Relationship between bacterial community composition and bottom-up versus top-down variables in four eutrophic shallow lakes

Bacterial community composition was monitored in four shallow eutrophic lakes during one year using denaturing gradient gel electrophoresis (DGGE) of PCR-amplified prokaryotic rDNA genes. Of the four lakes investigated, two were of the clearwater type and had dense stands of submerged macrophytes while two others were of the turbid type characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had high nutrient levels (total phosphorus, >100 micro g liter(-1)) while the other lakes had relatively low nutrient levels (total phosphorus, <100 micro g liter(-1)). For each lake, seasonal changes in the bacterial community were related to bottom-up (resources) and top-down (grazers) variables by using canonical correspondence analysis (CCA). Using an artificial model dataset to which potential sources of error associated with the use of relative band intensities in DGGE analysis were added, we found that preferential amplification of certain rDNA genes over others does not obscure the relationship between bacterial community composition and explanatory variables. Besides, using this artificial dataset as well as our own data, we found a better correlation between bacterial community composition and explanatory variables by using relative band intensities compared to using presence/absence data. While bacterial community composition was related to phytoplankton biomass in the high-nutrient lakes no such relation was found in the low-nutrient lakes, where the bacterial community is probably dependent on other organic matter sources. We used variation partitioning to evaluate top-down regulation of bacterial community composition after bottom-up regulation has been accounted for. Using this approach, we found no evidence for top-down regulation of bacterial community composition in the turbid lakes, while grazing by ciliates and daphnids (Daphnia and Ceriodaphnia) was significantly related to changes in the bacterial community in the clearwater lakes. Our results suggest that in eutrophic shallow lakes, seasonality of bacterial community structure is dependent on the dominant substrate source as well as on the food web structure.     

Characterization of bacterial communities in four freshwater lakes differing in nutrient load and food web structure

The phylogenetic composition of bacterioplankton communities in the water column of four shallow eutrophic lakes was analyzed by partially sequencing cloned 16S rRNA genes and by PCR-DGGE analysis. The four lakes differed in nutrient load and food web structure: two were in a clearwater state and had dense stands of submerged macrophytes, while two others were in a turbid state characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had very high nutrient levels (total phosphorus > 100 microg/l), while the other lakes were less nutrient rich (total phosphorus < 100 microg/l). Cluster analysis, multidimensional scaling and ANOSIM (analysis of similarity) were used to investigate differences among the bacterial community composition in the four lakes. Our results show that each lake has its own distinct bacterioplankton community. The samples of lake Blankaart differed substantially from those of the other lakes; this pattern was consistent throughout the year of study. The bacterioplankton community composition in lake Blankaart seems to be less diverse and less stable than in the other three lakes. Clone library results reveal that Actinobacteria strongly dominated the bacterial community in lake Blankaart. The relative abundance of Betaproteobacteria was low, whereas this group was dominant in the other three lakes. Turbid lakes had a higher representation of Cyanobacteria, while clearwater lakes were characterized by more representatives of the Bacteroidetes. Correlating our DGGE data with environmental parameters, using the BIOENV procedure, suggests that differences are partly related to the equilibrium state of the lake.     

Zooplankton, phytoplankton and the microbial food web in two turbid and two clearwater shallow lakes in Belgium

Components of the pelagic food web in four eutrophic shallow lakes in two wetland reserves in Belgium (Blankaart and De Maten) were monitored during the course of 1998–1999. In each wetland reserve, a clearwater and a turbid lake were sampled. The two lakes in each wetland reserve had similar nutrient loadings and occurred in close proximity of each other. In accordance with the alternative stable states theory, food web structure differed strongly between the clearwater and turbid lakes. Phytoplankton biomass was higher in the turbid than the clearwater lakes. Whereas chlorophytes dominated the phytoplankton in the turbid lakes, cryptophytes were the most important phytoplankton group in the clearwater lakes. The biomass of microheterotrophs (bacteria, heterotrophic nanoflagellates and ciliates) was higher in the turbid than the clearwater lakes. Biomass and community composition of micro- and macrozooplankton was not clearly related to water clarity. The ratio of macrozooplankton to phytoplankton biomass – an indicator of zooplankton grazing pressure on phytoplankton – was higher in the clearwater when compared to the turbid lakes. The factors potentially regulating water clarity, phytoplankton, microheterotrophs and macrozooplankton are discussed. Implications for the management of these lakes are discussed.     

