Effect of winter conditions on distributions of anoxic phototrophic bacteria in two meromictic lakes in Siberia,Russia

The year-to-year variations of vertical distribution and biomass of anoxic phototrophic bacteria were studied during ice periods 2003–2005 and 2007–2008 in meromictic lakes Shira and Shunet (Southern Siberia, Russian Federation). The bacterial layers in chemocline of both lakes were sampled with a thin-layer hydraulic multi-syringe sampler. In winter, biomass of purple sulphur bacteria varied considerably depending on the amount of light penetrating into the chemocline through the ice and snow cover. In relatively weakly stratified, brackish Shira Lake, the depth of chemocline varied between winters, so that light intensity for purple sulphur bacteria inhabiting this zone differed. In Shira Lake, increased transparency of mixolimnion in winter, high chemocline position and absence of snow resulted in light intensity and biomass of purple sulphur bacteria exceeding the summer values in the chemocline of the lake. We could monitor snow cover at the lake surface using remote sensing and therefore estimate dynamics and amount of light under ice and its availability for phototrophic organisms. In Shunet Lake, the light intensities in the chemocline and biomasses of purple sulphur bacteria were always lower in winter than in summer, but the biomasses of green sulphur bacteria were similar.     

The vertical distribution and abundance of <Emphasis Type="Italic">Gammarus lacustris</Emphasis> in the pelagic zone of the meromictic lakes Shira and Shunet (Khakassia,Russia)

The vertical distribution and abundance of Gammarus lacustris in the pelagic zone of two fishless meromictic lakes, L. Shira and L. Shunet, in Southern Siberia (Russia), was studied with the underwater video recording system and using vertical hauls. In both lakes, during summer stratification, Gammarus was distributed non-homogenously, with a stable peak in the metalimnion. The average depth of Gammarus population in the pelagic zone was significantly correlated with the depth of the thermocline. Gammarus abundances obtained using vertical plankton hauls with net were quite comparable with those obtained from video records. The peak abundance of Gammarus in the pelagic zone of the lakes observed with underwater video amounted up to 400 individuals m⁻², while the peak animal densities in the metalimnion reached 50 ind. m⁻³. The data are compared with previously published abundances of Gammarus in the littoral of Lake Shira. Both littoral and pelagic can be equally important habitats for amphipods in meromictic lakes. The absence of fish in the pelagic zone, high oxygen concentration, low water temperature, increased seston concentration, elevated water density in the metalimnion and the anoxic hypolimnion can be the most probable combination of factors that are responsible for the peak of Gammarus in the metalimnion of these lakes.     

Stratification and dynamics of microbial loop communities in Lake Fryxell, Antarctica

1. Lake Fryxell, situated in the McMurdo Dry Valleys, Antarctica, offers the opportunity to study microbial loop processes in the absence of crustacean zooplankton and other higher organisms. This is the first study of Lake Fryxell to provide detailed temporal and vertical variations of microbial loop organisms.
2. Protozoan communities are concentrated around the chemocline (9–10 m) in Lake Fryxell. Phototrophic nanoflagellates (PNAN), heterotrophic nanoflagellates (HNAN) and ciliates formed deep maxima of 14 580, 694 and 58 cells mL−1 respectively. Although abundance and biomass at the chemocline was high, diversity of protozoa was low, Plagiocampa accounting for> 80% of the total ciliate biomass.
3. In the mixolimnion (4.5–8 m), protozoa were less abundant, but more diverse, with 24 ciliate morphotypes being identified within this region of the water column. Inter-annual variability of protozoan biomass and abundance was greater in the mixolimnion than at the chemocline due to more variable nutrient and prey concentrations.
4. Physicochemical gradients in Lake Fryxell were very stable because the perennial ice cover reduced wind driven currents. As a consequence, ciliate species occurred in distinct depth strata, Monodinium being most abundant directly beneath the ice cover, Askenasia having maximum abundance at 8 m and Plagiocampa dominating ciliate biomass at the chemocline. The lack of vertical mixing reduced seasonal successions of PNAN and ciliate species. Three cryptophyte species dominated the PNAN community at all times (>79% of total biomass).     

