Abundance and biomass of heterotrophic microorganisms in Lake Tanganyika

1. This study focused on heterotrophic microorganisms in the two main basins (north and south) of Lake Tanganyika during dry and wet seasons in 2002. Bacteria (81% cocci) were abundant (2.28–5.30 × 10⁶ cells mL^?1). During the dry season, in the south basin, bacterial biomass reached a maximum of 2.27 g C m^?2 and phytoplankton biomass was 3.75 g C m^?2 (integrated over a water column of 100 m). 2. Protozoan abundance was constituted of 99% of heterotrophic nanoflagellates (HNF). Communities of flagellates and bacteria consisted of very small but numerous cells. Flagellates were often the main planktonic compartment, with a biomass of 3.42–4.43 g C m^?2. Flagellate biomass was in the same range and often higher than the total autotrophic biomass (1.60–4.72 g C m^?2). 3. Total autotrophic carbon was partly sustained by the endosymbiotic zoochlorellae Strombidium. These ciliates were present only in the euphotic zone and usually contributed most of the biomass of ciliates. 4. Total heterotrophic ciliate biomass ranged between 0.35 and 0.44 g C m^?2. In 2002, heterotrophic microorganisms consisting of bacteria, flagellates and ciliates represented a large fraction of plankton. These results support the hypothesis that the microbial food web contributes to the high productivity of Lake Tanganyika. 5. As the sole source of carbon in the pelagic zone of this large lake is phytoplankton production, planktonic heterotrophs ultimately depend on autochthonous organic carbon, most probably dissolved organic carbon (DOC) from algal excretion.     

Comparison of pelagic food webs in lakes along a trophic gradient and with seasonal aspects: influence of resource and predation

Composition and seasonal dynamics of phytoplankton, bacteria,and zooplankton (including heterotrophic flagellates, ciliates,rotifers and crustaceans) were studied in 55 lakes in NorthernGermany with different trophic status, ranging from mesotrophicto hypertrophic. Mean abundance and biomass of all groups increasedsignificantly with trophic level of the lake, but bacteria andmetazooplankton showed only a weak correlation and a slightincrease with chlorophyll concentration. Composition of phytoplanktonshowed a dominance of cyanobacteria in hypertrophic lakes, whereasthe importance of chrysophytes and dinophytes decreased withan increase in trophic status. Protozoans (heterotrophic flagellatesand ciliates) made up 24% (mesotrophic lakes) to 42% (hypertrophiclakes) of total zooplankton biomass on average, and were dominatedby ciliates (62–80% of protozoan biomass). Seasonally,protozoans can build up to 60% of zooplankton biomass in spring,when heterotrophic flagellates can contribute     

The Microbial Food Web in the Recently Flooded Sep Reservoir: Diel Fluctuations in Bacterial Biomass and Metabolic Activity in Relation to Phytoplankton and Flagellate Grazers

The spatial distribution of the bacterial biomass and production and of potential heterotrophic activity (PHA) weree measured every 4 h between 23 July (10:00 h) and 25 July (10:00 h) 1997 in a recently flooded oligo-mesotrophic reservoir (the Sep Reservoir, Puy-de-Dôme, France), in relation to temperature, the phytoplankton biomass and production, and the abundance of heterotrophic flagellates. The temperature varied slightly with time during the study, but the well-established thermal stratification agreed well with vertical distribution of the biological variables that were measured. Only the bacterial production and the PHA showed significant diel changes (t-test,p<0.05), with maxima at 18:00 h and minima at 02:00 h. A significant positive relation was found between bacterial abundance and that of heterotrophic flagellates, which, rather than being an association related to the thermal stratification of the water column, was considered to reflect a trophic relation between these two communities. A carbon balance analysis suggested that at least 30% of the C from primary production measured during the sampling period was used by bacteria, and that 42% of this secondary production, or 6% of the primary production, would be used for the development of the heterotrophic flagellates present. We conclude that the bacterioplankton forms, at least occasionally, an important source of carbon for higher trophic levels, and reject the hypothesis that bacterial production in the Sep Reservoir depends exclusively on organic matter of allochthonous origin.     

