Whole-lake microalgal biomass surveys were carried out in Lake Balaton to investigate the seasonal, spatial, and temporal changes of benthic algae, as well as to identify the drivers of the phytobenthos. Phytobenthos was controlled mainly by light: the highest benthic algal biomass was in the shallow littoral region characterized by large grain size (sand) with good light availability but lower nutrient content in the sediment. During the investigated period, phytoplankton biomass showed a significant decrease in almost the entire lake. At the same time, the biomass of benthic algae increased significantly in the eastern areas, increasing the contribution of total lake microalgae biomass (from 20 % to 27 %). Benthic algal biomass increase can be explained by the better light supply, owing to the artificially maintained high water level which greatly mitigates water mixing. The decrease in planktonic algal biomass could be attributed to increased zooplankton grazing, which is otherwise negatively affected by mixing. As a result of the high water level, the trophic structure of the lake has been rearranged in recent decades with a shift from the planktonic life form to the benthic one while the nutrient supply has largely remained unchanged.
Occurrence of the smallest phototrophic microorganisms (photoautotrophic picoplankton, APP) in Lake Balaton was discovered in the early 1980s. This triggered a series of systematic studies on APP and resulted in the setting of a unique long-term picoplankton dataset. In this review, we intend to summarize the obtained results and to give a new insight on APP ecology and diversity in Lake Balaton. According to the results, APP dynamics depends on trophic state, temperature, nutrient, and light availability, as well as grazing pressure. APP abundance in Lake Balaton decreased to a low level (1–2 × 10 5 cells mL −1 ) as a consequence of decreasing nutrient supply (oligotrophication) during the past more than two decades, and followed a characteristic seasonal dynamics with higher abundance values from spring to autumn than in winter. Concomitantly, however, the APP contribution to both phytoplankton biomass and primary production increased (up to 70% and 40–50%, respectively) during oligotrophication. Regarding annual pattern, picocyanobacteria are dominant from spring to autumn, while in winter, picoeukaryotes are the most abundant, most likely due to the different light and temperature optima of these groups. Within picocyanobacteria, single cells and microcolonies were both observed with mid-summer dominance of the latter which correlated well with the density of cladocerans. Community-level chromatic adaptation (i.e., dominance of phycoerythrin- or phycocyanin-rich forms) of planktonic picocyanobacteria was also found as a function of underwater light quality. Sequence analysis studies of APP in Lake Balaton revealed that both picocyanobacteria and picoeukaryotes represent a diverse and dynamic community consisting several freshwater genotypes (picocyanobacteria: Synechococcus, Cyanobium ; picoeukaryotes: Choricystis, Stichococcus , Mychonastes, Nannochloris , and Nannochloropsis ).
Lake Balaton, the largest lake in Central Europe, underwent severe eutrophication from the 1960s to the 1990s, due to phosphorus loadings from external anthropogenic sources. The subsequent and complex eutrophication control and lake restoration program resulted in a significant decrease in the external phosphorus loading to the lake. Consequently, Lake Balaton has been returning to its former meso-eutrophic character. In this paper, we explore the long-term dynamics of chlorophyll a (Chl a ) concentration, a proxy for phytoplankton biomass, and zooplankton biomass in Lake Balaton during its re-oligotrophication period from 2001 to 2017, and attempt to draw some conclusions on the subsequent changes in the fish stock. We found a proportional decrease in zooplankton and phytoplankton biomasses at moderate phytoplankton levels. However, below a certain phytoplankton concentration (< 10 μg l −1 Chl a ), the decrease in phytoplankton biomass was not coupled with a further decline in zooplankton biomass because the fraction of small phytoplankton, edible for zooplankton, showed a much smaller decrease in biomass compared with large non-edible phytoplankton. Thus, improvements in water quality (i.e., reduced nutrient loading), partly via concomitant changes in the phytoplankton size distribution, did not cause a large difference in the fish stock in this shallow lake.