Contrasting bacterioplankton community composition and seasonal dynamics in two neighbouring hypertrophic freshwater lakes

We characterized the bacterioplankton community and its seasonal dynamics in two neighbouring hypertrophic lakes by denaturing gradient gel electrophoresis (DGGE) analysis of short (193 bp) 16S ribosomal DNA polymerase chain reaction (PCR) products obtained with primers specific for the domain Bacteria. Lake Blankaart is turbid and has a high phytoplankton biomass and episodic cyanobacterial blooms, whereas biomanipulated Lake Visvijver is characterized by clearwater conditions and the establishment of a dense charophyte vegetation. Both lakes were dominated by bacterial groups commonly found in freshwater habitats (e.g. ACK4 cluster of Actinomycetes; ACK stands for clones isolated from the Adirondack mountain lakes) . Yet, cluster analysis and principal components analysis (PCA) revealed that taxon composition of the bacterioplankton community of the two lakes differs substantially and consistently throughout the season. During the study year (1998), the bacterioplankton community of both lakes showed a distinct seasonal pattern. Lake Blankaart showed a clear differentiation between winter, spring, summer and autumn. In Lake Visvijver, summer samples differed greatly from spring, autumn and winter samples. We hypothesize that the contrasting bacterioplankton in the two neighbouring shallow lakes is determined largely by the presence or absence of macrophytes.     

Factors regulating summer phytoplankton in a highly eutrophic Antarctic lake

Lakes from Maritime Antarctica are regarded as systems generally inhabited by metazoan plankton capable of imposing a top-down control on the phytoplankton during short periods, while lakes from Continental Antarctica lacking these communities would be typically controlled by scarcity of nutrients, following a bottom-up model. Otero Lake is a highly eutrophic small lake located on the NW of the Antarctic Peninsula, which has no metazoan plankton. During summer 1996, we studied the density, composition and vertical distribution of the phytoplankton community of this lake with respect to various abiotic variables, yet our results demonstrated neither light nor nutrient limitation of the phytoplankton biomass. Densities of heterotrophic nanoflagellates (HNAN) and ciliates from three different size categories were also studied. Extremely low densities of HNAN (0–155 ind. ml^–1) could be due to feeding competition by bacterivore nanociliates and/or predation by large ciliates. A summer bloom of the phytoflagellate Chlamydomonas aff. celerrima Pascher reached densities tenfold those of previous years (158.10³ ind. ml^–1), though apparently curtailed by a strong peak of large ciliates (107 ind. ml^–1) which would heavily graze on PNAN (phototrophic nanoflagellates). Top-down control can thus occur in this lake during short periods of long hydrologic residence time.     

Covariation between zooplankton community composition and cyanobacterial community dynamics in Lake Blaarmeersen (Belgium)

The cyanobacterial community composition in the mesotrophic Lake Blaarmeersen was determined by denaturing gradient gel electrophoresis (DGGE) of PCR-amplified 16S rRNA gene fragments during two consecutive years to assess the importance of different classes of explanatory variables (bottom-up and top-down factors, physical variables and phytoplankton) in cyanobacterial community dynamics. The most dominant cyanobacteria in Lake Blaarmeersen were Synechococcus (three genotypes), Limnothrix redekei and Anabaena/Aphanizomenon. Analyses of Similarity revealed that the cyanobacterial community in Lake Blaarmeersen differed significantly between the growing season and the winter season as well as between the epilimnion and hypolimnion during the stratified periods. Mantel tests revealed significant correlations between the DGGE data and bottom-up factors, physical variables, the phytoplankton community composition and, interestingly, the zooplankton community composition. In general, the zooplankton community composition (especially the cladoceran community) was more important in structuring the cyanobacterial community than the total zooplankton biomass. This study shows that grazing zooplankton communities can have a relatively strong impact on the cyanobacterial community dynamics and that this impact can be equally important as bottom-up processes regulated by nutrient concentrations and/or physical variables.     

