Effect of grazers and viruses on bacterial community structure and production in two contrasting trophic lakes

Background

Over the last 30 years, extensive studies have revealed the crucial roles played by microbes in aquatic ecosystems. It has been shown that bacteria, viruses and protozoan grazers are dominant in terms of abundance and biomass. The frequent interactions between these microbiological compartments are responsible for strong trophic links from dissolved organic matter to higher trophic levels, via heterotrophic bacteria, which form the basis for the important biogeochemical roles of microbial food webs in aquatic ecosystems. To gain a better understanding of the interactions between bacteria, viruses and flagellates in lacustrine ecosystems, we investigated the effect of protistan bacterivory on bacterial abundance, production and structure [determined by 16S rRNA PCR-DGGE], and viral abundance and activity of two lakes of contrasting trophic status. Four experiments were conducted in the oligotrophic Lake Annecy and the mesotrophic Lake Bourget over two seasons (early spring vs. summer) using a fractionation approach. In situ dark vs. light incubations were performed to consider the effects of the different treatments in the presence and absence of phototrophic activity.

Results

The presence of grazers (i.e. < 5-μm small eukaryotes) affected viral production positively in all experiments, and the stimulation of viral production (compared to the treatment with no eukaryotic predators) was more variable between lakes than between seasons, with the highest value having been recorded in the mesotrophic lake (+30%). Viral lysis and grazing activities acted additively to sustain high bacterial production in all experiments. Nevertheless, the stimulation of bacterial production was more variable between seasons than between lakes, with the highest values obtained in summer (+33.5% and +37.5% in Lakes Bourget and Annecy, respectively). The presence of both predators (nanoflagellates and viruses) did not seem to have a clear influence upon bacterial community structure according to the four experiments.

Conclusions

Our results highlight the importance of a synergistic effect, i.e. the positive influence of grazers on viral activities in sustaining (directly and indirectly) bacterial production and affecting composition, in both oligotrophic and mesotrophic lakes.     

Bottom-up versus top-down control of hypo- and epilimnion free-living bacterial community structures in two neighboring freshwater lakes

Bacterioplankton plays a central role in the microbial functioning of lacustrine ecosystems; however, factors that constrain its structural variation are still poorly understood. Here we evaluated the driving forces exerted by a large set of environmental and biological parameters on the temporal and spatial dynamics of free-living bacterial community structures (BCS) in two neighboring perialpine lakes, Lake Bourget and Lake Annecy, which differ in trophic status. We analyzed monthly data from a 1-year sampling period at two depths situated in the epi- and hypolimnia for each lake. Overall, denaturing gradient gel electrophoresis (DGGE) revealed significant differences in the BCS in the two lakes, characterized by a higher number of bands in the oligotrophic ecosystem (i.e., Lake Annecy). The temporal dynamics of BCS differed greatly between depths and lakes, with temporal scale patterns being much longer in the mesotrophic Lake Bourget. Direct-gradient multivariate ordination analyses showed that a complex array of biogeochemical parameters was the driving force behind BCS shifts in both lakes. Our results indicated that 60 to 80% of the variance was explained only by the bottom-up factors in both lakes, indicating the importance of nutrients and organic matter from autotrophic origin in controlling the BCS. Top-down regulation by flagellates together with ciliates or viruses was found only in the hypolimnion and not in the epilimnion for both lakes and explained less than 18% of the bacterial community changes during the year. Our study suggests that the temporal dynamics of the free-living bacterial community structure in deep perialpine lakes are dependent mainly on bottom-up factors and to a lesser extent on top-down factors, whatever the specific environmental conditions of these lakes.     