Dreissenid mussels (including the zebra mussel Dreissena polymorpha and the quagga mussel D. rostriformis) are among the world's most notorious invasive species, with large and widespread ecological and economic effects. However, their long-term population dynamics are poorly known, even though these dynamics are critical to determining impacts and effective management. We gathered and analyzed 67 long-term (>10 yr) data sets on dreissenid populations from lakes and rivers across Europe and North America. We addressed five questions: (1) How do Dreissena populations change through time? (2) Specifi- cally, do Dreissena populations decline substantially after an initial outbreak phase? (3) Do different measures of population performance (biomass or density of settled animals, veliger density, recruitment of young) follow the same patterns through time? (4) How do the numbers or biomass of zebra mussels or of both species combined change after the quagga mussel arrives? (5) How does body size change over time? We also considered whether current data on long-term dynamics of Dreissena populations are adequate for science and management. Individual Dreissena populations showed a wide range of temporal dynamics, but we could detect only two general patterns that applied across many populations: (1) Populations of both species increased rapidly in the first 1-2 yr after appearance, and (2) quagga mussels appeared later than zebra mussels and usually quickly caused large dedines in zebra mussel populations. We found little evidence that combined Dreissena populations declined over the long term. Different measures of population performance were not congruent; the temporal dynamics of one life stage or population attribute cannot generally be accurately inferred from the dynamics of another. We found no consistent patterns in the long-term dynamics of body size. The long-term dynamics of Dreissena populations probably are driven by the ecological characteristics (e.g., predation, nutrient inputs, water temperature) and their temporal changes at individual sites rather than following a generalized time course that applies across many sites. Existing long-term data sets on dreissenid populations, although dearly valuable, are inadequate to meet research and management needs. Data sets could be improved by standardizing sampling designs and methods, routinely collecting more variables, and increasing support.
Planktivorous silver carp and bighead carp (collectively, the bigheaded carps) have been stocked worldwide and their invasion has caused severe impacts on many freshwater ecosystems. Exploiting the chance provided by the specific hybrid bigheaded carp stock in Lake Balaton (Hungary) covering the entire morphological range between the two species (including gill raker morphology), we implemented a comprehensive study (1) to reveal the feeding habits of hybrid bigheaded carps living in a mesotrophic, lacustrine habitat and (2) to assess how biotic and abiotic environmental factors and gill raker morphology affect diet composition. We found that all bigheaded carps utilized primarily zooplankton and neglected the scarce and inefficiently digestible phytoplankton, irrespective of gill raker morphology. Moreover, we observed strikingly high levels of inorganic debris consumption, but the proportion of inorganic matter in the guts was not associated directly with the concentration of suspended inorganic particles. Variance in the diet composition of bigheaded carps was related mostly to environmental factors, including the wind-induced resuspension of inorganic particles and seasonally variable availability of food resources. In conclusion, the effects of abiotic environmental factors and available food resources could overwhelm the effect of gill raker morphology in shaping the feeding habits of bigheaded carps.
The colonization progress of the invasive bivalve dreissenids, the formerly dominant Dreissena polymorpha and the recently (2008) introduced Dreissena rostriformis bugensis was studied between 2009 and 2013 in the largest Central European shallow lake, Lake Balaton, Hungary. The density of dreissenid planktonic veligers, new settlers (post-veligers and early juveniles), and the population structure (density, length frequency, relative abundance) of the two species were monitored on experimentally introduced natural stone substrata, on different time scales. Dreissenids started dynamic settling following a sudden veliger bloom. As substratum saturation progressed, competition between species for places was suggested, which, after two years, led to an increased number of large individuals (>20mm) and also recruits of D. r. bugensis. By contrast, the population of D. polymorpha was confined to middle size (11–18mm) individuals of the first settler generation. On local substrata, where the benthic community was already established, the replacement of D. polymorpha by D. r. bugensis took longer, but it happened in a similar way. The invasion speed of D. r. bugensis in Lake Balaton resembled the speed obtained in other European water bodies where D. r. bugensis, similar to Lake Balaton, was introduced much later than D. polymorpha. However, a longer replacement process was found in North America, where both species invaded new habitats at the same time. This suggests that the speed, and probably the success, of D. r. bugensis invasion depends on new surface availability, and whether the two dreissenid species are introduced together or at different times.