Dynamics of Bacterial Community Composition and Activity during a Mesocosm Diatom Bloom

Bacterial community composition, enzymatic activities, and carbon dynamics were examined during diatom blooms in four 200-liter laboratory seawater mesocosms. The objective was to determine whether the dramatic shifts in growth rates and ectoenzyme activities, which are commonly observed during the course of phytoplankton blooms and their subsequent demise, could result from shifts in bacterial community composition. Nutrient enrichment of metazoan-free seawater resulted in diatom blooms dominated by a Thalassiosira sp., which peaked 9 days after enrichment (≈24 μg of chlorophyll a liter⁻¹). At this time bacterial abundance abruptly decreased from 2.8 × 10⁶ to 0.75 × 10⁶ ml⁻¹, and an analysis of bacterial community composition, by denaturing gradient gel electrophoresis (DGGE) of PCR-amplified 16S rRNA gene fragments, revealed the disappearance of three dominant phylotypes. Increased viral and flagellate abundances suggested that both lysis and grazing could have played a role in the observed phylotype-specific mortality. Subsequently, new phylotypes appeared and bacterial production, abundance, and enzyme activities shifted from being predominantly associated with the <1.0-μm size fraction towards the >1.0-μm size fraction, indicating a pronounced microbial colonization of particles. Sequencing of DGGE bands suggested that the observed rapid and extensive colonization of particulate matter was mainly by specialized α-Proteobacteria- and Cytophagales-related phylotypes. These particle-associated bacteria had high growth rates as well as high cell-specific aminopeptidase, β-glucosidase, and lipase activities. Rate measurements as well as bacterial population dynamics were almost identical among the mesocosms indicating that the observed bacterial community dynamics were systematic and repeatable responses to the manipulated conditions.     

How important is the crustacean plankton for the maintenance of water clarity in shallow lakes with abundant submerged vegetation?

• 1 We measured the abundance and biomass of filter‐feeding microcrustacean zooplankton and calculated their grazing impact on phytoplankton biomass during summer in five shallow, mesotrophic to eutrophic lakes. For three of the lakes data exist both from years with dense submerged vegetation and low turbidity (the clearwater state), as well as from years characterised by sparse vegetation and high turbidity (the turbid state). In the other two lakes data are available only for clearwater conditions.
• 2 In all lakes conditions of dense vegetation and clear water coincided with a low abundance of crustacean plankton during summer. In the three lakes that shifted, the calculated biovolume ingested by crustacean plankton (filtering rate) was 3–11 times lower during clearwater conditions compared with turbid conditions. Because phytoplankton biomass was lower during clearwater conditions, however, daily grazing pressure from microcrustacea (expressed as percentage of phytoplankton biomass) did not differ between states. In three of the five lakes, grazers were estimated to take less than 10% of the phytoplankton biomass per day, indicating filtration by zooplankton was not the most important mechanism to maintain clearwater conditions.
• 3 High densities of Cladocera were found in three of the lakes within dense stands of Charophyta. However, these samples were dominated by plant‐associated taxa that even during the night were rarely found outside the vegetation. This indicates that plant‐associated zooplankton has no major influence on the maintenance of water clarity outside the vegetation.
• 4 Spring peak abundance of Cladocera was observed in three of the lakes. In two of these, where seasonal development was studied in both the clearwater and the turbid state, spring peaks were lower during the clearwater state.
• 5 Predation, low food availability or a combination of both may explain the low zooplankton densities. Phytoplankton may be limited by low phosphorus availability in the lakes dominated by Charophyta. Our results indicate that the importance of zooplankton grazing may have minor importance for the maintenance of the clearwater state in lakes with dense, well‐established submerged vegetation.