Protistan bacterivory in an oligomesotrophic lake: Importance of attached ciliates and flagellates

Seasonal and depth variations of the abundance, biomass, and bacterivory of protozoa (heterotrophic and mixotrophic flagellates and ciliates) were determined during thermal stratification in an oligomesotrophic lake (Lake Pavin, France). Maximal densities of heterotrophic flagellates (1.9 × 10³ cells ml^–1) and ciliates (6.1 cells ml^–1) were found in the metalimnion. Pigmented flagellates dominated the flagellate biomass in the euphotic zone. Community composition of ciliated protists varied greatly with depth, and both the abundance and biomass of ciliates was dominated by oligotrichs. Heterotrophic flagellates dominated grazing, accounting for 84% of total protistan bacterivory. Maximal grazing impact of heterotrophic flagellates was 18.9 × 10⁶ bacteria 1^–1h^–1. On average, 62% of nonpigmented flagellates were found to ingest particles. Ciliates and mixotrophic flagellates averaged 13% and 3% of protistan bacterivory, respectively. Attached protozoa (ciliates and flagellates) were found to colonize the diatom Asterionella formosa. Attached bacterivores had higher ingestion rates than free bacterivorous protozoa and may account for 66% of total protozoa bacterivory. Our results indicated that even in low numbers, epibiotic protozoa may have a major grazing impact on free bacteria. Correspondence: C. Amblard.     

Pico- and Nanoplankton in Aquatic Ecosystems in the Valley of the Lakes and Great Lakes Depression (Mongolia)

The abundance and biomass of bacterioplankton, phototrophic picoplankton, and heterotrophic nanoflagellates has been determined in lakes, rivers, and reservoirs located in the Valley of the Lakes and Great Lakes Depression (Mongolia). The species richness of the heterotrophic flagellates and their consumption of bacteria are estimated. Pico- and nanoplankton are the most abundant in shallow mineral lakes Orog and Tatsyn and in the freshwater Durgun Reservoir. Heterotrophic nanoflagellates consume 26–92% (on average 66%) of the daily bacterioplankton production. Thus, flagellates are important in the transfer of bacterial carbon to the higher levels of planktonic trophic webs. A total of 30 species and their forms of heterotrophic flagellates from 14 large taxa are identified. The highest species diversity of these protists are found in the Durgun and Taishyr reservoirs.     

Channeling of bacterioplanktonic production toward phagotrophic flagellates and ciliates under different seasonal conditions in a river

The objective of this study was to analyze the flux of biomass through the communities of bacteria and phagotrophic protists in the cold and warm conditions occurring seasonally in Butrón River. Bacterial and heterotrophic protistan (flagellate and ciliate) abundance was determined by epifluorescence direct counts; protistan grazing on planktonic bacteria was measured from fluorescently labeled bacteria uptake rates; and the estimate of bacterial secondary production was obtained from [³H]thymidine incorporation rates. The abundance of bacterial, flagellate, and ciliate communities was similar during cold and warm situations. However, we observed that estimates of dynamic parameters, i.e., secondary bacterial production and protistan grazing, in both situations were noticeably different. In the warm situation, grazing rates of flagellates and ciliates (bacteria per protist per hour) were, respectively, 7 times and 18 times higher than those determined in the cold situation, and the grazing rates of the protistan communities (bacteria per protists present in 1 ml of water per hour) increased up to 5 times in the case of flagellates and 42 times in the case of ciliates. Estimates of bacterial secondary production were also higher during the warm situation, showing a ninefold increase. The percentage of bacterial production preyed upon by flagellates or ciliates was not significantly different between the two conditions. These results showed that in the different conditions of a system, the flux of biomass between the trophic levels may be quite different although this process may not be reflected in the abundance of each community of bacteria, flagellates, and ciliates. Offprint requests to: J. Iriberri.     