Variations in Abundance,Genome Size,Morphology, and Functional Role of the Virioplankton in Lakes Annecy and Bourget over a 1-Year Period

We sampled the surface waters (2–50 m) of two deep peri-alpine lakes over a 1-year period in order to examine (1) the abundance, vertical distribution, genome size, and morphology structures of the virioplankton; (2) the virus-mediated bacterial mortality; and (3) the specific genome size range of double-stranded DNA (dsDNA) phytoplankton viruses. Virus-like particle (VLP) concentrations varied between 4.16?×?10⁷ (January) and 2.08?×?10⁸ part mL^?1 (May) in Lake Bourget and between 2.7?×?10⁷ (June) and 8.39?×?10⁷ part mL^?1 (November) in Lake Annecy. Our flow cytometry analysis revealed at least three viral groups (referred to as virus-like particles 1, 2, and 3) that exhibited distinctive dynamics suggestive of different host types. Phage-induced bacterial mortality varied between 6.1 % (June) and 33.2 % (October) in Lake Bourget and between 7.4 % (June) and 52.6 % (November) in Lake Annecy, suggesting that viral lysis may be a key cause of mortality of the bacterioplankton. Virioplankton genome size ranged from 27 to 486 kb in Lake Bourget, while it reached 620 kb in Lake Annecy for which larger genome sizes were recorded. Our analysis of pulsed field gel electrophoresis bands using different PCR primers targeting both cyanophages and algal viruses showed that (1) dsDNA viruses infecting phytoplankton may range from 65 to 486 kb, and (2) both cyanophage and algal “diversity” were higher in Lake Annecy. Lakes Annecy and Bourget also differed regarding the proportions of both viral families (with the dominance of myoviruses vs. podoviruses) and infected bacterial morphotypes (short rods vs. elongated rods), in each of these lakes, respectively. Overall, our results reveal that (1) viruses displayed distinct temporal and vertical distribution, dynamics, community structure in terms of genome size and morphology, and viral activity in the two lakes; (2) the Myoviridae seemed to be the main cause of bacterial mortality in both lakes and this group seemed to be related to VLP2; and (3) phytoplankton viruses may have a broader range of genome size than previously thought. This study adds to growing evidence that viruses are diverse and play a significant role in freshwater microbial dynamics and more globally lake functioning. It highlights the importance of further considering this biological compartment for a better understanding of plankton ecology in peri-alpine lakes.     

The importance of herbivory by protists in lakes of a tropical floodplain system

Inland aquatic ecosystems play a critical role in the global carbon cycle, processing a great fraction of the organic matter coming from terrestrial ecosystems, and the microbial food web is crucial in this process. Thus, we aimed to evaluate whether the food resource of planktonic protozoa comes mainly from small primary producers or heterotrophic bacteria in tropical shallows lakes, assuming the hypothesis that, in general, picocyanobacteria would be the main food resource for protists. We also expected that the autotrophic fraction would be mainly related to protists at the surface of the environments, while the heterotrophic fraction would be more important at the lower strata of the water column. We performed size-fractionation experiments to evaluate the effects of predation of protists on heterotrophic bacteria and picocyanobacteria. We also sampled planktonic organisms at the subsurface and bottom of 20 lakes in a Neotropical floodplain. We found an herbivory preference of heterotrophic flagellates, while ciliates seem to exert a stronger impact on heterotrophic bacteria. We also found no relationship between heterotrophic bacteria and protists in the field data, whereas positive relationships between picocyanobacteria and protists were observed in environments where there was sunlight. Thus, both heterotrophic bacteria and picocyanobacteria were important components in the food webs of tropical shallow lakes. Moreover, the trophic cascade caused by zooplankton predation suggests that protists are efficient in transferring the energy from the base of microbial food webs to higher trophic levels.     

Budding and prosthecate bacteria from freshwater habitats of various trophic states

Budding and prosthecate bacteria were enumerated in spring and summer by viable counting procedures in several freshwater habitats in Australia including oligotrophic lakes, a mesotrophic lake, and eutrophic ponds.Caulobacter spp. were the most numerous type encountered. They were present in the highest concentrations (exceeding 1000/ml) in the mesotrophic lake during the summer. Their proportion to total viable heterotrophic bacteria was also highest (35.1 to 37.7) in this habitat. From 17 to 330/mlCaulobacter spp. were counted in the eutrophic habitats where their proportion to total viable numbers was less than 1.0%. In the oligotrophic lakes they varied from 5 to 23/ml and comprised greater than 5% of the total viable count.Hyphomicrobium- like bacteria were also numerous in the mesotrophic lake and in one oligotrophic lake during the summer sampling period.Ancalomicrobium spp. occurred in high concentrations (130/ml) in the mesotrophic lake. Budding bacteria of thePlanctomyces-Pasteuria group were most numerous in the eutrophic habitats where as many as 240/ml were counted; their proportion to total heterotrophs remained relatively constant regardless of trophic state, however. A similar pattern was observed withProsthecobacter spp.     

