Seasonal variations in PCR-DGGE fingerprinted viruses infecting phytoplankton in large and deep peri-alpine lakes

Double-stranded DNA viruses infecting eukaryotic algae (e.g., phycodnaviruses) and cyanobacteria (e.g., cyanophages) are now recognized as widespread and ubiquitous in aquatic environments. However, both the diversity and functional roles of these viruses in fresh waters are still poorly understood. We conducted a year-long study in 2011 of the community structure of planktonic virus groups in the upper lit layer of two important freshwater natural ecosystems in France, Lake Annecy (oligotrophic) and Lake Bourget (oligo-mesotrophic). Using PCR-DGGE to target a number of different structural and functional signature genes, i.e.,g20, g23, psbA, polB, and mcp, the phytoplankton viruses were shown to display temporal and spatial variability. There were marked seasonal changes in community structure for all viral groups in Lake Bourget, but only for T4-like myoviruses and psbA-containing cyanophages in Lake Annecy. The multivariate statistical analyses revealed that (1) various environmental factors can directly or indirectly explain the community structure observed for each phytoplankton viral group, and (2) temporal patterns of T4-like myovirus community structure were similar between the two lakes. In general, our results (1) suggest that the observed algal virus patterns were associated with significant shifts in phytoplankton biomass and/or structure, which in turn were shaped by the abiotic environment, and (2) support the Bank model proposed by Breitbart and Rohwer (Trends Microbiol 13:278–284, 2005). This study provides new evidence that freshwater lakes contain a significant diversity of algal viruses, and that the distribution of these viruses strongly mirrors that of their hosts.     

Viral abundance, production, decay rates and life strategies (lysogeny versus lysis) in Lake Bourget (France)

We have investigated the ecology of viruses in Lake Bourget (France) from January to August 2008. Data were analysed for viral and bacterial abundance and production, viral decay, frequency of lysogenic cells, the contribution of bacteriophages to prokaryotic mortality and their potential influence on nutrient dynamics. Analyses and experiments were conducted on samples from the epilimnion (2 m) and the hypolimnion (50 m), taken at the reference site of the lake. The abundance of virus‐like particles (VLP) varied from 3.4 × 10⁷ to 8.2 × 10⁷ VLP ml^?1; with the highest numbers and virus‐to‐bacterium ratio (VBR = 69) recorded in winter. Viral production varied from 3.2 × 10⁴ VLP ml^?1 h^?1 (July) to 2 × 10⁶ VLP ml^?1 h^?1 (February and April), and production was lower in the hypolimnion. Viral decay rate reached 0.12–0.15 day^?1, and this parameter varied greatly with sampling date and methodology (i.e. KCN versus filtration). Using transmission electron microscopy (TEM) analysis, viral lysis was responsible for 0% (January) to 71% (February) of bacterial mortality, while viral lysis varied between 0% (April) and 53% (January) per day when using a modified dilution approach. Calculated from viral production and burst size, the virus‐induced bacterial mortality varied between 0% (January) and 68% (August). A weak relationship was found between the two first methods (TEM versus dilution approach). Interestingly, flow cytometry analysis performed on the dilution experiment samples revealed that the viral impact was mostly on high DNA content bacterial cells whereas grazing, varying between 8.3% (June) and 75.4% (April), was reflected in both HDNA and LDNA cells equally. The lysogenic fraction varied between 0% (spring/summer) and 62% (winter) of total bacterial abundance, and increased slightly with increasing amounts of mitomycin C added. High percentages of lysogenic cells were recorded when bacterial abundance and activity were the lowest. The calculated release of carbon and phosphorus from viral lysis reached up to 56.5 µgC l^?1 day^?1 (assuming 20 fgC cell^?1) and 1.4 µgP l^?1 day^?1 (assuming 0.5 fgP cell^?1), respectively, which may represent a significant fraction of bacterioplankton nutrient demand. This study provides new evidence of the quantitative and functional importance of the virioplankton in the functioning of microbial food webs in peri‐alpine lakes. It also highlights methodologically dependent results.     

