Analysis of long-term ecological status of Lake Balaton based on the ALMOBAL phytoplankton database

Lake Balaton (Hungary), one of the largest lakes in Europe, has undergone eutrophication and restoration during the last two decades. The first quantitative phytoplankton records date back to the 1930s, and since that time thousands of data have been published or accumulated in counting protocols or computer sheets. These data provide material for both scientific analyses (e.g. effects of global change) and applications (e.g. estimation of reference state for the Water Framework Directive). The ALMOBAL phytoplankton database was developed to provide computing support for these applications. It stores data in standardized forms, handles synonyms and allows analyses to be conducted on the basis of floral records, numbers or biomass. The analysis includes records of about 3000 phytoplankton samples taken during the past 60 years from two representative basins in Lake Balaton. This article represents the first attempt at historical reconstruction of the ecological status and compares it with changes in trophic state and current water quality. The results indicate that the phytoplankton biomass and community structure found in the early 1960s could be regarded as reference conditions. Statistical analyses support the view that late summer phytoplankton assemblages are the most sensitive indicators of trophic change, and clearly show the eutrophication of the lake that occurred from the mid-1970s to the mid-1990s and the restoration during the last decade. An additional advantage is that, since quality estimation is based on relative biomass, the method can be used to reconstruct water quality in cases when counting protocols are available, but some basic data for quantitative estimates are missing.     

Relationships between phytoplankton and fish yields in Lake Balaton

Coefficients of determination between fish yields in Lake Balaton (kg/ ha) and different measurements of phytoplankton abundance were: phytoplankton biomass (0.48), chlorophyll a (0.45). The same indices for catch per unit of effort (CUE) proved to be 0.82 and 0.88, respectively. The correlation between fish yields and phytoplankton density has been described in second-degree polynomials.     

Identification versus counting protocols as sources of uncertainty in diatom-based ecological status assessments

The heterogeneity of oxygen distribution in a Hediste diversicolor burrow environment was investigated in a laboratory experiment using a 6-mm thick tank equipped with oxygen planar optodes. The two-dimensional oxygen distribution in a complete burrow was monitored every 2 min for 4 h. Oxygen concentrations fluctuated over a scale of minutes in the burrow lumen and wall (up to 2 mm) reflecting the balance between worm ventilation activity and oxygen consumption. The magnitude of the three surrounding micro-horizons (oxic, oscillating and anoxic) induced by the intermittent worm ventilation was spatially and temporally variable within the structure. Oxygen variations appeared to be controlled by distance from the sediment–water interface and the direction of water circulation. Moreover, there was an apparent ‘buffer effect’, induced by the proximity to the overlying water, which reduced the variations of lumen and wall oxygen in the upper part of the structure. These results highlight the heterogeneous distribution and dynamics of oxygen associated with H. diversicolor burrows and ventilation activity. They also highlight the necessity of integrating this complexity into the current burrow-base models in order to estimate the ecological importance of burrowing species in coastal ecosystems.     

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.     

Effect of streamlining taxa lists on diatom-based indices: implications for intercalibrating ecological status

Intercalibration of ecological status class boundaries between member states is a requirement of the European Union’s Water Framework Directive. Although a preliminary intercalibration of boundaries established for phytobenthos has been performed, a number of questions remain, including the extent to which variations in taxonomic concepts used in different member states influences the position of these boundaries. In this paper, the robustness of the diatom-based metrics used for intercalibration is assessed. Whilst use of genus-level identification led to a loss of ecological information, merging representatives of closely-related taxa has little effect on these metrics. Similarly, taxa that occur only rarely or never have high relative abundances in a dataset can also be ignored without the loss of ecological information. Similar results were obtained when modern taxonomic concepts were compared with concepts in use 80 years ago. Fine scale taxonomy may play a valuable role within member states; however, our results suggest that, at a continental scale, a simplified approach to diatom taxonomy should not affect intercalibration results.     