Virus and microbial loop dynamics over an annual cycle in three contrasting Antarctic lakes

1. Viral and microbial loop dynamics were investigated over an annual cycle in three contrasting saline Antarctic lakes – Highway Lake (salinity 4‰), Pendant Lake (salinity 19‰) and Ace Lake, a meromictic system (with a mixolimnion salinity of 18‰) in order to assess the importance of viruses in extreme, microbially dominated systems. 2. Virus like particles (VLP) showed no clear seasonal pattern, with high concentrations occurring in both winter and summer (range 0.89 × 10⁷ ± 0.038 to 12.017 × 10⁷ ± 1.28 mL^?1). VLP abundances reflected lake productivity based on chlorophyll a concentrations. Bacterial abundances and biomass did not correlate with VLP numbers except in Pendant Lake, the most productive of the three lakes studied. 3. Pendant Lake supported the highest bacterial biomass (range Highway: 18.44 ± 1.35 to 59.43 ± 2.80 ng C mL^?1; Ace: 14.42 ± 2.69 to 68.39 ± 2.95 ng C mL^?1; Pendant: 31.36 ± 3.94 to 115.95 ± 4.49 ng C mL^?1) so that virus to bacteria ratios (VBR) (range 30.48 ± 7.96 to 96.67 ± 8.21) were higher in Ace Lake (range 30.58 ± 3.98 to 80.037 ± 1.60) and Highway Lake (range 18.63 ± 3.12 to 126.74 ± 6.50). 4. Negative correlations occurred between VLP and cryptophytes (dominant phototrophic nanoflagellates), suggesting that they were not hosts to lytic viruses. Among the other protists only the heterotrophic nanoflagellates of Highway Lake (dominated by the marine choanoflagellate Diaphanoeca grandis) showed a positive correlation with VLP. 5. The VLP was negatively correlated with photosynthetically active radiation (PAR) and temperature, both of which increased with ice thinning and breakout, increasing viral decay. In winter VLP probably persisted in cold, dark water. 6. High VLP concentrations and high VBR (values at the upper end of those reported for marine and lacustrine systems) indicated that viruses, most of which were probably bacteriophage, are a major element within the microbial communities in extreme, saline lakes.     

Sensitivity and responses of diatoms to climate warming in lakes heavily influenced by humans

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Distribution of Viruses and Dissolved DNA along a Coastal Trophic Gradient in the Northern Adriatic Sea

The distribution of viral and other microbial abundances as well as the concentrations of dissolved DNA (D-DNA) along a trophic gradient in the northern Adriatic Sea were determined. Virus abundances, covering a range of 1.2 × 10⁹ to 8.7 × 10¹⁰ liter^-1 were on average 2.5-fold higher in eutrophic than in mesotrophic stations. A 2.5-fold enrichment was also measured for chlorophyll a concentrations, whereas the densities of bacteria and heterotrophic nanoflagellates were only approximately 1.5-fold higher. The frequency of bacteria containing mature phage increased linearly with bacterial abundance. Assuming that mature phage is only visible during the last 14 to 27% of the latent period (L. M. Proctor, A. Okubo, and J. A. Fuhrman, Microb. Ecol. 25:161-182, 1993), we estimated that between 3.5 and 7.3% of the bacterial population was infected at mesotrophic stations versus between 7.0 and 19.5% at eutrophic stations, indicating that the bacterial mortality due to viral lysis might increase with the degree of eutrophication. The frequency of bacteria with mature phage and the burst size varied significantly with the bacterial morphotype; rod-shape cells, the most abundant morphotype, showed low infection rates but a high burst size. Concentrations of D-DNA varied significantly with season but not with trophic conditions. The estimated percentage of viral DNA on total D-DNA concentrations averaged 17.1% (range, 0.7 to 88.3%). Some kind of interaction between heterotrophic nanoflagellates and viruses is proposed. We conclude (i) that the significance of viruses varies with changing trophic conditions and (ii) that viral activity may play a significant role in food web structure under changing trophic conditions.     