     

Vertical distribution of bacterial community structure in the sediments of two eutrophic lakes revealed by denaturing gradient gel electrophoresis (DGGE) and multivariate analysis techniques

Vertical distribution of bacterial community structure was investigated in the sediments of two eutrophic lakes of China, Lake Taihu and Lake Xuanwu. Profiles of bacterial communities were generated using a molecular fingerprinting technique, denaturing gradient gel electrophoresis (DGGE) followed by DNA sequence analysis, and the results were interpreted with multivariate statistical analysis. To assess changes in the genetic diversity of bacterial communities with changing depth, DGGE banding patterns were analysed by cluster analysis. Distinct clusters were recognized in different sampling stations of Lake Taihu. Canonical correspondence analysis (CCA) was carried out to infer the relationship between environmental variables and bacterial community structure. DGGE samples collected at the same sampling site clustered together in both lakes. Total phosphorus, organic matter and pH were considered to be the key factors driving the changes in bacterial community composition.     

Bottom-Up Effects on Biomass Versus Top-Down Effects on Identity: A Multiple-Lake Fish Community Manipulation Experiment

The extent to which ecosystems are regulated by top-down relative to bottom-up control has been a dominant paradigm in ecology for many decades. For lakes, it has been shown that predation by fish is an important determinant of variation in zooplankton and phytoplankton community characteristics. Effects of fish are expected to not only be a function of total fish biomass, but also of functional composition of the fish community. Previous research on the importance of trophic cascades in lakes has largely focused on the role of zooplanktivorous and piscivorous fish. We conducted a large-scale multiple-lake fish community manipulation experiment to test for the effect of differences in fish functional community composition on the trophic structure of lakes. We examine the effect of top-down and bottom-up factors on phytoplankton and zooplankton biomass as well as on their community composition. We put our data in a broader perspective by comparing our results to data of a survey that also included ponds with low fish densities as well as ponds with very high densities of fish. Our results indicate that the overall food web structure under relative high fish densities is primarily structured by bottom-up factors, whereas community characteristics seem to be primarily regulated by top-down factors. Our results suggest a subtle interplay between bottom-up and top-down factors, in which bottom-up factors dominate in determining quantities while top-down effects are important in determining identities of the communities.     

Comparative and experimental approaches to top-down and bottom-up regulation of bacteria

The regulation of bacterial community biomass and productivity by resources and predators is a central concern in the study of microbial food webs. Resource or bottom-up regulation refers to the limitation of bacteria by carbon and nutrients derived from allocthonous inputs, primary production, and heterotrophic production. Predatory or top-down regulation refers to the limitation of bacteria below levels supportable by resources alone. Large scale comparative studies demonstrate strong correlations between bacterial productivity and biomass, suggesting significant resource regulation. Comparisons of the abundances of heterotrophic flagellates and bacteria, however, imply that in some cases there may be top-down regulation of bacteria in eutrophic environments. Experimental studies in lakes support the importance of resource regulation and reveal little top-down control from protozoans. Increases in bacterial abundance and production with nutrient enrichment were limited in enclosure experiments with high abundances of the cladoceran, Daphnia. Regulation of bacteria by Daphnia may occur in many lakes seasonally and prevail in some lakes throughout the year where these animals sustain dense populations. In most situations, however, bacteria appear to be limited primarily by resources.     

Bottom-up versus top-down effects on ciliate community composition in four eutrophic lakes (China)