Application of a new multi-metric phytoplankton index to the assessment of ecological status in marine and transitional waters

Patterns of phytoplankton size spectra variation with gradients of environmental stress have been observed in freshwater, transitional waters and marine ecosystems, driving the development of size spectra based assessment tools.In this study, we have tested on transitional and coastal waters a new Index of Size spectra Sensitivity of Phytoplankton (ISS-Phyto), which integrates simple size spectra metrics, size class sensitivity to anthropogenic disturbance, phytoplankton biomass (chlorophyll a) and taxonomic richness thresholds. ISS-Phyto has been tested both among and within ecosystems along pressure gradients based on expert view assessment; the adequacy of symmetric and both left and right asymmetric models of phytoplankton size class sensitivity have been compared.The results showed that ISS-Phyto consistently discriminated between anthropogenic and natural disturbance conditions. Left asymmetric models of size spectra sensitivity, assuming greater disturbance tolerance with respect to eutrophication and organic enrichment of increasingly large size classes, showed the best fit comparing all ecosystems; in three of the four considered ecosystems (Varna, Helsinki, Mompás-Pasaia), they seemed to discriminate best between different levels of disturbance also within ecosystems. Moreover, they demonstrated significant and inverse patterns of variation along the overall pressure gradient as well as along the inorganic phosphorus (DIP), chlorophyll a and trophic index (TRIX) gradients.Therefore, ISS-Phyto, originally developed for transitional waters, seems to be an adequate assessment tool of ecological status also in coastal marine waters; moreover, it seems adequate to describe within ecosystem disturbance gradients. Hence, ISS-Phyto helps to understand the relationships between anthropogenic impact and ecosystem response from the individual point of view, with reference to the simple parameter of body size.     

Continuous monitoring of phytoplankton dynamics in Lake Balaton (Hungary) using on-line delayed fluorescence excitation spectroscopy

1. This study introduces delayed fluorescence (DF) excitation spectroscopy as an on‐line tool for in situ monitoring of the composition and biomass of various colour classes of phytoplankton when they are photosynthetically active (cyanobacteria, chlorophytes, chromophytes and cryptophytes). The DF data are validated by comparison with those from conventional methods (weekly microscopic counts and the measurement of chlorophyll concentration). 2. The composition of phytoplankton as assessed by DF agreed reasonably well with the results from microscopic counts, particularly when differences in chlorophyll‐specific DF integrals of the various colour classes were taken into account. 3. Integrals of DF spectra were converted into concentration of chlorophyll a using empirical factors derived from field data. The value of the conversion factor was nearly twice as high when the relative abundance of cyanobacteria was low (<15%) than when it was high. The converted DF‐chl time series agreed well with chlorophyll measurements particularly when blooms were developing. As the DF method is inherently free of the interference caused by pigment degradation products, the discrepancy between the two data sets increased during the collapse of blooms and when sediment resuspension was intense. 4. Fourier spectrum analysis of the time series of DF‐chl indicated that samples must be taken, at a minimum, every 2–3 days to capture the dynamics of phytoplankton. As a consequence, the dynamics of various algal blooms, including their timing, duration and net growth rate, could be estimated with greater confidence than by using conventional methods alone. 5. On‐line DF spectroscopy is an advanced technique for monitoring daily the biomass and composition of the photosynthetically active phytoplankton in aquatic environments, including turbid shallow lakes. At present, the detection limit is around 1 mg DF‐chl a m^?3 in terms of total biomass but confidence in estimates of phytoplankton composition declines sharply below about 5 mg chl a m^?3. 6. On‐line DF spectroscopy represents a promising approach for monitoring phytoplankton. It will be useful in water management where it can act as an early‐warning system of declines in water quality. In basic ecological research it can supplement manual methods. While default calibration spectra may be acceptable for routine monitoring, we suggest a careful individual calibration of the DF spectrometer for basic research. The statistical methods developed here help to assess the adequacy of various calibration sets.     