Increased Diversity of Predacious <Emphasis Type="Italic">Bdellovibrio</Emphasis>-Like Organisms (BLOs) as a Function of Eutrophication in Kumaon Lakes of India

Predation by Bdellovibrio-like organisms (BLOs) results in bacterial community succession in aquatic ecosystems. The effects of nutrient loading on the distribution and phylogeny of BLOs remain largely unknown. To this end, we present our findings on BLO diversity from four north-Indian lakes that are variable in their trophic status; Nainital is eutrophic, both, Bhimtal and Naukuchiatal are mesotrophic and Sattal remains oligotrophic, respectively. Initially, total heterotrophic bacteria and BLOs were quantified by most probable number (MPN) analyses using Pseudomonas putida and Escherichia coli as prey bacteria. Total bacterial numbers were at least two-logs higher in the eutrophic lake samples compared with oligotrophic lake. Similarly, BLO numbers were approximately 39-fold higher using Pseudomonas sp., which is likely the preferred prey within these lakes. Conversely, significant differences were not observed between mesotrophic and oligotrophic BLO numbers when E. coli was used as the prey. PCR-RFLP of small subunit rDNA (SSU rDNA) of BLOs, followed by cloning, sequencing, and taxonomic categorization revealed distinct differences such that, eutrophic lake consisted of higher BLO diversity compared with mesotrophic and oligotrophic lake, most likely due to both, higher numbers and availability of a diverse population of prey bacteria resulting from nutrient loading in this ecosystem.     

Viruses in the plankton of freshwater and saline Antarctic lakes

1. Virus‐like particle (VLP) abundances in nine freshwater to saline lakes in the Vestfold Hills, Eastern Antarctica (68° S) were determined in December 1999. In the ultra‐oligotrophic to oligotrophic freshwater lakes, VLP abundances ranged from 1.01 to 3.28 × 10⁶ mL^–1 in the top 6 m of the water column. In the saline lakes the range was between 6.76 and 36.5 × 10⁶ mL^–1. The lowest value was found in meromictic Ace Lake and the highest value in hypersaline Lake Williams. Virus to bacteria ratios (VBR) were lowest in the freshwater lakes and highest in the saline lakes, with a maximum of 23.4 in the former and 50.3 in the latter. 2. A range of morphologies among VLP was observed, including phages with short (Podoviridae) and long tails, icosahedric viruses of up to 300 nm and star‐like particles of about 80 nm diameter. 3. In these microbially dominated ecosystems there was no correlation between VLP and either bacterial numbers or chlorophyll a. There was a significant correlation between VLP abundances and dissolved organic carbon concentration (r=0.845, P < 0.01). 4. The data suggested that viruses probably attack a spectrum of bacteria and protozoan species. Virus‐like particle numbers in the freshwater lakes were lower than values reported for lower latitude systems. Those in the saline lakes were comparable with abundances reported from other Antarctic lakes, and were higher than most values published for lower latitude lakes and many marine systems. Across the salinity spectrum from freshwater through brackish to hypersaline, VLP concentrations increased roughly in relation to increasing trophy. 5. Given that Antarctic lakes have a plankton almost entirely made up of bacteria and protists, and that VLP abundances are high, it is likely that viruses play a pivotal role in carbon cycling in these extreme ecosystems.     

Climatic and anthropogenic effects on the sedimentation and geochemistry of Lakes Bourget,Annecy and Léman

During the last glaciation, the Rhone glacier extended as far as the region of Lyon and covered Lakes Bourget, Annecy and Léman. Glacial retreat successively freed Lakes Bourget and Annecy, it reached Geneva around 14 000 B.P. and the head of Lake Léman at about 12 000 B.P. Deposits situated between Nyon and the foot of the Jura provide a complete palynological zonation and serve as a type section for the lake core studies. A palynological horizon (about 1 800 A.D.) has been found in Lake Bourget and wood has been dated at 3 230 ± 65 B.P. (¹⁴C) in a core from Lake Léman. Three 6 m cores from each lake were sampled together with many 0.5 m cores from Léman. The short Lake Léman cores have been dated by¹³⁷Cs and provide a precise indication of changes in the basin during recent decades. Holocene climatic variations are evident in the Léman cores and are reflected by concentrations of carbonate and organic carbon which increase at the end of the Alleröd and reach a maximum in the Atlantic Period (climatic optimum). Human settlement on the shores of Lake Annecy is shown by increases in heavy metals; increased sedimentation rates suggest settlement at the beginning of the Christian era. All the cores show increases in the organic matter and nutrients; from the turn of the century in the Léman, and later for the other lakes where eutrophication started only a few decades ago. NAI-P shows a very recent increase, around 1965 in the Leman. Heavy metals (Hg, Cd and Pb) show increases at about 30 years ago in Annecy, at about the turn of the century in Lake Léman and in the mid-19th century in Lake Bourget.

