Microbial food web in a large shallow lake (Lake Balaton, Hungary)

Seasonal variations of phyto-, bacterio- and colourless flagellate plankton were followed across a year in the large shallow Lake Balaton (Hungary). Yearly average chlorophyll-a concentration was 11 µg 1^–1, while the corresponding values of bacterioplankton and heterotrophic nanoflagellate (HNF) plankton biomass (fresh weight) were 0.24 mg 1^–1 and 0.35 mg 1^–1, respectively. About half of planktonic primary production was channelled through bacterioplankton on the yearly basis. However, there was no significant correlation between phytoplankton biomass and bacterial abundance. Bacterial specific growth rates were in the range of 0.009 and 0.09 h^–1, and ended to follow the seasonal changes in water temperature. In some periods of the year, predator-prey relationships between the HNF and bacterial abundance were obvious. The estimated HNF grazing on bacteria varied between 3% and 227% of the daily bacterial production. On an annual basis, 87% of bacterial cell production was grazed by HNF plankton.     

An attempt to trace eutrophication in a shallow lake (Balaton,Hungary) using chironomids

Lake Balaton, the largest shallow lake in Central Europe, is about 20 000 years old. An enormous increase in tourism and the disproportionate building development of the last few decades has resulted in the acceleration of eutrophication in the lake. Widespread research to reveal the causes of water-quality deterioration and possible ways of protection against it have recently started. The investigation of the larvae of non-biting midges (Diptera: Chironomidae) in the sediment of the open-water zone has also begun. The contemporary faunal composition strongly correlates with the trophic gradient along the longitudinal axis of the lake. We therefore supposed that the eutrophication process should be identifiable from the analysis of subfossil chironomid head capsules from the upper (15 cm thick) layer of the sediment. We found that quantitative results could only be obtained when fragments as well as relatively intact head capsules are considered. Our data verify that the originally oligo-mesotrophic community has been gradually replaced by eutrophic species in a west to east direction. Large-bodied larvae belonging to the Chironomus plumosus group mix the sediment down to 15 cm as they build their tubes and consequently alter the original proportions of head capsules at the different levels. So the sequence of communities through the sediment-layers is not quite reliable.     

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.     

Distribution of submerged macrophytes along environmental gradients in large,shallow Lake Balaton (Hungary)

Distribution of submerged macrophytes was assessed at 15 littoral sites in large, shallow Lake Balaton using the echo sounding technique. Sites represented the conspicuous longitudinal and transverse environmental gradients. Absence/presence of plants, water depth, bottom consistency and distance from the shore have been derived from echo sounding data. Turbidity and meteorological variables were recorded at a single site from 2004 to 2006. Site-specific variability in underwater light conditions (an indicator of stress) and exposure to breaking waves (an indicator of disturbance) were estimated by simple models.     

Composition,density and feeding of crustacean zooplankton community in a shallow,temperate lake (Lake Balaton,Hungary)

Composition density and filtering rates of crustacean zooplankton were studied in the open water and among the macrophytes of the oligo-mesotrophic part of Lake Balaton from 1981 to 1983. From the individual filtering rates of the different populations and the densities community grazing rates were derived.Copepoda made up 79–90% of crustacean plankton community in the open lake and 95–97% of it in littoral zone. Among them the nauplii dominated. At the end of summer 1982, when Anabaenopsis was in bloom, the filter-feeding species (Eudiaptomus, Daphnia) practically disappeared, being replaced by cyclopoids. Daphnia had the highest filtering rates followed by those of Eudiaptomus and Diaphanosoma. Among copepods, the filtering rates in decreasing order were: ovigerous > all adults > copepodites > nauplii. > . The filtering rates of the different species varied both seasonally as well as from year to year. In 1983, when the concentration of organic seston decreased, filtering rate increased compared with those in the earlier years. During the water bloom in 1982, the rates decreased by 70% on the average.The community grazing rate was very low (3% per day) in the open lake and among macrophytes, both in 1981 and 1982; also the share of crustacean zooplankton in grazing was very low. In 1983, together with the improving of water quality, the community grazing rate increased 4-fold. In 1981 and 1983 the rates were influenced by water temperature but in 1982 by seston concentration.     