Stir-up effect of wind on a more-or-less stratified shallow lake phytoplankton community,Lake Balaton,Hungary

Microstratification of phytoplankton in the large shallow Lake Balaton (Hungary) was studied during a 24 h period. Dissolved O₂ showed biological stratification; flagellates exhibited a definite circadian rhythm. In the middle of the investigation a heavy storm broke out which was followed by the disappearance of differences between different layers of water. Storm-induced destratification is described by cluster-analysis. Abundances of dominant species changed differently in connection with the storm. Numbers of Nitzschia sp. increased due to stirring up from the sediment surface. Numbers of single-celled or colony-forming species (Cyclotella comta, Crucigenia quadrata, Coelosphaerium kuetzingianum) practically did not change. Numbers of all the three dominant filamentous species (Aphanizomenon fos-aquae f. klebahnii, Lyngbya limnetica, Planctonema lauterbornii) significantly decreased, which might be attributed to an unknown loss process and was followed by a competitive displacement by algae of small cell size.     

Orthophosphate uptake of planktonic microorganisms in Lake Balaton

Seasonal orthophosphate uptake of natural plankton has been studied in hypertrophic and mesotrophic basins of Lake Balaton. Initial uptake of carrier-free labelled phosphoric acid followed first order kinetics in accordance with a single two-compartment model. In isotope equilibrium from 1.2 to 9.7 per cent of radioactivity was found in the filtrate. Turnover times of orthophosphate in water were some minutes in summer and two orders of magnitude longer in winter and April. Turnover time of phosphorus in microorganisms has been estimated on the basis of equilibrium isotope distribution between water and particular phases and turnover time in water. It ranged from 28 to over 1 000 min. The shortest times were found in summer, and the longest in winter. Turnover time of orthophosphate in water increased in relation to phosphate additions according to Michaelis-Menten kinetics. The values of Michaelis-constant plus the ambient orthophosphate concentration (K + S_n) suggested an overestimation of real phosphate concentrations by chemical measurements, being much lower than soluble reactive phosphorus contents in most cases, as it is well-known in many other lakes. Both K + S_n values and maximum uptake velocities (V) were significantly higher in the hypertrophic basin than in the mesotrophic one, in contrast to essentially similar values of turnover times. Maximum uptake velocity per unit chlorophyll-a P _opt:V ratio showed extreme to moderate phosphorus deficiency for Lake Balaton plankton, which seemed to decrease somewhat in the course of eutrophication.     

Oligotrophication of Lake Balaton over a 20-year period and its implications for the relationship between phytoplankton and zooplankton biomass

Hydrobiologia - 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...     

太湖流域生态风险评价

随着城镇的急剧扩张和经济的快速增长,流域生态环境遭到极大冲击和破坏,致使生态系统出现资源退化、环境恶化与灾害风险加剧的趋势,生态环境面临前所未有的挑战.从复合生态系统入手,深入分析流域内各生态系统要素之间的相互作用与影响机制,综合考虑多风险源、多风险受体和生态终点共存情况下的风险大小,从风险源危险度、生境脆弱度及受体损失度三方面构建了流域生态风险评价技术体系,并选取太湖流域为实证区域,对太湖流域2000年、2008年两个时期生态风险的时空演化特征进行评价与分析.结果表明:太湖流域生态风险指数介于0.015-0.253之间,以中等和较低生态风险为主.至2008年,高、较高生态风险所占面积逐渐扩大,已由2000年的5.66％、13.42％增加至6.05％、18.42％,主要集中在流域北部的常州市区、江阴市大部分地区以及无锡市区.     

The role of storms in the summer succession of the phytoplankton community in a shallow lake (Lake Balaton, Hungary)

The authors carried out day-to-day studies on the phytoplanktonof the largest shallow lake of Central Europe in the summerof 1976, 1977 and 1978. Data were analysed with diversity andcluster analyses. (i) Wind-induced stirring-up of the sedimentplays important role in the summer phytoplankton successionof the shallow lake. (ii) After storms algae with very smallcell sizes show synchronous development. In long calm periodsalgae with larger cell sizes exhibit organised population dynamics.(iii) In calm periods a shift from r-selection to predominantK-selection can be suspected. After the next storm importanceof r-selectionists again increases. (iv) Effect of changinggrowth- and loss rates on the population dynamics of the mostabundant species is discussed based on measured generation timesand calculated elimination. The phenomena discussed in the papercan appear to different extents in the summer phytoplanktonsuccession of other shallow, polimictic lakes.     