17β-Estradiol (E2) and 17α-ethinyl estradiol (EE2), which are environmental estrogens, have been determined with LC-MS in freshwater. Their sensitive analysis needs derivatization and therefore is very hard to achieve in multiresidue screening. We analyzed samples from all the large and some small rivers (River Danube, Drava, Mur, Sava, Tisza, and Zala) of the Carpathian Basin and from Lake Balaton. Freshwater was extracted on solid phase and derivatized using dansyl chloride. Separation was performed on a Kinetex XB-C18 column. Detection was achieved with a benchtop orbitrap mass spectrometer using targeted MS analysis for quantification. Limits of quantification were 0.05 ng/L (MS1) and 0.1 ng/L (MS/MS) for E2, and 0.001 ng/L (MS1) and 0.2 ng/L (MS/MS) for EE2. River samples contained n.d.–5.2 ng/L E2 and n.d.–0.68 ng/L EE2. Average levels of E2 and EE2 were 0.61 and 0.084 ng/L, respectively, in rivers, water courses, and Lake Balaton together, but not counting city canal water. EE2 was less abundant, but it was still present in almost all of the samples. In beach water samples from Lake Balaton, we measured 0.076–0.233 E2 and n.d.–0.133 EE2. A relative high amount of EE2 was found in river Zala (0.68 ng/L) and in Hévíz-Páhoki canal (0.52 ng/L), which are both in the catchment area of Lake Balaton (Hungary).
We examined the distribution of submerged and emergent macrophyte species and the entire macrophyte community within and between five lake types (highland reservoirs, alkali lakes, large shallow lakes, small to medium sized shallow lakes, marshes) in the Pannon Ecoregion, Hungary. The lowest submerged, emergent and total species richness was found in alkali lakes. The highest submerged macrophyte richness was in small to medium sized lakes, while the highest emergent macrophyte species richness was in reservoirs, small to medium sized lakes, and marshes. The values of within-lake type beta diversity were generally lower than the values of alpha diversity, especially for submerged macrophytes, indicating between site homogeneity in species composition within the lake types. Emergent macrophyte communities contributed the most to within and between lake type diversity and total (gamma) diversity. Canonical correspondence analyses showed that the main environmental variables which influenced the distribution of submerged macrophytes were conductivity, Secchi transparency and water nitrogen contents. For emergent macrophytes conductivity, lake width, altitude and water depth proved to be the most influential variables. Our results contribute to the knowledge of large-scale distribution of macrophytes in the Pannon Ecoregion and to the identification of conservation value of lakes using macrophytes. The results support the importance of small lakes and artificial lakes in the conservation of macrophyte diversity compared to large and natural lakes in the Pannon Ecoregion.
Reliable estimation of the diet composition of filter-feeding Asian carps is essential to evaluate their effects on ecosystem functioning. In previous studies, the diet composition of these fishes was primarily determined based on the analysis of foregut contents. To assess the reliability of foregut content analysis in diet assessments, these were compared with gill raker filtrate analyses. Gill raker filtrates were found to be more reliable than foregut contents for determining food composition due to higher amounts of sample, significantly higher numbers of identifiable taxa (including both phytoplankton and zooplankton), and considerably higher numbers of intact planktonic individuals. The present findings indicate that diet composition analyses based on foregut samples alone are likely to underestimate the number of individuals and the biomass of planktonic species which are less resistant to digestive processes.
As lake biota are basically determined by hydromorphological characteristics of the lake basin, estab- lishment of a lake typology is the first step in lake quality assessment and management. Based on descriptor vari- ables (size, depth, lake bed material, water-regime and macrophyte coverage), 16 hydromorphological lake-types were established for the Pannonian Ecoregion. Our analyses revealed that descriptor variables have considerable effect both on biomass and composition of phytoplankton. As the phytoplankton biomass of several lake-types can be merged, a simplified typology can be proposed. Since the phytoplankton composition of various lake-types dif- fered significantly in terms of diversity and relative abundance of functional groups, well delineated composition- based types cannot be established for Pannonian lakes. Although the Pannonian Ecoregion is the westernmost part of the Eurasian steppe zone, the proposed typology of the studied area can be extended to lakes of the greater Pannonian region.
After an earthquake, the condition of industrial systems, structures and equipment, especially at nuclear power plants has to be assessed since this information is needed for accident management and for the decision on the continuation of the operation. The damaging potential of the earthquake can be characterized by maximum horizontal acceleration of the ground motion, response spectra, cumulative absolute velocity and different instrumental intensity values. These quantities can be correlated with the earthquake characteristics, magnitude, distance, etc. Some of the indices are selective to damage mechanism and can be correlated to load-bearing features of the structures. Different indicators of damaging potential of earthquakes are analyzed in the paper from the point of view of applicability for post-event condition assessment at nuclear power plants and for using as criteria for restart of the operation.
