Heterotrophic microorganisms and viruses in the water of the Gorky reservoir during a period of anomalously high water temperature

During the anomalously hot summer of 2010, the water temperature in the Gorky reservoir reached 27–33°C. Pronounced cyanobacterial blooms occurred in the limnetic part of the reservoir. The average values for bacterioplankton abundance (11.58 ± 1.25 × 10⁶ cell/mL), biomass (886 ± 96 mg/m³), and production [169 ± 32 mg C/(m³ day)] were twice as high as in the year with temperatures comparable to long-term average values. These parameters were higher in the limnetic part than in the river one. The abundance (4.86 ± 0.75 × 10³ cell/mL) and biomass (138 ± 9 mg/m³) of heterotrophic nanoflagellates were 2.3 and 1.7 times higher, respectively, than in years with regular temperature regimes. The average number of plank-tonic viral particles (N _v) in 2010 was 48.89 ± 9.54 × 10⁶ particles/mL, while virus-induced bacterial mortality (VMB) accounted for 26.9 ± 4.6% of the bacterial production. The N _v and VMB values in the limnetic part of the reservoir were, respectively, 1.5 and 1.8 times higher than in the river one.     

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.     

Trophic relationships between planktonic bacteria,heterotrophic nanoflagellates and viruses in a mesoeutrophic reservoir

The abundance, biomass, and production (Р _В) of bacrerioplankton; the taxonomic composition, abundance, biomass of heterotrophic nanoflagellates (HNF) and the rate of consumption of bacteria by HNFs; and the abundance of virioplankton, frequency of visibly infected bacterial cells, virus-induced mortality of bacterioplankton, and viral production were estimated in the mesoeutrophic Rybinsk Reservoir. The rate of bacterial mortality due to viral lysis (7.8–34.1%, on average 17.2 ± 2.0% of daily Р _В) was lower than the consumption of bacteria by the HNF community (15.4–61.3%, on average 32.0 ± 4.2% of daily Р _В). While consuming bacteria, HNFs simultaneously absorbed a significant number of viruses residing on the surface and inside the bacterial cells.     

Microbial plankton across Drake Passage

We determined biomass and activity of microbial plankton across the Polar Front (PF) in Drake Passage during January 1994. Temperature was around 0°C south and between 3 and 5°C north of the PF. Both biomass and activities of microorganisms were significantly lower in the Antarctic waters south of the PF than in the sub-Antarctic waters north of it. Thus, values of chlorophyll a, integrated between 0 and 200 m, reached 150 mgm⁻² north, but only 25 mg m⁻² south of the PF. Likewise, bacteria varied between 10¹⁴ and 4×10¹³ cells m⁻². However, the abundance of heterotrophic nanoflagellates was extremely low throughout Drake Passage (around 3×10¹⁰ cells m⁻²). Bacterial doubling times were long (mean of 25 days). Bacterivory was estimated from the abundance of predators and prey and from temperature. The grazing impact on bacterioplankton biomass was insignificant (less that 0.05% per day) and low on bacterial heterotrophic production (15% per day). Neither biomass nor the activities of microorganisms were found to increase at the PF. The microbial food web was uncoupled and the bacteria did not seem to be controlled by predation.     

Distribution of viruses in the water column of the ice-covered Rybinsk Reservoir and their contribution to heterotrophic bacteria mortality

Virioplankton and bacterioplankton abundance has been determined in the pelagic and littoral zones of the Rybinsk Reservoir during the ice-covered period. The role of viruses in heterotrophic bacterioplankton infection and mortality is assessed. At water temperatures between 0.3 and 0.9°C, the number of planktonic virus particles and planktonic bacteria varies from 37.1 × 10⁶ to 84.1 × 10⁶ particles/mL, (57.3 ± 2.1) × 10⁶ particles/mL on average and from 2.50 × 10⁶ to 6.11 × 10⁶ cells/mL, (3.66 ± 0.16) × 10⁶ cells/mL on average, respectively. The ratio of the virus number to the bacteria number varies from 8.8 to 27.9, being 16.5 ± 0.7 on average. Visually infected cells comprise 0.3–0.5% (1.5 ± 0.2% on average) of the total number of bacterioplankton. Infected bacterial cells contain from 5 to 107 (17 ± 4 on average) mature virus particles. The average virus-induced mortality of bacteria accounts for 13.0 ± 1.9% (variations range from 2 to 55%) of the daily bacterial production, indicating that viruses play an important role in the regulation of bacterioplankton production and abundance in the Rybinsk Reservoir during the ice-covered period.     