Transformations between organic and inorganic sediment phosphorus in Lake Balaton

In order to estimate microbial P content and biological P uptake in sediments, the tungstate precipitation method of Orrett & Karl (1987) was used in sediment extracts. This method allows a simple and rapid separation of organic and inorganic ³²P radioactivity. Either inorganic ³²P (as carrierfree H₃ ³²PO₄) or organic ³²P (as ³²P-labelled algal material) was added to surface sediment suspensions of shallow Lake Balaton. Inorganic ³²P was rapidly transformed into organic ³²P, and this process was completely inhibited by formaline. P content of living benthic microorganisms was estimated from steady state distribution of the radioactivity. Transformation of algal organic P into inorganic P could also be detected.In extremely P limited Lake Balaton benthic microorganisms were shown to supplement their high P requirements by inorganic P uptake. The velocity of the inorganic into organic P transformation, i.e. the rate of microbial P uptake, was comparable to P uptake in the water column. Microbial P uptake contributed significantly to total P fixation by sediments, particularly at low ( 100 µg P l^–1) phosphate additions.     

Stability and change of phytoplankton communities in a highly dynamic environment—the case of large,shallow Lake Balaton (Hungary)

Time series data of key environmental variables (water temperature, global radiation, vertical light attenuation, internal P load) and biomass of four colour classes of photosynthetically active algae were collected during 2003 and 2004 with daily resolution. Using these data, seasonal patterns of phytoplankton were analyzed as a function of the dynamic environment. Abstraction of the environmental state as a point in multi-dimensional space was used to identify habitat templates of bloom-forming groups and derive an indicator of environmental stability/physical disturbance. These templates were synthesized into a simple threshold model that sufficiently simulated development and collapse of various blooms. Blooms were, however, rare events related to specific environments with strong, unidirectional forcing. Tentative quantification of disturbance and compositional stability/community change allowed discriminating disturbance-driven changes and autogenic succession with reasonable success. The two processes were found to be equally important in shaping the composition and biomass of phytoplankton.     

Mobility of phosphorus fractions in the sediments of Lake Balaton

Sediment phosporus was studied by a combination of the fractionation procedure of Hieltjes & Lijklema and the isotopic dilution technique in a mesotrophic (Tihany) and a hypertrophic (Keszthely) basin of Lake Balaton.In the calcareous sediments the largest part of phosphorus was bound to calcium. Iron-bound and residual P showed higher concentrations at Keszthely than at Tihany. There was little loosely adsorbed P at both locations. Vertical differences in P fractions of the sediments were more pronounced at Keszthely than at Tihany. Exchangeability of the fractions decreased in the following sequence: iron-bound > loosely adsorbed > calcium-bound > residual. Phosphorus, particularly in the calcium-bound fraction, was more mobile at Keszthely, and its exchangeability decreased rapidly downwards at both stations. Four times more potentially mobile phosphorus has been accumulated by the upper sediment layer at Keszthely than at Tihany.     

Phytopatological problems and solutions in the walnut orchards along Lake Balaton

European or Persian Walnut (Juglans regia) is an important and healthy food as well as base material of timber industry. Several pests (pathogens and insect pests) may cause serious damages on walnut. These are less known on the crop land of the tree. Results of some years of our experiments including bacteriological and mycological studies, are presented in this paper. The optimum time of chemical protection against the walnut blight (Xanthomonas arboricola pv. juglandis) was determined. Occurrences of pathogenic fungi were surveyed in an orchard and on home garden trees in Hungary (18 fungus species were identified). The following experimental results are reported on the pathogenic fungi: cultivar resistance to walnut anthracnose (Gnomonia leptostyla), dying of wood parts in the cultivar collection, application of the spore trap, in vitro fungicide testing against Phomopsis juglandina.     