     

Flow cytometric analysis of bacteria- and virus-like particles in lake sediments

Flow cytometry (FCM) was successfully used to analyze freshwater bacteria and viruses in lake sediments after relatively simple sample treatment and optimization of dilution/fixation/staining procedures. Biological particles from Lakes Geneva and Bourget were first separated from the sediments by using both Sodium Pyrophosphate (0.01 M final concentration) and Polyoxyethylene-Sorbitan Monooleate (10% final concentration) and sonicating for 3 min in a water bath. The best results (based on FCM signature and the highest virus and bacterial yields from the sediments) were obtained by formaldehyde fixation carried out within less than one hour (2% final concentration, vs. no fixation or using glutaraldehyde at different concentrations), SYBR-Green II staining (x1/20,000 stock solution concentration, vs. use of SYBR-Gold and SYBR-Green I dyes at different concentrations). There was a considerable loss of particles after only a few days of storage at either 4 or -22 degrees C. For FCM analysis, the samples were diluted in Tris-EDTA buffer (pH 8) and heated for 10 min at 75 degrees C after incubating for 5 min in the dark. The bacterial and viral counts paralleled those obtained using epifluorescence microscopy (EFM), but EFM always gave lower counts than FCM. Analysis of the distribution of the viruses in the water column and in the sediments of Lakes Bourget revealed a marked gradient, with larger quantities in the top layer of the sediment than in the water above it. These results are discussed, as well as the possible novel application of flow cytometry in the study of aquatic viral ecology.     

Local human pressures influence gene flow in a hybridizing Daphnia species complex

Anthropogenic environmental changes are considered critical drivers of the genetic structure of populations and communities through, for example, the facilitation of introgressive hybridization between syntopic species. However, the mechanisms by which environmental perturbations trigger changes in the genetic structure of populations and communities, such as the processes that determine the directionality of hybridization and patterns of mitochondrial introgression over many generations, remain largely unexplored. In this study, the changes in genetic structure of hybridizing members of the Daphnia longispina species complex were reconstructed over the last 100 years for three large temperate lakes under strong anthropogenic pressures via palaeogenetic analyses of resting egg banks. Drastic changes in the genetic structure of the Daphnia community, associated with hybridization events between D. longispina and D. galeata and subsequent introgression, were detected in Lakes Geneva and Bourget. In Lake Bourget, these changes were induced by the successful establishment of D. galeata with rising phosphorus levels and reinforced by the sensitivity of D. longispina to fish predation pressure. In Lake Geneva, the pattern of hybridization during eutrophication is more likely a function of the original taxonomic composition of the species complex in this lake. Lakes seem to require at least a meso‐oligotrophic status to allow D. galeata populations to establish and accordingly no D. galeata genotypes were found in the egg bank of oligotrophic Lake Annecy. In contrast to the generally assumed pattern of unidirectional hybridization in this species complex, bidirectional hybridization was recorded in Lakes Geneva and Bourget. Our results also demonstrate complex genetic trajectories within this species complex and highlight the irreversibility of changes in the genotypic architecture of populations driven by local human pressures. Finally, we show that extensive hybridization and introgression do not necessarily result in a large and homogenous hybrid swarm.     