Previous studies have shown that ciliate plankton is generally controlled by food resources (e.g., algae) and predators (e.g., metazooplankton). Among lakes with similar trophic levels but different distributions of phyto- and metazooplankton, the main forces acting on ciliate assemblages may be different. We investigated the relationship between ciliate communities and bottom-up versus top-down variables based on a survey of four subtropical eutrophic lakes (China). Two of the lakes (Chaohu, Taihu) are located on the Mid-lower Yangtze Plain near sea level, and the other two (Dianchi, Xingyunhu) on the Yunnan-Kweichow Plateau at 1700 m above sea level. Blooms of cyanobacteria developed during summer in Lakes Chaohu and Taihu and throughout the year in Lakes Dianchi and Xingyunhu. Ciliate functional feeding groups differed significantly between lakes. The results of canonical correspondence analysis (CCA) and variation partitioning showed that cyanobacteria significantly influence ciliate species, whereas ‘edible’ algae (cryptophytes, diatoms) and cladocerans were the important variables in explaining the ciliate community structure of Lakes Dianchi and Xingyunhu compared with Lakes Taihu and Chaohu. Our results highlight the importance of consistent cyanobacterial blooms in shaping the ciliate community in subtropical eutrophic shallow lakes by interacting with top-down and bottom-up factors.     

Bacterioplankton Community Shifts in an Arctic Lake Correlate with Seasonal Changes in Organic Matter Source

Seasonal shifts in bacterioplankton community composition in Toolik Lake, a tundra lake on the North Slope of Alaska, were related to shifts in the source (terrestrial versus phytoplankton) and lability of dissolved organic matter (DOM). A shift in community composition, measured by denaturing gradient gel electrophoresis (DGGE) of 16S rRNA genes, occurred at 4°C in near-surface waters beneath seasonal ice and snow cover in spring. This shift was associated with an annual peak in bacterial productivity ([¹⁴C]leucine incorporation) driven by the large influx of labile terrestrial DOM associated with snow meltwater. A second shift occurred after the flux of terrestrial DOM had ended in early summer as ice left the lake and as the phytoplankton community developed. Bacterioplankton communities were composed of persistent populations present throughout the year and transient populations that appeared and disappeared. Most of the transient populations could be divided into those that were advected into the lake with terrestrial DOM in spring and those that grew up from low concentrations during the development of the phytoplankton community in early summer. Sequencing of DNA in DGGE bands demonstrated that most bands represented single ribotypes and that matching bands from different samples represented identical ribotypes. Bacteria were identified as members of globally distributed freshwater phylogenetic clusters within the α- and β-Proteobacteria, the Cytophaga-Flavobacteria-Bacteroides group, and the Actinobacteria.     

The Native Bacterioplankton Community in the Central Baltic Sea Is Influenced by Freshwater Bacterial Species

The Baltic Sea is one of the largest brackish environments on Earth. Despite extensive knowledge about food web interactions and pelagic ecosystem functioning, information about the bacterial community composition in the Baltic Sea is scarce. We hypothesized that due to the eutrophic low-salinity environment and the long water residence time (>5 years), the bacterioplankton community from the Baltic proper shows a native “brackish” composition influenced by both freshwater and marine phylotypes. The bacterial community composition in surface water (3-m depth) was examined at a single station throughout a full year. Denaturing gradient gel electrophoresis (DGGE) showed that the community composition changed over the year. Further, it indicated that at the four extensive samplings (16S rRNA gene clone libraries and bacterial isolates from low- and high-nutrient agar plates and seawater cultures), different bacterial assemblages associated with different environmental conditions were present. Overall, the sequencing of 26 DGGE bands, 160 clones, 209 plate isolates, and 9 dilution culture isolates showed that the bacterial assemblage in surface waters of the central Baltic Sea was dominated by Bacteroidetes but exhibited a pronounced influence of typical freshwater phylogenetic groups within Actinobacteria, Verrucomicrobia, and Betaproteobacteria and a lack of typical marine taxa. This first comprehensive analysis of bacterial community composition in the central Baltic Sea points to the existence of an autochthonous estuarine community uniquely adapted to the environmental conditions prevailing in this brackish environment.     