Seasonal and spatial variability of virio-, bacterio-, and picophytoplanktonic abundances in three peri-alpine lakes

Flow cytometry (FCM) was used to assess microbial community abundances and patterns in three natural, large and deep peri-alpine hydrosystems, i.e., lakes Annecy (oligotrophic), Bourget, and Geneva (mesotrophic). Picocyanobacteria, small eukaryotic autotrophs, heterotrophic prokaryotes, and viruses were studied in the 0–50 m surface layers to highlight the impact of both physical and chemical parameters as well as possible biotic interactions on the functioning of microbial communities. Some specificities were recorded according to the trophic status of each ecosystem such as the higher number of viruses and heterotrophic bacteria in mesotrophic environments (i.e., Lakes Geneva and Bourget) or the higher abundance of picocyanobacteria in the oligotrophic Lake Annecy. However, both seasonal (temperature) and spatial (depth) variations were comparatively more important than the trophic status in driving the microbial communities’ abundances in these three lakes, as revealed by principal component analysis (PCA). A strong viral termination of the heterotrophic bacterial blooms could be observed in autumn for each lake, in parallel to the mixing of the upper lit layers. As virus to bacteria ratio (VBR) was indeed very high at this period with values varying between 87 and 114, such important relationships between viruses and bacteria were likely. The magnitudes of seasonal variations in VBR, with the highest values ever reported so far, were largely greater than the magnitude of theoretical variations due to the trophic status, suggesting also a strong seasonality in virioplankton production associated to prokaryotic dynamics. FCM analyses allowed discriminating several viral groups. Virus-Like Particles group 1 (VLP1) and group 2 (VLP2) were always observed and significantly correlated to bacteria for the former and chlorophyll a and picocyanobacteria for the latter, suggesting that most of VLP1 and VLP2 could be bacteriophages and cyanophages, respectively. On the basis of these results, new ways of investigation emerge concerning the study of relationships between specific picoplanktonic groups; and overall these results provide new evidence of the necessity to consider further viruses for a better understanding of lake plankton ecology. Handling editor: Luigi Naselli-Flores     

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.     

Bacterial dynamics in two high-arctic lakes

The heterotrophic planktonic bacteria in two high-arctic lakes were studied by direct microscope count and the enzymatic uptake of ¹⁴C labelled glucose which generally conformed to Michaelis-Menten kinetics. Bacterial numbers and activity in oligo-trophic Char Lake ranged from 0.1 to 2.0×10^?3 bacteria/l and a maximum uptake velocity (V_max) of 1.8 × 10^?3μg glucose l^? h^?1. Nearby Meretta Lake received waste water from the Department of Transport Base at Resolute and this eutrophication was reflected in higher bacterial numbers of 2-80 × 10⁸/1 and Kmax of 0.1 × 10^?1-7.5 × 10^?1 fig glucose l^?1 h^?1 The Kmax per cell in Char Lake was 3 × 10^?11μg glucose l^?1 h^?1 and changed little between the period of solid ice cover in May and ice-free conditions in August. Bacterial cycles could not be related to phytoplankton cycles in either lake. Comparison of kinetic data from several lakes suggests a relationship between the bacterial uptake rate of glucose and phytoplankton production. Both bacterial numbers and activity in Char Lake may be very close to the minima to be expected in undisturbed freshwater environments.     

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.     

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.

     

Viral dynamics and patterns of lysogeny in saline Antarctic lakes

Antarctic lakes are extreme ecosystems with microbially dominated food webs, in which viruses may be important in controlling community dynamics. A year long investigation of two Antarctic saline lakes (Ace and Pendant Lakes) revealed high concentrations of virus like particles (VLP) (0.20–1.26 × 10⁸ ml⁻¹), high VLP: bacteria ratios (maximum 70.6) and a seasonal pattern of lysogeny differing from that seen at lower latitudes. Highest rates of lysogeny (up to 32% in Pendant Lake and 71% in Ace Lake) occurred in winter and spring, with low or no lysogeny in summer. Rates of virus production (range 0.176–0.823 × 10⁶ viruses ml⁻¹ h⁻¹) were comparable to lower latitude freshwater lakes. In Ace Lake VLP did not correlate with bacterial cell concentration or bacterial production but correlated positively with primary production, while in Pendant Lake VLP abundance correlated positively with both bacterial cell numbers and bacterial production but not with primary production. In terms of virus and bacterial dynamics the two saline Antarctic lakes studied appear distinct from other aquatic ecosystems investigated so far, in having very high viral to bacterial ratios (VBR) and a very high occurrence of lysogeny in winter.     