Relation between nematode communities and trophic state in southern Swedish lakes

The aim of this study was to examine whether littoral nematode community patterns are shaped by lake trophic state. It was hypothesized that trophic level is associated negatively with the proportion of omnivores and positively with the percentages of bacterial feeders, but not at all with the diversity, abundance, and biomass of freshwater nematodes. Sediment samples were taken at littoral sites of eight southern Swedish lakes of different trophy in spring and autumn 2007. Trophic level was found to strongly influence species richness, as oligotrophic and mesotrophic lakes supported the greatest species numbers, whereas nematode abundance, biomass, and Shannon index were unaffected. Furthermore, our results indicated effects on the nematode community’s trophic structure, with a larger proportion of predatory nematodes in oligotrophic and mesotrophic lakes but no differences in the other feeding types (bacteria, algae and suction feeders, omnivorous species). Multivariate analysis indicated a shift in species compositions along the threshold from mesotrophic to eutrophic conditions, with the presence of Tobrilus gracilis, Monhystera paludicola, Brevitobrilus stefanskii, and Ethmolaimus pratensis related to the latter. Nematode communities in oligotrophic and mesotrophic lakes were characterized by a similar species composition, with pronounced occurrences of Eumonhystera longicaudatula, Semitobrilus cf. pellucidus, Prodesmodora circulata, and Rhabdolaimus terrestris. Overall, the results suggested that lake trophic state is a major factor structuring littoral nematode communities, although intra-lake variations might be of importance as well.     

Short-term variations in abundances and potential activities of viruses,bacteria and nanoprotists in Lake Bourget

Samples were collected at four depths every 6 h over a 42-h period during two contrasting seasons (June vs. December) from Lake Bourget, France, for evidence of circadian fluctuations in the concentrations and potential activities of viruses, prokaryotes and protists in relation to environmental conditions: temperature, chlorophyll a and dissolved organic carbon (DOC) concentrations. Considerable vertical and temporal fluctuations were observed for all variables. Circadian variations were noted for DOC and chlorophyll a concentrations. Despite the external abiotic forcing (light, water movements), the fluctuations of microbial variables (including viruses) in most cases were apparently linked to biotic factors and interactions. Standing stocks and activities, as well as the number and levels of correlations among the microbial components, were, surprisingly, higher in winter than in summer. We speculate that this was because trophic interactions prevailed over the seasonal forcing (i.e. temperature) in shaping the observed differences.     

Distribution of picocyanobacteria and virioplankton in mesotrophic and eutrophic reservoirs: The role of viruses in mortality of picocyanobacteria

The quantitative distribution and relationships of picocyanobacteria and viruses were studied in the mesotrophic Sheksna and mesoeutrophic Rybinsk reservoirs belonging to the system of the Volga-Baltic Waterway. The share of picocyanobacteria in the sum biomass and production of phytoplankton decreased along with increase in the productivity in the waterbody. However, the abundance of virioplankton increased in such conditions. The extent of infection by phages and virus-induced mortality of picocyanobacteria were two times higher on average in the mesoeutrophic reservoir compared to the mesotrophic one: 14 and 7% of the number of picocyanobacteria, 21 and 11% of daily production of picocyanobacteria, respectively. Mortality in picocyanobacteria resulted from lysis by viruses increased (up to 52% of daily production of picocyanobacteria) at the local parts where water was blooming with phytoplankton and at the parts affected by communal pollution.     

Unexpected importance of potential parasites in the composition of the freshwater small-eukaryote community

The diversity of small eukaryotes (0.2 to 5 mum) in a mesotrophic lake (Lake Bourget) was investigated using 18S rRNA gene library construction and fluorescent in situ hybridization coupled with tyramide signal amplification (TSA-FISH). Samples collected from the epilimnion on two dates were used to extend a data set previously obtained using similar approaches for lakes with a range of trophic types. A high level of diversity was recorded for this system with intermediate trophic status, and the main sequences from Lake Bourget were affiliated with ciliates (maximum, 19% of the operational taxonomic units [OTUs]), cryptophytes (33%), stramenopiles (13.2%), and cercozoa (9%). Although the comparison of TSA-FISH results and clone libraries suggested that the level of Chlorophyceae may have been underestimated using PCR with 18S rRNA primers, heterotrophic organisms dominated the small-eukaryote assemblage. We found that a large fraction of the sequences belonged to potential parasites of freshwater phytoplankton, including sequences affiliated with fungi and Perkinsozoa. On average, these sequences represented 30% of the OTUs (40% of the clones) obtained for each of two dates for Lake Bourget. Our results provide information on lacustrine small-eukaryote diversity and structure, adding to the phylogenetic data available for lakes with various trophic types.     