Weak Coupling between Heterotrophic Nanoflagellates and Bacteria in a Eutrophic Freshwater Environment

In a study on the dynamics and trophic role of the heterotrophic nanoflagellate (HNAN) assemblage in the microbial food web of a eutrophic oxbow lake abundances, biomass, and production rates of HNAN and their potential prey organisms, namely heterotrophic bacteria and autotrophic picoplankton, were monitored for a period of 2 years. No coupling between HNAN abundance and biomass and the abundance and biomass of their picoplanktonic prey was observed for the investigation period. The ratio of heterotrophic bacterial to HNAN abundance ranged from 2.2 x 103 to 8.6 x 103 (mean: 4.2 x 103 +/- 1.8 x 103). HNAN carbon consumption could account for only 10% to 40% of bacterial secondary production. The lack of coupling between HNAN and their potential prey and the low HNAN abundance relative to bacterial abundance suggested (a) that HNAN grazing was an insignificant factor in the regulation of bacterial abundance and (b) that HNAN abundance was regulated by predation rather than by prey abundance. This hypothesis was supported by the fact that HNAN growth rates were high (in the range of 0.45 d-1 to 1.00 d-1 during spring and summer, yearly mean: 0.52 d-1), and only weakly correlated with prey abundance and biomass. The results indicated strong top-down control of HNAN and consequently a weak coupling of HNAN and picoplankton in the investigated eutrophic freshwater environment.     

Benthic–pelagic coupling: a comparison of the community structure of benthic and planktonic heterotrophic protists in shallow inlets of the southern Baltic

1. The taxonomic composition, abundance and biomass of heterotrophic protists (ciliates, heterotrophic flagellates (HF), rhizopods and actinopods) in the sediment and water column of shallow inlets of the Southern Baltic was studied under a variety of environmental conditions during 1996–1997. A shallow, highly eutrophic station and a deeper, less eutrophic station were compared.
2. Community biomass ranged from 0.12 to 0.34 μg C cm?3 in the water column and from 1.5 to 105 μg C cm?3 in the sediment. Heterotrophic protists dominated zooplankton biomass at both stations (73% and 84% mean contribution), while they were of minor importance within the zoobenthos. Expressed per unit area, benthic biomass contributed a significant part (44% and 49%) to the total heterotrophic protistan community at both stations.
3. Although the methodology for counting ciliates and HF was focussed on a high taxonomic resolution, the results reveal some general trends in the distribution of heterotrophic protists: protozooplankton biomass was dominated by flagellates (80% mean biomass contribution) at the shallow station and by ciliates (73% mean biomass contribution) at the deep station. In the benthos at both stations, ciliates were the dominant protozoans, followed by the hitherto little‐studied rhizopods (25% and 35% mean biomass contribution) and flagellates.
4. The degree of benthic–pelagic coupling differed between taxonomic groups. Benthic and pelagic communities of ciliates showed little taxonomic overlap. In contrast, many heterotrophic flagellate species were found both in the benthos and in the pelagic. These benthic–pelagic species contributed significantly to the biomass of HF in the water column. The planktonic rhizopod community consisted of a subset of those species found in the benthos.
5. The abundance of benthic and pelagic protists was positively correlated at the shallow station, but taxonomic data indicate that the direct exchange between benthic and pelagic communities was only partly responsible.     