Selection of phytoplankton associations in Lake Balaton,Hungary, in response to eutrophication and restoration measures,with special reference to the cyanoprokaryotes

Seasonal and spatial patterns of aquatic primary production were compared in a tidal creek (Estero Pargo) bordered by mangroves and open waters of Terminos Lagoon, Mexico. Comparisions were made during a 17-month period in 1990–91 that spanned dry, rainy, and storm or 'Norte' seasons. Annual net primary productivity was 478 g C m-2 yr-1 in the lagoon and 285 g C m-2 yr-1 in the tidal creek. In some months, there were significant differences in primary production between the two sites. In both areas, the highest productivity occurred in summer at the start of the rainy season (June 1991), and the lowest production occurred during the dry season from February to May. Aquatic primary production was lower during the dry season of 1991 in comparison to 1990, possibly related to unusually low precipitation during 1991. Seasonal changes in water column productivity were correlated to variations in light and precipitation. The effect of runoff from mangrove forests was assessed by serial additions of surface water from a fringe forest to bottle incubations of lagoon water. Small additions of mangrove water stimulated primary production in lagoon waters during all seasons. The net productivity was extremely sensitive to aliquot volume; small amounts (0.3 and 1.7% of total volume) were stimulatory, increasing rates by > 50% in 7 of 12 experiments. The greatest effect occurred in September, 1990, when productivity tripled after an amendment with 1 ml (0.3% by volume) of mangrove water. Additions greater than 3% of total volume generally led to reduction in net productivity probably due to the inhibitory effect of humic substances. In many tropical systems, tidal exchange of estuarine waters with mangrove forests is likely to be important to enhancing water column productivity by exporting organic nutrients and other growth-enhancing substances to the estuary. This revised version was published online in July 2006 with corrections to the Cover Date.     

Trophic relationships between primary producers and fish yields in Lake Balaton

Relationships between chlorophyll a content of the water, the shoreline-length: water area ratio and the annual total fish yield as catch per unit effort (CUE: kg ha^–1 100 h^–1 as annual mean values) have been calculated by multivariable regression. The determination coefficient (r ² = 0.913) showed a significant dependence of fish yield on morphometry of different lake areas. Accordingly, fish carrying capacity of the open water areas, calculated from chlorophyll a content and S/A, ranged from 12 to 34%, but that of the littoral zone between 66 and 88%. These findings have also been supported by echo-sounding records of the horizontal distribution of fish.Bream (Abramis brama L.) contributes the majority (70–80%) of fish stock and yield. Its food mainly consists of zooplankton and benthic invertebrates in ratios that are widely variable with season and depend on the age of fish. Average daily food consumption of individuals (age group 3 + and over) varies between 2 and 5 g. Bream consumes two- to three-times more food in the SW basin than in the NE one. This means that the present stocks inhabiting areas from NE to SW consume annually 13249–20085 t yr^–1 of food. According to estimated calorific values, the annual energy consumption of local populations along the longitudinal axis of the lake varies between 93 and 141 kJ m^–2 yr^–1. The efficiency of energy transfer from primary producers to fish is low and varies from 0.04 to 0.1%.     

The food of bream (Abramis brama L.) in two basins of Lake Balaton of different trophic status

Contrary to earlier observations the food composition of bream (Abramis brama L.) in Lake Balaton has changed in parallel with eutrophication. These changes were probably caused by increased population density and connected with density dependent growth as well as sharpened inter- and intraspecific competition in the cyprinid community. According to the frequency of occurrence, zooplankton followed by benthic invertebrates comprised the majority of the bream's food. By weight, however, benthic food dominated in both basins. Size-related dietary changes were pronounced, but statistically not always significant, suggesting the overall importance of the most abundant and available prey types. Bream did not strongly select any of the zooplankton groups, however, seasonal changes in food composition alternated in parallel with the trophic gradient along the longitudinal axis of the lake.