Grazing effects of a freshwater bivalve (Corbicula leana Prime) and large zooplankton on phytoplankton communities in two Korean lakes

This study examined the effects of a freshwater filter feeding bivalve (Corbicula leana Prime) and large zooplankton (>200 μm, mostly cladocerans and copepods) on the phytoplankton communities in two lakes with contrasting trophic conditions. A controlled experiment was conducted with four treatments (control, zooplankton addition, mussel addition, and both zooplankton and mussel addition), and each established in duplicate 10-l chambers. In both lakes there were significant effects of mussel grazing on phytoplankton density and biomass. The effects were greater in mesotrophic Lake Soyang than in hypertrophic Lake Ilgam. Effects of zooplankton grazing did not differ between these lakes, and zooplankton effects on phytoplankton were much less than the effects of mussels. Although mussels exerted a varying effect on phytoplankton according to their size, mussels reduced densities of almost all phytoplankton taxa. Total mean filtering rate (FR) of mussels in Lake Soyang was significantly greater than that in Lake Ilgam (p=0.002, n=5). Carbon fluxes from phytoplankton to mussels (977–2,379 μgC l^?1d^?1) and to zooplankton (76–264 μgC l^?1 d^?1) were always greater in Lake Ilgam due to the greater phytoplankton biomass (p<0.01, n=6). Based on the C-flux to biomass ratios, the mussels consumed 170–754% (avg. 412%) of phytoplankton standing stock in Lake Soyang, and 38–164% (avg. 106%) in Lake Ilgam per day. The C-flux to biomass ratio for mussels within each lake was much greater than for large zooplankton. Mussels reduced total phosphorus concentration by 5–34%, while increasing phosphate by 30–55% relative to the control. Total nitrogen also was reduced (by 9–25%), but there was no noticeable change in nitrate among treatments. The high consumption rate of phytoplankton by Corbicula leana even in a very eutrophic lake suggests that this mussel could affect planktonic and benthic food web structure and function by preferential feeding on small seston and by nutrient recycling. Control of mussel biomass therefore might be an effective tool for management of water quality in shallow eutrophic lakes and reservoirs in Korea.     

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.     

Seasonal Viral Loop Dynamics in Two Large Ultraoligotrophic Antarctic Freshwater Lakes

The effect of viruses on the microbial loop, with particular emphasis on bacteria, was investigated over an annual cycle in 2003–2004 in Lake Druzhby and Crooked Lake, two large ultraoligotrophic freshwater lakes in the Vestfold Hills, Eastern Antarctica. Viral abundance ranged from 0.16 to 1.56 × 10⁹ particles L^-1;1 and bacterial abundances ranged from 0.10 to 0.24 × 10⁹ cells L^-1;1, with the lowest bacterial abundances noted in the winter months. Virus-to-bacteria ratios (VBR) were consistently low in both lakes throughout the season, ranging from 1.2 to 8.4. lysogenic bacteria, determined by induction with mitomycin C, were detected on three sampling occasions out of 10 in both lakes. In Lake Druzhby and Crooked Lake, lysogenic bacteria made up between 18% and 73% of the total bacteria population during the lysogenic events. Bacterial production ranged from 8.2 to 304.9 × 10⁶ cells L^-1;1 day^-1;1 and lytic viral production ranged from 47.5 to 718.4 × 10⁶ viruslike particles L^-1;1 day^-1;1. When only considering primary production, heterotrophic nanoflagellate (HNF) grazing and viral lysis as the major contributors to the DOC pool (i.e., autochthonous sources), we estimated a high contribution from viruses during the winter months when >60% of the carbon supplied to the DOC pool originated from viral lysis. In contrast, during the summer <20% originated from viral lysis. Our study shows that viral process in ultraoligotrophic Antarctic lakes may be of quantitative significance with respect to carbon flow especially during the dark winter period.     