The distribution of protozoa across a trophic gradient, factors controlling their abundance and importance in the plankton food web

The relative contribution of protozoan biomass to whole planktoniccommunities (phytoplankton, picophytoplankton, bactenoplankton,protozoa and zooplankton) and factors important in controllingprotozoan abundance were investigated at two eutrophic coastalsites and two meso-oligo trophic offshore sites in the centralbasin of Lake Erie, USA, from May through August in 1993 and1994. The abundance and biomass of heterotrophic nanoflagellates(HNAN) and ciliates (and also other plankton components) weresignificantly higher at the coastal sites than at the offshoresites. HNAN dominated numerically at all sites most of the time,but the biomass of phototrophic nano flagellates (PNAN) wasas high as that of HNAN, indicating that the average size ofPNAN was larger. Percent protozoan carbon content was alwayshigher at the offshore sites than the coastal sites, due torelatively lower phyto-and zooplankton biomass at the offshoresites. The percent contribution of heterotrophic protozoans(both HNAN and ciliates) also showed the same trend. Correlationsbetween protozoan abundance and other parameters were strongerat the offshore sites than the coastal sites. When correlatingdata over the coastal to offshore transect, both HNAN and ciliateabundances were significantly correlated with total phosphorus(TP) and the abundance of bacteria, nauplii and copepods. Theseresults suggest that both bottom-up and top-down factors maybe important in controlling protozoan abundance, and suggestthat protozoans are important as a carbon link in the microbialfood web of Lake Erie. ¹Present address: Ecosystem Restoration Department, South FloridaWater Management District, West Palm Beach, FL 33416-4680, USA     

Abundance,biomass, and production of heterotrophic bacteria in a large plain reservoir during the ice-covered period

The abundance, biomass, production, and size and morphological structure of heterotrophic bacteria have been determined and their contribution to the total plankton biomass has been assessed in the Rybinsk Reservoir (the Upper Volga) during the ice-covered period. Structural and functional characteristics of bacteria, except for their cell sizes in winter, are lower than those in the growing season. However, heterotrophic bacteria have made the main contribution (89.3%) to the total biomass of plankton community in winter. During the growing season, phytoplankton is a major component of plankton (48.8%), while the bacteria average 36.5%. The total plankton biomass under the ice averages 148 mg C/m³, which is 2.6 times lower than in the period of open water. During the ice-covered period, the number of protists is small and they consume an insignificant part of the bacterial production; viral lysis is the main reason for bacterial mortality.     

Bacterial quantity and microbial reactivity in Tugur bay of the Okhotsk Sea

The paper presents the results of investigation of the total abundance and the biomass of bacterioplankton, the abundance of heterotrophic bacteria, and the activity of microbiological processes involved in the carbon cycle in the water of the Bay of Tugur of the Sea of Okhotsk. In different regions of the bay, the total abundance of bacterioplankton was found to vary from 0.51 x 10(6) to 2.54 x 10(6) cells/ml; the bacterioplankton biomass, from 8.5 to 46.5 micrograms C/l; the abundance of heterotrophic bacteria, from 0.06 x 10(3) to 2.12 x 10(3) cells/ml; the bacterial assimilation of CO2, glucose, acetate, and protein hydrolysate, from 0.8 to 6.3, from 0.11 to 1.88, from 0.07 to 0.56, and from 0.01 to 0.22 mg C/(m3 day), respectively; the degradation of organic matter ranged from 28 to 221 mg C/(m3 day); and the intensity of methane oxidation, from 0.0005 to 0.17 microliter CH4/l. The spatial pattern and the functional characteristics of bacterioplankton in the Bay of Tugur were found to be dependent on the tidal dynamics.     

Bacterial Abundance and the Activity of Microbiological Processes in the Bay of Tugur of the Sea of Okhotsk