Abundance and distribution of picoplankton in tropical, oligotrophic Lake Kivu, eastern Africa

1. We used flow cytometry to characterize freshwater photosynthetic picoplankton (PPP) and heterotrophic bacteria (HB) in Lake Kivu, one of the East‐African great lakes. Throughout three cruises run in different seasons, covering the four major basins, phycoerythrin‐rich cells dominated the PPP. Heterotrophic bacteria and PPP cell numbers were always high and spatial variations were modest. This represents an important difference from temperate and high latitude lakes that show high fluctuations in cell abundance over an annual cycle. 2. Three populations of picocyanobacteria were identified: one corresponded to single‐cells (identified as Synechococcus by epifluorescence microscopy, molecular methods and pigment content), and the two other that most probably correspond to two and four celled colonies of the same taxon. The proportion of these two subpopulations was greater under stratified conditions, with stronger nutrient limitation. 3. High PPP concentrations (c. 10⁵ cell mL^?1) relative to HB (c. 10⁶ cell mL^?1) were always found. Lake Kivu supports relatively less bacteria than phytoplankton biomass than temperate systems, probably as a consequence of factors such as temperature, oligotrophy, nutrient limitation and trophic structure. 4. A review of PPP concentration across aquatic systems suggests that the abundance of Synechococcus‐like cyanobacteria in large, oligotrophic, tropical lakes is very high. 5. Photosynthetic picoplankton cell abundances in the oligotrophic tropical lakes Kivu and Tanganyika are comparable to those of eutrophic temperate lakes. This apparently contradicts the view that PPP abundance increases with increasing eutrophy. More data on PPP in tropical lakes are needed to explore further this particular pattern.     

Relationship between bacteria, phytoplankton and heterotrophic nanoflagellates along the trophic gradient

Bacterial and heterotrophic nanoflagellates (HNF) abundance, as well as bacterial production and chlorophylla levels, were measured at five sites extending from the coastal zone toward the open Adriatic in the period from March to October 1995. The investigated areas were grouped into trophic categories according to concentrations of chlorophylla. All the biotic-para-meters increased along the trophic gradient, leading to eutrophy, but they did not increase at the same rate. The bacterial biomass: phytoplankton biomass (BB: chla) ratio decreased from about 10 in the very oligotrophic area to 0.8 at the eutrophic site. In contrast, the bacterial abundance: HNF abundance ratio (B: HNF) increased from 1000 bacteria per 1 flagellate in the oligotrophic system to 1700 bacteria flagellate⁴ in the eutrophic area. Decreasing BB: chla and increasing B: HNF ratios along the trophic gradient might reflect the different structures of the microbial food web. Relationships between bacterial abundance and production, and chla and HNF showed that bacterial abundance along the trophic gradient was regulated by the interplay between nutrient supply and grazing pressure. But in the oligotrophic system, bacterial abundance was more closely related to bacterial production and chla than in the eutrophic system, suggesting stronger control of bacterial abundance by substrate supply. On the other hand, the coupling between bacteria and HNF, and uncoupling between bacterial abundance and production in the eutrophic system, showed that the importance of bacteriovory increased in richer systems.     

Picophytoplankton, nanophytoplankton, heterotrohpic bacteria and viruses in the Changjiang Estuary and adjacent coastal waters

Flow cytometry (FCM) was used to examine the abundances anddistributions of different picophytoplankton groups (i.e. Synechococcus,Prochlorococcus and picoeukaryotes), nanophytoplankton, heterotrophicbacteria and viruses were examined in the Changjiang Estuary,China and adjacent coastal waters during autumn 2004. Watertemperature and light availability were found to be criticalfactors for picophytoplankton growth. Positive correlationswere found between picophytoplankton, heterotrophic bacteriaand viruses, and a seaward-increasing trend in the V-I (thegroup yielding high green fluorescence according to FCM) populationwithin viruses was detected. The importance of nanophytoplanktonis progressively usurped by picophytoplankton with increasingdistance offshore. Picoeukaryotes are the most successful groupamong picophytoplankton in near-shore eutrophic waters, whereasProchlorococcus surpasses other groups within the pico- andnanophytoplankton community in offshore oligotrophic regionsof the East China Sea Shelf.