Organic carbon burial in lake sediments in the middle and lower reaches of the Yangtze River Basin, China

Lakes are important in the global and regional carbon cycle, and lake sediments potentially store substantial quantities of organic carbon. The middle and lower reaches of the Yangtze River basin (MLYB) are some of the largest agricultural areas in China with an extremely high density of lakes and rivers. The lakes in the region have undergone dramatic changes over the past several decades. In this study, six cores from five lakes (the macrophyte-dominated: Shijiuhu Lake and Honghu Lake; the algae-dominated: Chaohu Lake, Taihu Lake, and Nanyihu Lake) in the MLYB were collected from 2002 A.D. to 2008 A.D. Mass accumulation rates (MARs) of sediment derived from ²¹⁰Pb and ¹³⁷Cs along with total organic carbon content (TOC) were used to determine organic carbon accumulation rates (OC ARs) over the last 100 years. The TOC in the five lakes exhibited a significant increase since the mid or late 20th century, which was consistent with the increase in the lake water trophic status due to nutrient input. The average organic carbon accumulation rates for the Taihu Lake, Nanyihu Lake, Chaohu Lake, Shijiuhu Lake, and Honghu Lake were calculated to be 16.6, 28.9, 9.8, 25.4, and 113.2 g C m^?2 year^?1, respectively, over the past 100 years. Based on the average OC AR of 32.1 g C m^?2 year^?1 from the five lakes, carbon burial in lake sediments may be as much as 6.8 × 10¹³ g C in the MLYB over the past 100 years.     

Primary production of aquatic macrophytes and their epiphytes in two shallow lakes (Peipsi and Võrtsjärv) in Estonia

In shallow lakes with large littoral zones, epiphytes and submerged macrophytes can make an important contribution to the total annual primary production. We investigated the primary production (PP) of phytoplankton, submerged macrophytes, and their epiphytes, from June to August 2005, in two large shallow lakes. The production of pelagic and littoral phytoplankton and of the dominant submerged macrophytes in the littoral zone (Potamogeton perfoliatus in Lake Peipsi and P. perfoliatus and Myriopyllum spicatum in Lake Võrtsjärv) and of their epiphytes was measured using a modified ¹⁴C method. The total PP of the submerged macrophyte area was similar in both lakes: 12.4 g C m^?2 day^?1 in Peipsi and 12.0 g C m^?2 day^?1 in Võrtsjärv. In Peipsi, 84.2% of this production was accounted for by macrophytes, while the shares of phytoplankton and epiphytes were low (15.6 and 0.16%, respectively). In Võrtsjärv, macrophytes contributed 58%, phytoplankton 41.9% and epiphytes 0.1% of the PP in the submerged macrophyte area. Epiphyte production in both lakes was very low in comparison with that of phytoplankton and macrophytes: 0.01, 5.04, and 6.97 g C m^?2 day^?1, respectively, in Võrtsjärv, and 0.02, 1.93, and 10.5 g C m^?2 day^?1, respectively, in Peipsi. The PP of the littoral area contributed 10% of the total summer PP of Lake Peipsi sensu stricto and 35.5% of the total summer PP of Lake Võrtsjärv.     

Virus dynamics in a large epishelf lake (Beaver Lake,Antarctica)

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Photosynthesis,mixotrophy and microbial plankton dynamics in two high Arctic lakes during summer