Bacterial Communities of Three Saline Meromictic Lakes in Central Asia

Meromictic lakes located in landlocked steppes of central Asia (~2500 km inland) have unique geophysiochemical characteristics compared to other meromictic lakes. To characterize their bacteria and elucidate relationships between those bacteria and surrounding environments, water samples were collected from three saline meromictic lakes (Lakes Shira, Shunet and Oigon) in the border between Siberia and the West Mongolia, near the center of Asia. Based on in-depth tag pyrosequencing, bacterial communities were highly variable and dissimilar among lakes and between oxic and anoxic layers within individual lakes. Proteobacteria, Bacteroidetes, Cyanobacteria, Actinobacteria and Firmicutes were the most abundant phyla, whereas three genera of purple sulfur bacteria (a novel genus, Thiocapsa and Halochromatium) were predominant bacterial components in the anoxic layer of Lake Shira (~20.6% of relative abundance), Lake Shunet (~27.1%) and Lake Oigon (~9.25%), respectively. However, few known green sulfur bacteria were detected. Notably, 3.94% of all sequencing reads were classified into 19 candidate divisions, which was especially high (23.12%) in the anoxic layer of Lake Shunet. Furthermore, several hydro-parameters (temperature, pH, dissolved oxygen, H₂S and salinity) were associated (P< 0.05) with variations in dominant bacterial groups. In conclusion, based on highly variable bacterial composition in water layers or lakes, we inferred that the meromictic ecosystem was characterized by high diversity and heterogenous niches.     

Sulfate reduction and methanogenesis in the Shira and Shunet meromictic lakes (Khakass Republic, Russia)

The biogeochemical and molecular biological study of the chemocline and sediments of saline meromictic lakes Shira and Shunet (Khakass Republic, Russia) was performed. A marked increase in the rates of sulfate reduction and methanogenesis was revealed at the medium depths of the chemocline. The rates of these processes in the bottom sediments decreased with depth. The numbers of Bacteria, Archaea, and of sulfate-reducing bacteria (SRB) were determined by fluorescence in situ hybridization with rRNA specific oligonucleotide probes labeled with horseradish peroxidase and subsequent tyramide signal amplification. In the chemocline, both the total microbial numbers and those of Bacteria were shown to increase with depth. The archaea and SRB were present in almost equal numbers. In the lake sediments, a drastic decrease in microbial numbers with depth was revealed. SRB were found to prevail in the upper sediment layer and archaea in the lower one. This finding correlates with the measured rates of sulfate reduction and methanogenesis.     

Sulfate reduction and methanogenesis in the Shira and Shunet meromictic lakes (Khakasia, Russia)

The biogeochemical and molecular biological study of the chemocline and sediments of saline meromictic lakes Shira and Shunet (Khakasia, Russia) was performed. A marked increase in the rates of sulfate reduction and methanogenesis was revealed at the medium depths of the chemocline. The rates of these processes in the bottom sediments decreased with depth. The numbers of the members of domains Bacteria, Archaea, and of sulfate-reducing bacteria (SRB) were determined by fluorescence in situ hybridization with rRNA specific oligonucleotide probes labeled with horseradish peroxidase and subsequent tyramide signal amplification. In the chemocline, both the total microbial numbers and those of Bacteria were shown to increase with depth. The archaea and SRB were present in almost equal numbers. In the lake sediments, a drastic decrease in microbial numbers with depth was revealed. SRB were found to prevail in the upper sediment layer and archaea in the lower one. This finding correlated with the measured rates of sulfate reduction and methanogenesis.     