Contributions of benthic and planktonic primary producers to nitrate and ammonium uptake fluxes in a nutrient-poor shallow coastal area (Corsica, NW Mediterranean)

By using the stable isotope ¹⁵N, we have measured in situ the uptake of nitrate and ammonium by the seagrass Posidonia oceanica, its leaf epiphyte community, the brown macroalgae Halopteris scoparia and the suspended particulate organic matter (SPOM). In Revellata Bay (Gulf of Calvi, Western Corsica), which is a very nutrient-poor region, the specific uptake rates (V) (μg N g N⁻¹ h⁻¹) of SPOM measured at ambient concentrations are 10-1000 higher than those of benthic primary producers. Macroalgae have intermediary V, between the seagrass leaf and leaf epiphytes. V are quite variable and the reasons for this variability remain unclear.Despite the difference of specific uptake rates found between benthic and pelagic primary producers, when integrating the uptake fluxes for a water column of 10 m depth, the contribution of benthic primary producers to N uptake fluxes (g N m⁻² h⁻¹) is significant, corresponding on average to 40% of total uptake flux. This results from the dominance in terms of N biomass of benthic primary producers in this shallow nutrient-poor area. When reported for the entire volume of the Revellata Bay, the contribution of benthic primary producers is reduced to 5-10% of total N uptake flux.Although this contribution could appear relatively low, it results in a significant direct transfer of inorganic nitrogen from the water column to the benthic compartment. By this transfer, the benthic plants act as a biological pump incorporating the pelagic N into the benthic compartment for a time longer than the characteristic time of phytoplankton dynamics (month-years vs. day-week).     

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.     

Diversity patterns of trait-based phytoplankton functional groups in two basins of a large, shallow lake (Lake Balaton, Hungary) with different trophic state

The application of functional approaches in understanding phytoplankton community-level responses to changes in the environment has become increasingly widespread in recent years. Eutrophication is known to cause profound modifications in ecosystem processes; however, the underlying relationships between environmental drivers and phytoplankton diversity and functioning are complex and largely unknown. Therefore, in the present study, we investigated and compared the temporal diversity patterns of phytoplankton functional groups in the mesotrophic eastern and eutrophic western basin of the shallow Lake Balaton situated in Hungary. Diversity data were derived from taxonomic composition and biomass data corresponding to the years 2005–2006 and 2008–2009. With the use of cluster analysis, phytoplankton species were classified into eight distinct groups representing different combinations of functionally relevant traits including greatest axial linear dimension; surface-to-volume ratio; photosynthetic pigment composition; N₂ fixation; phagotrophic potential; growth form/complexity (unicell, filamentous, colony-, or coenobium-forming); and motility. Our results have revealed that there is a significant inverse relationship between the functional group diversity used in our study and trophic state (total phytoplankton biomass) as opposed to species diversity, where no correlation was observed. In addition, group evenness showed an even stronger negative correlation with trophic state, while species evenness yielded only a weak relationship. The observed variability in functional group diversity suggests that such an approach could provide an efficient means of revealing structural changes in phytoplankton communities, establishing new hypotheses and highlighting fundamental points in ecosystem functioning.     

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.     

Nitrogen uptake and the importance of internal nitrogen loading in Lake Balaton

1. The importance of various forms of nitrogen to the nitrogen supply of phytoplankton has been investigated in the mesotrophic eastern and eutrophic western basin of Lake Balaton.
2. Uptake rates of ammonium, urea, nitrate and carbon were measured simultaneously. The uptake rates were determined using N and C methodologies, and N2‐fixation was measured using the acetylene‐reduction method. The light dependence of uptake was described with an exponential saturation equation and used to calculate surface‐related (areal) daily uptake.
3. The contribution of ammonium, urea and nitrate to the daily nitrogen supply of phytoplankton varied between 11 and 80%, 17 and 73% and 1 and 15%, respectively. N2‐fixation was negligible in the eastern basin and varied between 5 and 30% in the western region of the lake. The annual external nitrogen load was only 10% of that utilized by algae.
4. The predominant process supplying nitrogen to the phytoplankton in the lake is the rapid recycling of ammonium and urea in the water column. The importance of the internal nutrient loading is emphasized.     