Dimethylsulfoniopropionate Turnover Is Linked to the Composition and Dynamics of the Bacterioplankton Assemblage during a Microcosm Phytoplankton Bloom

Processing of the phytoplankton-derived organic sulfur compound dimethylsulfoniopropionate (DMSP) by bacteria was studied in seawater microcosms in the coastal Gulf of Mexico (Alabama). Modest phytoplankton blooms (peak chlorophyll a [Chl a] concentrations of ~2.5 μg liter⁻¹) were induced in nutrient-enriched microcosms, while phytoplankton biomass remained low in unamended controls (Chl a concentrations of ~0.34 μg liter⁻¹). Particulate DMSP concentrations reached 96 nM in the enriched microcosms but remained approximately 14 nM in the controls. Bacterial biomass production increased in parallel with the increase in particulate DMSP, and nutrient limitation bioassays in the initial water showed that enrichment with DMSP or glucose caused a similar stimulation of bacterial growth. Concomitantly, increased bacterial consumption rate constants of dissolved DMSP (up to 20 day⁻¹) and dimethylsulfide (DMS) (up to 6.5 day⁻¹) were observed. Nevertheless, higher DMSP S assimilation efficiencies and higher contribution of DMSP to bacterial S demand were found in the controls compared to the enriched microcosms. This indicated that marine bacterioplankton may rely more on DMSP as a source of S under oligotrophic conditions than under the senescence phase of phytoplankton blooms. Phylogenetic analysis of the bacterial assemblages in all microcosms showed that the DMSP-rich algal bloom favored the occurrence of various Roseobacter members, flavobacteria (Bacteroidetes phylum), and oligotrophic marine Gammaproteobacteria. Our observations suggest that the composition of the bacterial assemblage and the relative contribution of DMSP to the overall dissolved organic sulfur/organic matter pool control how efficiently bacteria assimilate DMSP S and thereby potentially divert it from DMS production.     

Quantitative responses of lake phytoplankton to eutrophication in Northern Europe

Based on the currently largest available dataset of phytoplankton in lakes in northern Europe, we quantified the responses of three major phytoplankton classes to eutrophication. Responses were quantified by modelling the proportional biovolumes of a given group along the eutrophication gradient, using generalized additive models. Chlorophyll-a (Chl-a) was chosen as a proxy for eutrophication because all classes showed more consistent responses to Chl-a than to total phosphorus. Chrysophytes often dominate in (ultra-) oligotrophic lakes, and showed a clear decrease along the eutrophication gradient. Pennate diatoms were found to be most abundant at moderate eutrophication level (spring-samples). Cyanobacteria often dominate under eutrophic conditions, especially in clearwater lakes at Chl-a levels >10 μg l⁻¹ (late summer samples). We compare the relationships among types of lakes, based on the lake typology of the northern geographic intercalibration group, and among countries sharing common lake types. Significant differences were found especially between humic and clearwater lakes, and between low- and moderately alkaline lakes, but we could not identify significant differences between shallow and deep lakes. Country-specific differences in response curves were especially pronounced between lakes in Norway and Finland, while Swedish lakes showed an intermediate pattern, indicating that country-specific differences reflect large-scale geographic and climatic differences in the study area.     

Distribution of planktonic ciliates in highly coloured subtropical lakes: comparison with clearwater ciliate communities and the contribution of myxotrophic taxa to total autotrophic biomass

SUMMARY 1. The planktonic ciliate communities of eleven organically coloured north and central Florida lakes representing a variety of trophic conditions were examined during 1979–80. The total abundance and biomass of ciliates were not significantly different from comparable clearwater lakes and only minor taxonomic replacements were noted at the order level.
2. Timing of population peaks of oligotrophic lakes was dissimilar to clearwater lakes of the same trophic state, but seasonality in meso-trophic and eutrophic lakes resembled patterns described for comparable clearwater lakes.
3. Various ciliate components were strongly correlated with chlorophyll a concentrations, but only moderately correlated to dominant phytoplankton groups. No significant correlations were found between ciliate components and bacterial abundance.
4. Myxotrophic taxa numerically dominated oligotrophic systems, particularly during midsummer, and accounted for a large percentage of the total ciliate biomass. Estimates of the ciliate contribution to total autotrophic biomass indicate that these zoochlorellae-bearing protozoa may account for much of the autotrophic biomass during midsummer periods in coloured lakes, and thus may lead to an overestimation of phytoplankton standing crops available to zooplankton grazers if chlorophyll a is used as a surrogate measure of algal biomass.     