Photosynthesis and microbial plankton dynamics of two lakes in the Kongsfjorden catchment of Spitzbergen (Svalbard Archipelago, 78°N) were investigated during the summer of 2000. One of the lakes (Tvillingvatnet) served as the water supply for the village of Ny Ålesund. The other was a deeper, larger unnamed lake in the Ossian Sarsfjella reserve—named Lake OS in this study. Both lakes can be classified as oligotrophic on the basis of chlorophyll a and inorganic nutrient concentrations. Chlorophyll a concentrations ranged between 0.33 and 1.65 µg l ^-1 in Tvillingvatnet and 0.2 and 0.3 µg l ^-1 in Lake OS. The phytoplankton was dominated by chrysophytes and cryptophytes, with the diatom Rhizosolenia and a number of dinoflagellate species. Rates of photosynthesis were typically low, between 24.5 µg and 1.0 mg l ^-1 day ^-1 in Tvillingvatnet (photosynthetic efficiency 0.004–0.26), and between 3.1 and 29.5 µg l ^-1 day ^–1 in Lake OS (photosynthetic efficiency 0.0019–0.0085). Among the phytoflagellates (PNAN) there were a number of mixotrophs which reached their peaks of abundance before other PNAN. Mixotrophy appeared to provide a competitive advantage. Grazing rates for the mixotroph Dinobryon ranged between 0.063 and 1.12 pg C cell ^-1 day ^-1 in Lake OS and between 0.22 and 1.11 pg cell ^-1 day ^-1 in Tvillingvatnet, with rates increasing between July and August. However, Dinobryon removed less than 1% of bacterial biomass day ^-1, while the heterotrophic nanoflagellates (HNAN) removed up to 28% of bacterial biomass day ^-1. Bacterial concentrations were low, with a maximum of 28.8×10 ⁸ l ^-1 in Tvillingvatnet and 23.6×10 ⁸ l ^-1 in Lake OS. Ciliated protozoan and rotifer diversity in the plankton was low.     

Prevalence of Viral Photosynthetic and Capsid Protein Genes from Cyanophages in Two Large and Deep Perialpine Lakes

Cyanophages are important components of aquatic ecosystems, but their genetic diversity has been little investigated in freshwaters. A yearlong survey was conducted in surface waters of the two largest natural perialpine lakes in France (Lake Annecy and Lake Bourget) to investigate part of this cyanophage diversity through the analysis of both structural (e.g., g20) and functional (e.g., psbA) genes. We found that these cyanophage signature genes were prevalent throughout the year but that the community compositions of g20 cyanomyoviruses were significantly different between the two lakes. In contrast, psbA-containing cyanophages seemed to be more similar between the two ecosystems. We also found that a large proportion of g20 sequences grouped with cyanomyophage isolates. psbA sequences, belonging to phages of Synechococcus spp., were characterized by distinct triplet motifs (with a novel viral triplet motif, EFE). Thus, our results show that cyanophages (i) are a diverse viral community in alpine lakes and (ii) are clearly distinct from some other freshwater and marine environments, suggesting the influence of unique biogeographic factors.     

Mixotrophic algae in three ice-covered lakes of the Pocono Mountains, U.S.A.

1. The occurrence and grazing activity of mixotrophic (phagotrophic) algae in three icecovered freshwater lakes of different trophic status were examined (oligotrophic Lake Giles, mesotrophic Lake Lacawac, eutrophic Lake Waynewood), Microbial population densities were low (4.1–7.2 × 10⁵ bacteria ml^?1 and 1.2–2.4 × 10³ nanoplanktonic protists ml^?1). All three nanoplankton communities were dominated by chloroplast-bearing forms (60–96%). 2. Mixotrophs formed up to 48% of the phototrophic nanoplankton in Lake Lacawac and were responsible for up to ～90% of the observed uptake of bacteria-sized particles. The abundance of mixotrophic algae in Lakes Giles and Waynewood were extremely low (3 and 2% of the phototrophic algae, respectively), and heterotrophs dominated nanoplankton bacterivory. 3. The overall impact of nanoplankton feeding activity on the bacterial assemblage was low under the ice in Lakes Giles and Waynewood. Removal rates of bacteria based on our particle uptake experiments were 1.0 and 4.0% of the bacterial standing stock day^?1 in these lakes, respectively. Removal rates were higher in Lake Lacawac and ranged from 4.9 to 11% of the bacterial standing stock day^?1 on 2 successive sampling days.     