Changes of organic matter quality along the longitudinal axis of a large shallow lake (Lake Balaton)

Hydrobiologia - Dissolved organic matter (DOM) is quantitatively the most significant pool of organic matter in lakes. Within DOM, the pool of dissolved organic carbon (DOC) is dominated...     

Phytoplankton production and phosphorus supply in Cayuga Lake (1968–1973)

The average annual rate of carbon production in Cayuga Lake, New York, was determined by ¹⁴C uptake and by a phosphorus supply method. The rate of phosphorus supply was converted to units of carbon production using observed sestonic C : P ratios. The two methods gave good agreement and annual carbon production was estimated to be 130 gC m⁻² yr⁻¹. Research supported by Federal Hatch Funds and by U.S. Department of Interior, Office of Water Resources Research.     

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.     

Forests losing large quantities of nitrogen have elevated 15N:14N ratios

Summary Urea (U) and ammonium nitrate (AN) had been applied to a Scots pine (Pinus sylvestris L.) forest in northern Sweden for 18 consecutive years at four doses resulting in total N applications ranging from 0 to 1980 kg ha^–1. The ¹⁵N abundance ( ¹⁵N) of the grass Deschampsia flexuosa (L.) Trin. increased linearly (from –0.7 to 11.0) with application rate in the case of U. The response to AN was in the same direction but smaller. While others have shown that the initial response of nitrogen-limited systems to additions of N is a change of ¹⁵N abundance towards that of added N, this study shows that further and excessive additions leads to a retention of ¹⁵N. Monitoring ¹⁵N abundance over time in dose-response trials of this type thus opens new possibilities to estimate critical loads of N and the point of nitrogen saturation.     

Phytoplankton biomass and distribution in a shallow eutrophic lake (Lake George,Uganda)

Summary Lake George, a shallow lake in western Uganda, supports a permanent and dense crop of phytoplankton, and may be regarded as eutrophic although the ambient concentrations of inorganic nitrogen and phosphorus are low. The figures for the annual nutrient loadings (Viner and Smith, 1973) would suggest, however, a eutrophic lake when analysed on the scheme of eutrophication proposed by Vollenweider (1968).The horizontal distribution of chlorophyll a shows a concentric pattern, with the maximum values occurring towards the center. This horizontal variation is thought to reflect water movements, and a general pattern of water flow is proposed.The majority of the species show no seasonal variation but populations of both Anabaena and Melosira show annual variations. These two species also have a horizontal distribution pattern which is the reverse of other species. This pattern is used to support the proposed movements of water.Phytoplankton generation times are discussed in relation to the diurnal cycle occurring in Lake George.     

A15N tracer study to compare nitrogen supply by Azolla and ammonium sulphate to IR8 rice plants grown under flooded conditions

Summary A pot experiment was carried out using a Bangladesh sandy loam paddy soil of pH 6.9 to compare the rates at which nitrogen from Azolla and ammonium sulphate was available to a high yielding rice variety, IR8, grown for 60 days in pots with 4 cm standing flood water.¹⁵N tracer studies confirm that nitrogen from ammonium sulphate was more available to the rice plants than from Azolla. An application of 6, 9 and 18 mg N of Azolla pot^–1 (each pot contained 250 g soil) increased shoot dry matter yields by 13, 29 and 49% for an uptake of 19, 36 and 85% more nitrogen; the corresponding increases on using ammonium sulphate were 33, 54 and 114% for an increased uptake of 57, 90 and 177% more nitrogen, respectively. About 34% of applied¹⁵N of Azolla was taken up by the rice plants in 60 days but 61% of¹⁵N of the ammonium sulphate was absorbed during this period. About 45% of the Azolla-N was released in 60 days, 55% remained in the soils as undecomposed material and 11% was lost as gas. The gaseous loss of¹⁵N from ammonium sulphate was 14%; 25% remained in the soils.     