Effects of different fish species and biomass on plankton interactions in a shallow lake

Thirty-six enclosures, surface area 4 m², were placed in Little Mere, a shallow fertile lake in Cheshire, U.K. The effects of different fish species (common carp, common bream, tench and roach) of zooplanktivorous size, and their biomass (0, 200 and 700 kg ha^–1) on water chemistry, zooplankton and phytoplankton communities were investigated. Fish biomass had a strong effect on mean zooplankton size and abundance. When fish biomass rose, larger zooplankters were replaced by more numerous smaller zooplankters. Consequently phytoplankton abundance rose in the presence of higher densities of zooplanktivorous fish, as zooplankton grazing was reduced. Fish species were also significant in determining zooplankton community size structure. In enclosures with bream there were significantly greater densities of small zooplankters than in enclosures stocked with either carp, tench and, in part, roach. When carp or roach were present, the phytoplankton had a greater abundance of Cyanophyta than when bream or tench were present. Whilst top-down effects of fish predation controlled the size partitioning of the zooplankton community, this, in turn apparently controlled the bottom-up regeneration of nutrients for the phytoplankton community. At the zooplankton–phytoplankton interface, both top-down and bottom-up processes were entwined in a reciprocal feedback mechanism with the extent and direction of that relationship altered by changes in fish species. This has consequences for the way that top-down and bottom-up processes are generalised.     

On the annual variation of phytoplankton biomass in Finnish inland waters

Annual variations in phytoplankton biomass in 63 lakes in Southern and Central Finland are discussed. Biomass is rather small during winter (January–April), usually <0.05 mg l^–1 (fresh weight) and there are no differences between oligotrophic and eutrophic lakes. In early spring and in autumn biomass varies widely, depending mainly on water temperature. Phytoplankton biomass is smaller in July than in June and August in oligotrophic lakes (biomass <0.20 mg l^–1 fresh weight) and mesotrophic (biomass 1.0–2.5 mg l^–1) lakes, but greater in eutrophic (biomass 2.5–10.0 mg l^–1) and hypereutrophic (biomass >10.0 mg l^–1) lakes. The standard deviation of phytoplankton biomass in Finnish inland waters is usually smallest in July, which facilitates the comparison of phytoplankton between different kinds of lakes.     

Longitudinal Changes in the Bacterial Community Composition of the Danube River: a Whole-River Approach

The Danube River is the second longest river in Europe, and its bacterial community composition has never been studied before over its entire length. In this study, bacterial community composition was determined by denaturing gradient gel electrophoresis (DGGE) analysis of PCR-amplified portions of the bacterial 16S rRNA gene from a total of 98 stations on the Danube River (73 stations) and its major tributaries (25 stations), covering a distance of 2,581 km. Shifts in the bacterial community composition were related to changes in environmental conditions found by comparison with physicochemical parameters (e.g., temperature and concentration of nutrients) and the concentration of chlorophyll a (Chl a). In total, 43 distinct DGGE bands were detected. Sequencing of selected bands revealed that the phylotypes were associated with typical freshwater bacteria. Apparent bacterial richness in the Danube varied between 18 and 32 bands and correlated positively with the concentration of P-PO₄ (r = 0.56) and negatively with Chl a (r = −0.52). An artificial neural network-based model explained 90% of the variation of apparent bacterial richness using the concentrations of N-NO₂ and P-PO₄ and the distance to the Black Sea as input parameters. Between the cities of Budapest and Belgrade, apparent bacterial richness was significantly lower than that of other regions of the river, and Chl a showed a pronounced peak. Generally, the bacterial community composition developed gradually; however, an abrupt and clear shift was detected in the section of the phytoplankton bloom. Large impoundments did not have a discernible effect on the bacterial community of the water column. In conclusion, the riverine bacterial community was largely influenced by intrinsic factors.