Heterotrophic nanoflagellates in water column and bottom sediments of the Rybinsk Reservoir: Species composition,abundance, biomass and their grazing impact on bacteria

Abundance, biomass, and taxonomic composition of heterotrophic nanoflagellates (HNFs) have been determined in the water column and bottom sediments of the large lowland meso-eutrophic reservoir (Rybinsk Reservoir, Upper Volga) in summer. The role of HNFs in the consumption of the bacterial production is estimated. In the reservoir, 55 species from 15 large taxa, including 35 species from the plankton, are identified and 45 species are from benthos samples. The orders Kinetoplastida, Choanomonada, and Chrysomonadida are distinguished by the highest species diversity. Abundance and biomass of HNFs in the water column average 991 ± 326 cells/mL and 41.4 ± 14.1 mg/m³, while in the bottom sediments they are (236 ± 61) × 10³ cells/mL and 10.7 ± 4.0 μg/mL, respectively. The biomass of HNFs average 11.2% of the bacterial biomass in the water column and only 0.8% of that in the sediments. Flagellates are found to be a major factor which control the development of bacterioplankton grazing, on average, 32.3% of its daily production, whereas their impact on bacteriobenthos is insignificant, as they consume, on average, only 2.0% of its production.     

Structure and functioning of the microbial loop in a boreal reservoir

The abundance and biomass of the main components of the microbial plankton food web (“microbial loop”)—heterotrophic bacteria, phototrophic picoplankton and nanoplankton, heterotrophic nanoflagellates, ciliates and viruses, production of phytoplankton and bacterioplankton, bacterivory of nanoflagellates, bacterial lysis by viruses, and the species composition of protists—have been determined in summer time in the Sheksna Reservoir (the Upper Volga basin). A total of 34 species of heterotrophic nanoflagellates from 15 taxa and 15 species of ciliates from 4 classes are identified. In different parts of the reservoir, the biomass of the microbial community varies from 26.2 to 64.3% (on average 45.5%) of the total plankton biomass. Heterotrophic bacteria are the main component of the microbial community, averaging 63.9% of the total microbial biomass. They are the second (after the phytoplankton) component of the plankton and contribute on average 28.6% to the plankton biomass. The high ratio of the production of heterotrophic bacteria to the production of phytoplankton indicates the important role of bacteria, which transfer carbon of allochthonous dissolved organic substances to a food web of the reservoir.     

Temperature regulation of marine heterotrophic prokaryotes increases latitudinally as a breach between bottom‐up and top‐down controls

Planktonic heterotrophic prokaryotes make up the largest living biomass and process most organic matter in the ocean. Determining when and where the biomass and activity of heterotrophic prokaryotes are controlled by resource availability (bottom‐up), predation and viral lysis (top‐down) or temperature will help in future carbon cycling predictions. We conducted an extensive survey across subtropical and tropical waters of the Atlantic, Indian and Pacific Oceans during the Malaspina 2010 Global Circumnavigation Expedition and assessed indices for these three types of controls at 109 stations (mostly from the surface to 4,000 m depth). Temperature control was approached by the apparent activation energy in eV (ranging from 0.46 to 3.41), bottom‐up control by the slope of the log‐log relationship between biomass and production rate (ranging from ?0.12 to 1.09) and top‐down control by an index that considers the relative abundances of heterotrophic nanoflagellates and viruses (ranging from 0.82 to 4.83). We conclude that temperature becomes dominant (i.e. activation energy >1.5 eV) within a narrow window of intermediate values of bottom‐up (0.3–0.6) and top‐down 0.8–1.2) controls. A pervasive latitudinal pattern of decreasing temperature regulation towards the Equator, regardless of the oceanic basin, suggests that the impact of global warming on marine microbes and their biogeochemical function will be more intense at higher latitudes. Our analysis predicts that 1°C ocean warming will result in increased biomass of heterotrophic prokaryoplankton only in waters with <26°C of mean annual surface temperature.     