This paper presents the results of investigation of the total abundance and the biomass of bacterioplankton, the abundance of heterotrophic bacteria, and the activity of microbiological processes involved in the carbon cycle in the water of the Bay of Tugur of the Sea of Okhotsk. In different regions of the bay, the total abundance of bacterioplankton was found to vary from 0.51 × 10⁶ to 2.54 × 10⁶ cells/ml; the bacterioplankton biomass, from 8.5 to 46.5 g C/l; the abundance of heterotrophic bacteria, from 0.06 × 10³ to 2.12 × 10³ cells/ml; the bacterial assimilation of CO₂, glucose, acetate, and protein hydrolysate, from 0.8 to 6.3, from 0.11 to 1.88, from 0.07 to 0.56, and from 0.01 to 0.22 mg C/(m³ day), respectively; the degradation of organic matter ranged from 28 to 221 mg C/(m³ day); and the intensity of methane oxidation, from 0.0005 to 0.17 l CH₄/l. The spatial pattern and the functional characteristics of bacterioplankton in the Bay of Tugur were found to be dependent on the tidal dynamics.     

The structure of the planktic microbial community in the lower reaches of the Ob River near Salekhard

We studied the distribution of the main components of planktic food web: bacteria, heterotrophic flagellates, and ciliates in the lower reaches of the Ob River near the town of Salekhard. In the early summer, the abundance, biomass, and diversity of protozoans were low; the plankton community was dominated by heterotrophic bacteria whose biomass was higher than that of phyto- and zooplankton. Downstream of Salekhard, biomass and abundance of bacterioplankton were increased due to household and industrial wastewater. The data obtained showed the importance of heterotrophic organisms in trophic dynamics and self-purification of the lower reaches of the Ob River.     

Occurrence of mixotrophic flagellates in relation to bacterioplankton production, light regime and availability of inorganic nutrients in unproductive lakes with differing humic contents

1. Field data from five unproductive Swedish lakes were used to investigate the occurrence of mixotrophic flagellates in relation to bacterioplankton, autotrophic phytoplankton, heterotrophic flagellates and abiotic environmental factors. Three different sources of data were used: (i) a 3‐year study (1995–97) of the humic Lake Örträsket, (ii) seasonal measurements from five lakes with widely varying dissolved organic carbon (DOC) concentrations, and (iii) whole lake enrichment experiments with inorganic nutrients and organic carbon. 2. Mixotrophic flagellates usually dominated over autotrophic phytoplankton in Lake Örträsket in early summer, when both bacterial production and light levels were high. Comparative data from the five lakes demonstrated that the ratio between the biomasses of mixotrophic flagellates and autotrophic phytoplankton (the M/A‐ratio) was positively correlated to bacterioplankton production, but not to the light regime. Whole lake carbon addition (white sugar) increased bacterial biomass, and production, reduced the biomass of autotrophs by a factor of 16, and increased the M/A‐ratio from 0.03 to 3.4. Collectively, the results indicate that the dominance of mixotrophs among phytoplankton was positively related to bacterioplankton production. 3. Whole lake fertilisation with nitrogen (N) and phosphorus (P) demonstrated that the obligate autotrophic phytoplankton was limited by N. N‐addition increased the biomass of the autotrophic phytoplankton but had no effect on mixotrophic flagellates or bacteria, and the M/A‐ratio decreased from 1.2 to 0.6 after N‐enrichment. Therefore, we suggest that bacteria under natural conditions, by utilising allochthonous DOC as an energy and carbon source, are able to outcompete autotrophs for available inorganic nutrients. Consequently, mixotrophic flagellates can become the dominant phytoplankters when phagotrophy permits them to use nutrients stored in bacterial biomass. 4. In Lake Örträsket, the biomass of mixotrophs was usually higher than the biomass of heterotrophs during the summer. This dominance could not be explained by higher grazing rates among the mixotrophs. Instead, ratios between mixotrophic and heterotrophic biomass (the M/H‐ratio) were positively related to light availability. Therefore, we suggest that photosynthesis can enable mixotrophic flagellates to outcompete heterotrophic flagellates.     