A general one-dimensional vertical ecosystem model of Lake Shira (Russia,Khakasia): description,parametrization and analysis

A one-dimensional ecological model of the meromictic brackish Lake Shira (Russia, Khakasia) was developed. The model incorporates state-of-the-art knowledge about the functioning of the lake ecosystem using the most recent field observations and ideas from PCLake, a general ecosystem model of shallow freshwater lakes. The model of Lake Shira presented here takes into account the vertical dynamics of biomasses of the main species of algae, zooplankton and microbial community, as well as the dynamics of oxygen, detritus, nutrients and hydrogen sulphide from spring to autumn. Solar radiation, temperature and diffusion are modelled using real meteorological data. The parameters of the model were calibrated to the field data, after applying different methods of sensitivity analysis to the model. The resulting patterns of phytoplankton and nutrients dynamics show a good qualitative and quantitative agreement with the field observations during the whole summer season. Results are less satisfactory with respect to the vertical distribution of zooplankton biomass. We hypothesize that this is due to the fact that the current model does not take the sex and age structure of zooplankton into account. The dynamics of oxygen, hydrogen sulphide and the modelled positions of the chemocline and thermocline are again in good agreement with field data. This resemblance confirms the validity of the approach we took in the model regarding the main physical, chemical and ecological processes. This general model opens the way for checking various hypotheses on the functioning of the Lake Shira ecosystem in future investigations and for analysing options for management of this economically important lake.     

Densities and distribution of flagellates and ciliates in the chemocline of saline,meromictic Lake Shunet (Siberia,Russia)

The vertical and seasonal distributions of the phytoflagellate Cryptomonas spp., and its most common, the planktonic ciliate predators (Oligotrichida, Scuticociliatida, Hypotrichida and Prostomatida) were investigated in chemocline region of small saline, meromictic lake Shunet (Siberia, Russia) during 2003 and 2005. The lake has a pronounced chemocline, with abundance of purple and green sulphur bacteria. Vertical distribution of the Cryptomonas populations near the oxic/anoxic boundary layer was studied at close intervals in water sampled using a hydraulically operated thin-layer sampler. In both summer and winter, Cryptomonas peaked in water stratum 5–10 cm above anoxic zone or in the anoxic zone water column in the chemocline (about 5 m). Ciliate densities and biomass were also much higher in chemocline than in mixolimnion. The range of diurnal migration of Cryptomonas population was not very wide, and it was restricted to layers with high light intensity. The ciliates were sometimes detected above the upper border of the anoxic zone but also several centimetres below this zone.     

Ecology of purple sulfur bacteria in the highly stratified meromictic Lake Shunet (Siberia, Khakassia) in 2002–2009

Phototrophic sulfur bacteria form dense accumulations in the chemocline zones of stratified lakes where light reaches the sulfide-containing layers of water. Many works are dedicated to the ecophysiology of these microorganisms in meromictic lakes. However, the role of these microorganisms in the trophic network of these ecosystems, the ways of biomass utilization, and the contribution to the turnover of biogenic elements have so far been insufficiently understood. This work deals with the analysis of many years?? seasonal dynamics of the biomass of purple sulfur bacteria and the physicochemical conditions of their environment in Lake Shunet (Siberia, Khakassia, Russia), unraveling the causes of their anomalous development in the chemocline of this lake, as well as the comparative analysis of such type of ecosystems. Lake Shunet is characterized by markedly pronounced stratification and the high density of purple sulfur bacteria (PSB) in the chemocline, which is comparable to that of Lake Mahoney (Canada) where the number of PSB is the greatest among those known in the world. It was shown that, in the period 2002?C2009, the total amount of bacterio-chlorophyll a in the water column of Lake Shunet increased and did not correlate with the seasonal variations in temperature and illumination in the chemocline. It was established that PSB cells in the purple layer experienced the effect of self-shading. The sedimentation rate of purple sulfur bacteria in Lake Shunet was low due to the pronounced density gradient in the chemocline zone. Thus, the high number of PSB in the chemocline was due to the combination of strong illumination, a high sulfide concentration, and a high water density gradient, which was responsible for stable stratification and contributed to the accumulation of the cells in a narrow layer. The data obtained could be useful for the paleoreconstruction of climatically deter-mined changes in the level of the lake and its periods of meromixis by the presence of carotenoids and bacte-riochlorophylls in the bottom sediments.     