Ciliates are the Dominant Grazers on Pico- and Nanoplankton in a Shallow,Naturally Highly Eutrophic Lake

Abundance and biomass of the microbial loop members [bacteria, heterotrophic nanoflagellates (HNF), and ciliates] were seasonally measured in the naturally eutrophic and shallow (2.8 mean depth) Lake Võrtsjärv, which has a large open surface area (average 270 km²) and highly turbid water (Secchi depth <1 m). Grazing rates (filter feeding rates) on 0.5-, 3-, and 6-μm-diameter particles were measured to estimate pico- and nanoplankton grazing (filter feeding) by micro- and metazooplankton. Among grazers, HNF had a low abundance (<50 cells mL^?1) and, due to their low specific filtering rates, they only grazed a minor fraction of the bacterioplankton (≤4.2% of total grazing). Ciliates were relatively abundant (≤158 cells mL^?1) and, considering their high specific feeding rates, were able to graze more than 100% of the bacterial biomass production in the open part of the lake, whereas the average daily grazing accounted for 9.3% of the bacterial standing stock. Ciliates were potentially important grazers of nanoplanktonic organisms (on average, approximately 20% of the standing stock of 3-μm-size particles was grazed daily). Metazooplankton grazed a minor part of the bacterioplankton, accounting for only 0.1% of standing stock of bacteria. Grazing on nanoplankton (3–6 μm) by metazooplankton was higher (0.4% of standing stock). The hypothesis is proposed that ciliates dominate due to a lack of top–down regulation by predators, and HNF have a low abundance due to strong grazing pressure by ciliates.     

Top‐down effects on pelagic heterotrophic nanoflagellates (HNF) in a large river (River Rhine): do losses to the benthos play a role?

1. The abundance, production and control of pelagic heterotrophic nanoflagellates (HNF) in the Lower River Rhine (Germany) were investigated. Field samples (live‐counting technique) were taken at least every 2 weeks at Cologne (km 685) over a period of 20 months. In addition, Lagrangian sampling was carried out 160 km downstream of Cologne (Kalkar–Grieth, km 845) over a period of 12 months. Potential HNF growth rates and loss rates caused by planktonic predators were estimated in the laboratory (size fractionation experiments) and compared with the changes in HNF‐density in a water parcel flowing downstream. 2. Mean abundance (±SD) ranged from 7 ± 6 to 4890 ± 560 individuals mL^–1 and was positively correlated with discharge. Heterotrophic nanoflagellates abundance increased up to 30‐fold during flood events, whereas there were only minor changes in the taxonomic composition. 3. HNF growth rate ranged from 0.16 ± 0.12 to 1.98 ± 0.10 day^–1. Between 0 and 95% (mean: 32%) of the gross production was lost to planktonic predators; a larger portion between 0 and 195% (mean: 64%) of the HNF gross production was lost by other means. 4. There may be an important role for benthic predators in the control of pelagic HNF. First, production of HNF was high and grazing losses to planktonic predators low at times when HNF abundance was low. Secondly, high in situ loss rates (not explicable by planktonic predators), which were positively related to temperature, indicate the importance of biotic interactions. Thirdly, the dependence of HNF abundance on discharge indicated a decrease grazing intensity with rising water levels (increase in water volume/colonised river bed ratio). 5. The impact of discharge on planktonic HNF mediated by the grazing impact of benthic predators was modelled, showing a good fit with the field data.     

Structural and functional characteristics of planktonic microbial communities of the Rybinsk Reservoir in the impact area of a large industrial center

In sites of the Rybinsk Reservoir with different levels of contamination by the municipal and industrial wastewater from the city of Cherepovets, the primary production of phytoplankton and the quantitative distribution of bacterioplankton, heterotrophic nanoflagellates, and viruses, as well as the content of finely disperse detritus, were investigated. The structural and functional characteristics of microbial communities form the contaminated sites were determined. Heterotrophic nanoflagellates were shown to play a major role in the consumption of 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.     

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.     

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.     

Abundance and Biomass Responses of Microbial Food Web Components to Hydrology and Environmental Gradients within a Floodplain of the River Danube

This study investigated the relationships of time-dependent hydrological variability and selected microbial food web components. Samples were collected monthly from the Kopa?ki Rit floodplain in Croatia, over a period of 19 months, for analysis of bacterioplankton abundance, cell size and biomass; abundance of heterotrophic nanoflagellates and nanophytoplankton; and concentration of chlorophyll a. Similar hydrological variability at different times of the year enabled partition of seasonal effects from hydrological changes on microbial community properties. The results suggested that, unlike some other studies investigating sites with different connectivity, bacterioplankton abundance, and phytoplankton abundance and biomass increased during lentic conditions. At increasing water level, nanophytoplankton showed lower sensitivity to disturbance in comparison with total phytoplankton biomass: this could prolong autotrophic conditions within the floodplain. Bacterioplankton biomass, unlike phytoplankton, was not impacted by hydrology. The bacterial biomass less affected by hydrological changes can be an important additional food component for the floodplain food web. The results also suggested a mechanism controlling bacterial cell size independent of hydrology, as bacterial cell size was significantly decreased as nanoflagellate abundance increased. Hydrology, regardless of seasonal sucession, has the potential to structure microbial food webs, supporting microbial development during lentic conditions. Conversely, other components appear unaffected by hydrology or may be more strongly controlled by biotic interactions. This research, therefore, adds to understanding on microbial food web interactions in the context of flood and flow pulses in river-floodplain ecosystems.     