In this study, we tested whether the spatial distribution of waterbirds is influenced by shoreline urbanization or other habitat characteristics. We conducted monthly censuses along shoreline sections of a continental lake (Lake Balaton, Hungary) to assess the abundance of 11 common species that use this lake as a feeding and staging area during migration and winter. We estimated the degree of urbanization of the same shoreline sections and also measured other habitat characteristics (water depth, extent of reed cover, biomass of zebra mussels, distances to waste dumps and to other wetlands). We applied linear models and model averaging to identify habitat variables with high relative importance for predicting bird distributions. Bird abundance and urbanization were strongly related only in one species. Other habitat variables exhibited stronger relationships with bird distribution: (1) diving ducks and coots preferred shoreline sections with high zebra mussel biomass, (2) gulls preferred sites close to waste dumps, and (3) the abundances of several species were higher on shoreline sections close to other wetlands. Our findings suggest that the distribution of waterbirds on Lake Balaton is largely independent of shoreline urbanization and influenced by food availability and connectivity between wetlands.
The restoration and management of shallow, pond-like systems are hindered by limitations in the applicability of the well-known models describing the relationship between nutrients and lake phytoplankton biomass in higher ranges of nutrient concentration. Trophic models for naturally eutrophic small, shallow, endorheic lakes have not yet been developed, even though these are the most frequent standing waters in continental lowlands. The aim of this study was to identify variables that can be considered as main drivers of phytoplankton biomass and to build a predictive model. The influence of potential drivers of phytoplankton biomass (nutrients, other chemical variables, land use, lake use and lake depth) from 24 shallow eutrophic lakes was tested using data in the Pannonian ecoregion (Hungary and Romania). By incorporating lake depth, TP, TN and lake use as independent and Chl-a as dependent variables into different models (multiple regression model, GLM and multilayer perception model) predictive models were built. These models explained >50% of the variance. Although phytoplankton biomass in small, shallow, enriched lakes is strongly influenced by stochastic effects, our results suggest that phytoplankton biomass can be predicted by applying a multiple stressor approach, and that the model results can be used for management purposes.
During the unusually long European drought between 2000 and 2003, the water level of the large and shallow Lake Balaton, Hungary (area = 596 km2, mean depth = 3.25 m), decreased by 28%. Although food availability for zooplankton remained unchanged, and the fish stock declined more than the water mass, the density of populations of several planktonic rotifers, cladocerans, calanoid copepodes, and veligers decreased by 60‐90% simultaneously with the water‐level decrease and regenerated only after the drought. The generally strong turbulence of the lake was intensified during the four consecutive years of low water, as verified by instrumental monitoring of the turbulence intensity and by the estimation of the turbulent kinetic‐energy dissipation rate. In our tank experiments, turbulence conditions similar to those that existed in the lake during low water were simulated, but mineral suspended material was minimized and food was regularly resupplied. Under these experimental conditions, zooplankton taxa showing the highest mortality were the same as those that were most susceptible in situ. Increased turbulence coupled with the water‐level decrease is especially unfavorable for rotifer Keratella, the cladoceran Daphnia, Bosmina, and copepodit and adult stages of the calanoid copepod Eudiaptomus gracilis in this lake.
Low velocity surface layers can significantly increase ground accelerations during earthquakes. When saturated sandy sediments are present, because of pore pressure increase, decrease of soil strength or even liquefaction can occur. Some volume change follows the dissipation of excess pore pressure after the earthquake resulting surface settlements. To determine the liquefaction probability and post-liquefaction settlement is very important for critical facilities e.g. for the site of Paks Nuclear Power Plant, Hungary. Pore pressure increase and so the liquefaction and surface settlements depend on the characteristics of seismic loading and soil parameters. To quantify the extent of these phenomena is rather difficult. Uncertainties arise both from the probabilistic nature of the earthquake loading and from the simplifications of soil models as well. In the paper, the most important semi-empirical and dynamical effective stress methods for liquefaction and post-liquefaction settlement assessment are summarized. Most significant contributors to the uncertainties are highlighted, and particular examples through the investigation of Paks NPP site are given. Finally, a probabilistic procedure is proposed where the uncertainties will be taken into account by applying a logic tree methodology. At the same time, the uncertainties are reduced by the use of site-specific UHRS and stress reduction factors.