Microbiological and isotopic-geochemical investigations of meromictic lakes in Khakasia in winter

Microbiological and isotopic-geochemical investigations of the brackish meromictic lakes Shira and Shunet were performed in the steppe region of Khakasia in winter. Measurements made with a submersed sensor demonstrated that one-meter ice transmits light in a quantity sufficient for oxygenic and anoxygenic photosynthesis. As in the summer season, in the community of phototrophic bacteria found in Lake Shira, the purple sulfur bacteria Amoebobacter purpureus dominated, whereas, in Lake Shunet, the green sulfur bacteria Pelodictyon luteolum were predominant. Photosynthetic production, measured using the radioisotopic method, was several times lower than that in summer. The rates of sulfate reduction and production and oxidation of methane in the water column and bottom sediments were also lower than those recorded in summer. The process of anaerobic methane oxidation in the sediments was an exception, being more intense in winter than in summer. The data from radioisotopic measurements of the rates of microbial processes correlate well with the results of determination of the isotopic composition of organic and mineral carbon (delta13C) and hydrogen sulfide and sulfate (delta34S) and suggest considerable seasonal variations in the activity of the microbial community in the water bodies investigated.     

Asynchronous vertical migrations of zooplankton in stratified lakes

Asynchronous vertical migrations of calanoid copepods Arctodiaptomus salinus were studied in two stratified lakes in the south of Siberia using the method of two-section enclosures. It was found that the presence of a pronounced thermocline and a depth maximum of phytoplankton (Lake Shira) contributes to the appearance of intensive individual migrations of copepods between areas of the epi- and hypolimnion.     

Microbial community of the chemocline of the meromictic Lake Shunet (Khakassia,Russia) during summer stratification

The spatio-temporal organization of the bacterial community inhabiting the chemocline of the stratified meromictic Lake Shunet (Khakassia, Russia) was investigated from May to September 2005 by means of microscopy, analysis of photosynthetic pigments, and PCR-DGGE with subsequent 16S rDNA analysis. The samples were collected with a multisyringe stratification sampler, sampling being performed every 5 cm. It was demonstrated that, during the period of investigation, there were no large changes in the bacterial community of the chlemocline, at least among the detected forms. During the whole period of study, purple sulfur bacteria related to Lamprocystis purpurea (Chromatiaceae) were predominant in the chemocline. Beneath the layer of purple bacteria, green sulfur bacteria were revealed that were phylogenetically distant from strain ShNPel02, which was previously isolated from this lake. Development of phytoflagellates of the genus Cryptomonas was observed in the upper zone of the chemocline. In the chemocline of Lake Shunet, the numbers of picoplankton cyanobacteria of the genus Synechococcus increased from May to September. It was demonstrated that the application of universal bacterial primers for DGGE resulted in the same qualitative distributional pattern of predominant species as microscopic studies.     

Ace Lake: three decades of research on a meromictic,Antarctic lake

Ace Lake (Vestfold Hills, Antarctica) has been investigated since the 1970s. Its close proximity to Davis Station has allowed year-long, as well as summer only, investigations. Ace Lake is a saline meromictic (permanently stratified) lake with strong physical and chemical gradients. The lake is one of the most studied lakes in continental Antarctica. Here, we review the current knowledge of the history, the physical and chemical environment, community structure and functional dynamics of the mixolimnion, littoral benthic algal mats, the lower anoxic monimolimnion and the sediment within the monimolimnion. In common with other continental meromictic Antarctic lakes, Ace Lake possesses a truncated food web dominated by prokaryote and eukaryote microorganisms in the upper aerobic mixolimnion and an anaerobic prokaryote community in the monimolimnion, where methanogenic Archaea, sulphate-reducing and sulphur-oxidizing bacteria occur. These communities are functional in winter at subzero temperatures, when mixotrophy plays an important role in survival in dominant photosynthetic eukaryotic microorganisms in the mixolimnion. The productivity of Ace Lake is comparable to other saline lakes in the Vestfold Hills, but higher than that seen in the more southerly McMurdo Dry Valley lakes. Finally, we identify gaps in the current knowledge and avenues that demand further investigation, including comparisons with analogous lakes in the north polar region.     