Analysis of long-term variation in phytoplankton biovolume in the northern basin of Lake Biwa

The long-term variation in phytoplankton biovolume in the northern basin of Lake Biwa was analyzed using periodic phytoplankton census data from January 1979 to December 2009. Population densities obtained from census data were transformed into biovolumes, and phytoplankton species were categorized into three size fractions: net phytoplankton (≥4,000 μm³ cell^?1, ≥ca. 20 μm in diameter), large nanophytoplankton (100–4,000 μm³ cell^?1, ca. 6–20 μm in diameter), and small nanophytoplankton (<100 μm³ cell^?1, <ca. 6 μm in diameter). Although the annual total biovolume gradually decreased over time, the total biovolumes in winter and spring were found to increase. Furthermore, a decrease in the biovolume of net phytoplankton and an increase in that of small nanophytoplankton were observed. Because of succession in the phytoplankton community, the average cell volume of the phytoplankton community decreased from 269 μm³ cell^?1 in the 1980s to 56 μm³ cell^?1 in the 2000s. Lake warming accompanied with the intensification of thermal stratification and the augmentation of wind speed were observed at Lake Biwa over the study period. Serial analysis correcting for autocorrelation revealed that oligotrophication in the epilimnion, induced by lake warming and limitation of light available for phytoplankton growth by wind-induced water mixing, was a potential factor in the succession of the phytoplankton community.     

Dust mediated transfer of phosphorus to alpine lake ecosystems of the Wind River Range,Wyoming, USA

Alpine lakes receive a large fraction of their nutrients from atmospheric sources and are consequently sensitive to variations in both the amount and chemistry of atmospheric deposition. In this study we explored the spatial changes in lake water chemistry and biology along a gradient of dust deposition in the Wind River Range, Wyoming. Regional differences were explored using the variation in bulk deposition, lake water, sediment, and bedrock geochemistry and catchment characteristics. Dust deposition rates in the Southwestern region averaged 3.34 g m^?2 year^?1, approximately three times higher than deposition rates in the Northwestern region (average 1.06 g m^?2 year^?1). Dust-P deposition rates ranged from 87 µg P m² day^?1 in the Northwestern region to 276 µg P m² day^?1 in the Southwestern region. Subalpine and alpine lakes in the Southwestern region had greater total phosphorus (TP) concentrations (5–13 µg L^?1) and greater sediment phosphorus (SP) concentrations (2–5 mg g^?1) than similar lakes elsewhere in the region (1–8 µg L^?1 TP, 0.5–2 mg g^?1 SP). Lake phosphorus concentrations were related to dissolved organic carbon (DOC) across vegetation gradients, but related to the percent of bare rock, catchment area to lake area, and catchment steepness across dust deposition gradients. Modern phytoplankton and zooplankton biomasses were two orders of magnitude greater in the Southwest than in the Northwest, and alpine lakes in the Southwest had a unique diatom species assemblage with relatively higher concentrations of Asterionella formosa, Pseudostaurosira pseudoconstruens, and Pseudostaurosira brevistriata. These results suggests that catchment controls on P export to lakes (i.e. DOC) are overridden in dominantly bare rock basins where poor soils cannot effectively retain dust deposited P.     

Estimation of reference conditions for phytoplankton in a naturally eutrophic shallow lake

Classification of waters using biological quality elements and determination of the degree of deviation from reference levels is a key issue in the Water Framework Directive of EU. Lakes in reference conditions with sufficient biological data are available for several boreal lake types with the exception of naturally eutrophic lakes. An empirical approach is one alternative for estimating the reference conditions of such lakes. We used the water transparency of the naturally eutrophic Lake Tuusulanjärvi recorded in August in the early 1910s to estimate reference values for phytoplankton biomass and chlorophyll a concentrations. Three phytoplankton samples during August 2000–2001 corresponded to the estimated reference values for total biomass (<5.6 mg l^?1) and chlorophyll a (<28 μg l^?1), as did the simultaneous Secchi depths. The phytoplankton assemblage in these samples with 24 eutrophy indicators (17% of the total taxa number) corresponded in general the species list from the early 1900s, which as such could be regarded as reference assemblage. Furthermore, in August 2000, 3 years after intensive fish removal a prominent decrease in cyanobacterial biomass and toxin concentration was observed. The costs of the measures and studies in Lake Tuusulanjärvi during 1989–2003 have been approximately 2.5 million euros.