Leaf vs. epiphyte nitrogen uptake in a nutrient enriched Mediterranean seagrass (Posidonia oceanica) meadow

In situ nitrogen uptake by leaves and epiphytes was studied in a Mediterranean seagrass (Posidonia oceanica) meadow impacted from a fish farm and a pristine meadow, using ¹⁵NH₄ and ¹⁵NO₃ as tracers. In the impacted meadow both leaves and epiphytes yielded higher N concentrations and showed higher specific N uptake, suggesting a linkage between N uptake and its accumulation. Epiphytes took up N faster than leaves in relation to their corresponding biomass, but when assessed per unit area, N uptake was higher in leaves. Leaf N uptake was negatively correlated with epiphyte N uptake. With increasing epiphyte load on leaves, N leaf uptake decreased while N epiphyte uptake increased, indicating that epiphyte overgrowth hinders N uptake by P. oceanica leaves. Epiphyte contribution to total N uptake increased, while that of leaves decreased at the impacted meadow. However, 2-3 times less N was transferred daily from the water column to the benthic compartment, through seagrass and epiphyte uptake on total, at the impacted meadow. Therefore, it is probably still the loss of the key species - the seagrass - which plays the most important role in N cycling in seagrass ecosystems.     

Nitrate deposition in northern hardwood forests and the nitrogen metabolism of Acer saccharum marsh

It is generally assumed that plant assimilation constitutes the major sink for anthropogenic Nitrate NO ₃ ^– deposited in temperate forests because plant growth is usually limited by nitrogen (N) availability. Nevertheless, plants are known to vary widely in their capacity for NO ₃ ^– uptake and assimilation, and few studies have directly measured these parameters for overstory trees. Using a combination of field and greenhouse experiments, we studied the N nutrition of Acer saccharum Marsh. in four northern hardwood forests receiving experimental NO ₃ ^– additions equivalent to 30 kg N ha^–1 year^–1. We measured leaf and fine-root nitrate reductase activity (NRA) of overstory trees using an in vivo assay and used ¹⁵N to determine the kinetic parameters of NO ₃ ^– uptake by excised fine roots. In two greenhouse experiments, we measured leaf and root NRA in A. saccharum seedlings fertilized with 0–3.5 g NO ₃ ^– –N m^–2 and determined the kinetic parameters of NO ₃ ^– and NH ₄ ⁺ uptake in excised roots of seedlings. In both overstory trees and seedlings, rates of leaf and fine root NRA were substantially lower than previously reported rates for most woody plants and showed no response to NO ₃ ^– fertilization (range = non-detectable to 33 nmol NO ₂ ^– g^–1 h^–1). Maximal rates of NO ₃ ^– uptake in overstory trees also were low, ranging from 0.2 to 1.0 mol g^–1 h^–1. In seedlings, the mean V _max for NO ₃ ^– uptake in fine roots (1 mol g^–1 h^–1) was approximately 30 times lower than the V _max for NH ₄ ⁺ uptake (33 mol g^–1 h^–1). Our results suggest that A. saccharum satisfies its N demand through rapid NH ₄ ⁺ uptake and may have a limited capacity to serve as a direct sink for atmospheric additions of NO ₃ ^– .     

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.     

Phytoplankton community structure and succession in the water column of Lake Naivasha,Kenya: a shallow tropical lake

Lake Naivasha is a shallow freshwater lake in the Rift Valley of Kenya. Since the 1980s, when the lake showed a seasonal shift between diatom and cyanobacterial dominance it has become moderately eutrophic. Its algal assemblage is now dominated by a persistent Aulacoseira italica population both numerically and in terms of contribution to overall primary production. Algal and cyanobacterial counts were used to derive Simpson's diversity, succession rate and total community succession, focusing on the 10 most numerically abundant taxa. 170 species were identified, 43 of which were in common with the 143 found in 1979–80, before the increase in trophic state. Most diatoms are indicators of moderate to high nutrient conditions. There is little horizontal or vertical variation in successional processes throughout the lake and although the absolute abundance of cells varies widely, proportional composition is relatively stable. In Crescent Island lagoon, the only regularly stratified site, hypolimnetic succession rates are lower than those in the epilimnion. Overall, community composition is controlled by mixing (and hence light regime) and nutrient availability. With `endless summer' conditions and full mixing, there is a successional pattern of `muted seasonality' adapted to physical instability and environmental stability.