Microbial Community Structure and Dynamics in the Largest Natural French Lake (Lake Bourget)

We investigated the dynamics and diversity of heterotrophic bacteria, autotrophic and heterotrophic flagellates, and ciliates from March to July 2002 in the surface waters (0–50 m) of Lake Bourget. The heterotrophic bacteria consisted mainly of “small” cocci, but filaments (>2 μm), commonly considered to be grazing-resistant forms under increased nanoflagellate grazing, were also detected. These elongated cells mainly belonged to the Cytophaga-Flavobacterium (CF) cluster, and were most abundant during spring and early summer, when mixotrophic or heterotrophic flagellates were the main bacterial predators. The CF group strongly dominated fluorescent in situ hybridization–detected cells from March to June, whereas clear changes were observed in early summer when Beta-proteobacteria and Alpha-proteobacteria increased concomitantly with maximal protist grazing pressures. The analysis of protist community structure revealed that the flagellates consisted mainly of cryptomonad forms. The dynamics of Cryptomonas sp. and Dinobryon sp. suggested the potential importance of mixotrophs as consumers of bacteria. This point was verified by an experimental approach based on fluorescent microbeads to assess the potential grazing impact of all protist taxa in the epilimnion. From the results, three distinct periods in the functioning of the epilimnetic microbial loop were identified. In early spring, mixotrophic and heterotrophic flagellates constituted the main bacterivores, and were regulated by the availability of their resources mainly during April (phase 1). Once the “clear water phase” was established, the predation pressure of metazooplankton represented a strong top-down force on all microbial compartments. During this period only mixotrophic flagellates occasionally exerted a significant bacterivory pressure (phase 2). Finally, the early summer was characterized by the highest protozoan grazing impact and by a rapid shift in the carbon pathway transfer, with a fast change-over of the main predators contribution, i.e., mixotrophic, heterotrophic flagellates and ciliates in bacterial mortality. The high abundance of ciliates during this period was consistent with the high densities of resources (heterotrophic nanoflagellates, algae, bacteria) in deep layers containing the most chlorophyll. Bacteria, as ciliates, responded clearly to increasing phytoplankton abundance, and although bacterial grazing impact could vary largely, bacterial abundance seemed to be primarily bottom-up regulated (phase 3).     

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.     

Micro-organism activity in aggregate layers in shallow eutrophic brackish water as influenced by wind induced mixing; an experimental approach

In the shallow eutrophic waters of the Darss-Zingst-ecosystem (Baltic Sea) wind induced mixing of the water column, and resuspension of sediment is an important phenomenon. Incubation experiments (7 days) with natural water were performed in the laboratory under simulated mixing conditions in order to assess the effects on abundance, biomass and activity of bacteria, heterotrophic nanoflagellates and ciliates in the water column and upper bottom layers. Under calm mixing conditions a bottom layer developed, rich in organic aggregates. Bacterial biomass increased through increase of bacteria abundance and doubled cell volume. Consequently, biomass of flagellates and, more pronounced, of ciliates increased. The bottom layer, comprising only 15% of the incubated volume, containedca. 60% of the bacterial biomass and ca. 55% of production, 75 and 95% of flagellate biomass, and ca. 70% of ciliate biomass. It is concluded that bottom layers rich in organic aggregates, although not being present permanently, play an important role in the shallow Darss-Zingst-ecosystem.     

Differences of the protozoan biomass and grazing during spring and summer in the Indian sector of the Southern Ocean