Production of Heterotrophic Bacterioplankton in a Large Meso-Eutrophic Reservoir: The Importance of Extracellular Organic Carbon Released by Phytoplankton

The spatial distribution and seasonal dynamics of heterotrophic bacterioplankton production have been studied and the value of autochthonous sources of substrates for bacteria has been evaluated in the pelagic zone of the meso-eutrophic Rybinsk Reservoir (the Upper Volga). During the vegetation period, the bacterial production ranges from 32 to 1352 (on average 444 ± 44) mg C/(m² × day). The total input of organic carbon from the processes of extracellular production of phytoplankton, viral lysis of prokaryotic cells, and feeding of protists provides 9–64% (32 ± 3% on average) of the daily carbon demand for heterotrophic bacterioplankton.     

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.     

Biomass, production and heterotrophic activity of bacterioplankton in the Great Astrolabe Reef lagoon (Fiji)

 Biomass, production and heterotrophic activity of bacterioplankton were determined for two weeks in the Great Astrolabe Reef lagoon, Fiji. Bacterial and Bacterial activities were distributed homogeneously throughout the water column (20 to 40 m deep) and varied little from site to site inside the lagoon. Bacterioplankton biomass and production also varied little over a diel period with coefficients of variation of 9 and 22%, respectively. On average, over the whole study, bacterial abundance was 0.77×10⁹ cells l^-1 and bacterial production averaged 0.36 μg-at. C l^-1 d^-1. Bacterial abundance and production were greater in the lagoon than in oceanic waters. Attachment to particles seems to provide an advantage for bacterioplankton growth because specific growth rates for attached bacterioplankton were, on average, significantly greater than that of the free community. Growth efficiency, determined by correlating the net increase of bacterial biomass and the net decrease of dissolved organic carbon (DOC) in dilution cultures, was very low (average 6.6%). Using carbon growth efficiency and bacterial production rates, heterotrophic activity was estimated to average 5.4 μg-at. C l^-1 d^-1. The turn-over rate of DOC (average 114 μg-at. C l^-1) due to bacterial consumption was estimated to be 0.048 d^-1 during the period of study. Accepted: 25 July 1998     

Nanoflagellate predation on auto- and heterotrophic picoplankton in the oligotrophic Mediterranean Sea

Dynamics of autotrophic and heterotrophic prokaryotes and theirconsumption by nanoflagellates were studied in the euphoticzone at nine stations located from the Levantine Basin (34°E)to the Balearic sea (5°E) in June 1999. Bacterial biomassconstituted the largest proportion of living biomass at allstations. Integrated bacterial production at the furthest eaststation, was sixfold lower than integrated bacterial productionat the furthest west (13 and 75 mg C m^-2 d^-1 respectively).Estimated heterotrophic nanoflagellate bacterivory accountedfor 45–87% of bacterial production. Small protists (<3µm) dominated the bacterivore assemblage and accountedfor more than 90% of the heterotrophic bacterial consumption.Our results indicated that there was no negative selection againstSynechococcus and that both picoplankton groups were grazedaccording to their standing stocks. An estimated consumptionof Synechococcus derived from food vacuole content analysisof nanoflagellates revealed that they consumed from 0.5 to 45%(mean 13%) of Synechococcus stock per day. These data are amongthe first documenting the relative grazing impact of heterotrophicnanoflagellates on bacteria and Synechococcus in situ. Assumingthat there was no selection for or against Prochlorococcus,heterotrophic nanoflagellates could ingest from 1.4 to 21% (mean6%) of Prochlorococcus stock per day. The amount of organiccarbon obtained by heterotrophic nanoflagellates from photosyntheticprokaryotes represented 27% of the total amount of carbon obtainedfrom total prokaryotes     

Heterotrophic activity of bacterioplankton along the gradients of the chlorophyll <Emphasis Type="Italic">a</Emphasis> concentration and water temperature in marine and limnetic ecosystems

The dependence of the heterotrophic activity of bacterioplankton (V, μg C L^–1 h^–1) on the concentration of chlorophyll a (Chl, μg L^–1) and the water temperature (T) was examined for lakes (37°29′–80°36′ N) and marine polar waters (69°16′–80°36′ N). It was shown that ~76% of the V variations was related to changes in Chl and T.