. Thus, routine monitoring provides very little insight into environmental variability of shallow lakes as habitats for phytoplankton. 3. The internal P load was estimated from daily rate of resuspension and P desorption capacity of sediments. The latter was assumed to be proportionate to the potentially mobile inorganic P content of SPM. A comparison with net primary production and nutrient status of phytoplankton showed that the proposed method of estimating time series of internal P load captured seasonal trends. 4. The daily rate of resuspension was high whereas that of internal P load was low in Lake Balaton relative to other shallow lakes. The latter reflects favourable behaviour of the calcite‐rich sediments. As a consequence, carrying capacity of Basin 1 of Lake Balaton was P‐determined. 5. The timing of external and internal loads was radically different. While the former showed mostly seasonal changes, large pulses characterised the latter. As a consequence, internal load may supply more P to phytoplankton growth during the critical summer months than external load. However, the relative importance of these sources may show substantial interannual variability. 6. Large resuspension events often followed each other during periods of 10–15 days. It has been shown that disturbances in this frequency range are of key importance in maintaining the diversity of phytoplankton. We propose that resuspension can be perceived not only as a disturbance factor but also as a factor that periodically relaxes nutrient stress. The former feature may dominate the instantaneous effect, whereas the latter may determine the persistent effect of resuspension on succession of phytoplankton.