The dynamics of protozoa were investigated during two cruises in the Indian sector of the Southern Ocean: the early spring ANTARES 3 cruise (28 September to 8 November 1995) and the late summer ANTARES 2 cruise (6 February to 8 March 1994). Biomass and feeding activity of protozoa were measured as well as the biomass of their potential prey – bacteria and phototrophic flagellates – along the 62°E meridian. The sampling grid extended from the Polar Frontal region to the Coastal and Continental Shelf Zone in late summer and to the ice edge in spring, crossing the Antarctic Divergence. Protozoan biomass, although low in absolute terms, contributed 30% and 20% to the total microbial biomass (bacteria, phytoplankton and protozoa) in early spring and late summer, respectively. Nanoprotozoa dominated the total protozoan biomass. The geographical and seasonal distribution of protozoan biomass was correlated with that of phototrophic flagellates. However, bacterial and phototrophic flagellate biomass were inversely correlated. Phototrophic flagellates dominated in the Sea Ice Zone whereas bacteria were predominant at the end of summer in the Polar Frontal region and Coastal and Continental Shelf Zone. Furthermore, bacteria were the most important component of the microbial community (57% of the total microbial biomass) in late summer. Phototrophic flagellates were ingested by both nano-and microprotozoa. In contrast, bacteria were only ingested by nanoprotozoa. Protozoa controlled up to 90% of the daily bacterial production over the period examined. The spring daily protozoan ingestion controlled more than 100% of daily phototrophic flagellate production. This control was less strong at the end of summer when protozoan grazing controlled 42% of the daily phototrophic flagellate production. Accepted: 30 October 1999     

Protozooplankton in the Weddell Sea,Antarctica: Abundance and distribution in the ice-edge zone

Summary Protozooplankton were sampled in the iceedge zone of the Weddell Sea during the austral spring of 1983 and the austral autumn of 1986. Protozooplankton biomass was dominated by flagellates and ciliates. Other protozoa and micrometazoa contributed a relatively small fraction to the heterotrophic biomass. During both cruises protozoan biomass, chlorophyll a concentrations, phytoplankton production and bacterial biomass and production were low at ice covered stations. During the spring cruise, protozooplankton, phytoplankton, and bacterioplankton reached high concentrations in a welldeveloped ice edge bloom 100 km north of the receding ice edge. During the autumn cruise, the highest concentrations of biomass were in open water well-separated from the ice edge. Integrated protozoan biomass was <12% of the biomass of phytoplankton during the spring cruise and in the autumn the percentages at some stations were >20%. Bacterial biomass exceeded protozooplankton biomass at ice covered stations but in open water stations during the fall cruise, protozooplankton biomass reached twice that of bacteria in the upper 100m of the water column. The biomass of different protozoan groups was positively correlated with primary production, chlorophyll a concentrations and bacterial production and biomass, suggesting that the protozoan abundances were largely controlled by prey availability and productivity. Population grazing rates calculated from clearance rates in the literature indicated that protozooplankton were capable of consuming significant portions of the daily phyto- and bacterioplankton production.     

Phage Bacteriolysis, Protistan Bacterivory Potential, and Bacterial Production in a Freshwater Reservoir: Coupling with Temperature

Phage abundance and infection of bacterioplankton were studied from March to November 2003 in the Sep Reservoir (Massif Central, France), together with temperature, chlorophyll, bacteria (abundance and production), and heterotrophic nanoflagellates (abundance and potential bacterivory). Virus abundance (VA) ranged from 0.6 to 13 × 10¹⁰ viruses l⁻¹, exceeding bacterial abundance (BA) approximately sixfold on average. In terms of carbon, viruses corresponded to up to 25% of bacterial biomass. A multiple regression model indicated that BA was the best predictor for VA (R² = 0.75). The frequency of infected bacteria (estimated from the percentage of visibly infected cells) varied from 1% to 32% and was best explained by a combination of temperature (R² = 0.20) and bacterial production (R² = 0.25). Viruses and flagellates contributed about equally to bacterial mortality. Both factors destroyed 55% of bacterial production, with a shift from phage bacteriolysis in early spring to protistan bacterivory in late summer. The vertical differences in most of the biological variables were not significant, contrasting with the seasonal differences (i.e., spring vs. summer-autumn). All biological variables under study were indeed significantly coupled to temperature. We regarded this to be the consequence of the enhanced discharge of the reservoir in 2003 (compared to previous years). This substantially weakened the stability and the thermal inertia of the water column, thereby establishing temperature as a stronger forcing factor in setting the conditions for optimal metabolic activity of microbial communities.