Vertical Distribution of Archaea and Bacteria in a Meromictic Lake as Determined by Fluorescent In Situ Hybridization

The prokaryotic cells distribution in the water column of the coastal saline meromictic Lake Faro (Messina, Italy) was investigated by microscopic counting techniques. Water samples were collected at a central station from the surface to the bottom, when waters were characterized by a marked stratification. A “red-water” layer, caused by a dense growth of photosynthetic sulfur bacteria, was present at a depth of 15 m, defining a transition area between oxic (mixolimnion) and anoxic (monimolimnion) layers. Fluorescently labeled 16S rRNA oligonucleotide, group-specific probes were used to determine the abundance of Bacteria and Archaea, and their subgroups, Green Sulfur Bacteria (GSB), Sulfate Reducing Bacteria (SRB), Cyanobacteria and Chromatium okenii, and Crenarchaeota and Euryarchaeota, as key elements of the microbial community. Bacteria decreased from surface to bottom, while Archaea increased with depth and reached the maximum value at 30 m, where they outnumbered the Bacteria. Bacteria and picophytoplankton prevailed in the mixolimnion. At the chemocline high numbers of prokaryotic cells were present, mainly represented by Cyanobacteria, Chromatium okenii and Euryarchaeota. GSB, SRB, and Crenarchaeota prevailed below the chemocline. Although Archaea constitute a minor fraction of microbial community, they could represent active contributors to the meromictic Lake Faro ecosystem.     

Feeding spectra of <Emphasis Type="Italic">Arctodiaptomus salinus</Emphasis> (Calanoida,Copepoda) using fatty acid trophic markers in seston food in two salt lakes in South Siberia (Khakasia,Russia)

During two vegetation seasons (2004–2005), we compared feeding spectra of Arctodiaptomus salinus (Calanoida, Copepoda) populations inhabiting two neighboring salt lakes, Shira and Shunet, Khakasia, Russia, using fatty acid (FA) trophic markers. Sestonic FA composition in two lakes moderately differed, whereas levels of diatom FA markers were higher in Lake Shunet and of Cyanobacteria and green algae markers in Lake Shira. In general, markers in storage lipids—triacylglycerols (TAG) of A. salinus—reflected the differences in sestonic composition of the two lakes. Nevertheless, TAG fraction was also enriched by FA trophic markers of the minor components of seston, which were selectively ingested by the animals. In Lake Shira, A. salinus had significantly higher concentrations of bacterial FA markers in TAG. In Lake Shunet, TAG of A. salinus contained significantly higher relative amounts of 18:4ω3, 18:5ω3 and C22 polyunsaturated fatty acids (PUFA), which indicated marked contribution of cryptophytes or (and) flagellates into the diet. Laboratory experiments showed feeding on Cryptomonas and sulfur purple bacteria in Lake Shunet and ciliates and colonial picoplankton in both lakes, and generally confirmed the differences in FA trophic markers in A. salinus between the lakes. The two populations of A. salinus markedly differed in levels of essential long-chain PUFA, eicosapentaenoic (EPA) and docosahexaenoic (DHA) acids, although the levels of these FA in seston were quite similar between the two lakes. The higher levels of the essential ω3 PUFA in A. salinus in Lake Shunet may be an adaptive response of the animals to a vertical stratification of physico-chemical conditions and significantly higher salinity levels at the boundary of adjacent bottom layer in this lake.     

Microbiological and Isotopic-Geochemical Investigations of Meromictic Lakes in Khakasia in Winter

Microbiological and isotopic-geochemical investigations of the brackish meromictic lakes Shira and Shunet were performed in the steppe region of Khakasia in winter. Measurements made with a submersed sensor demonstrated that one-meter ice transmits light in a quantity sufficient for oxygenic and anoxygenic photosynthesis. As in the summer season, in the community of phototrophic bacteria found in Lake Shira, the purple sulfur bacteria Amoebobacter purpureus dominated, whereas, in Lake Shunet, the green sulfur bacteria Pelodictyon luteolum were predominant. Photosynthetic production, measured using the radioisotopic method, was several times lower than that in summer. The rates of sulfate reduction and production and oxidation of methane in the water column and bottom sediments were also lower than those recorded in summer. The process of anaerobic methane oxidation in the sediments was an exception, being more intense in winter than in summer. The data from radioisotopic measurements of the rates of microbial processes correlate well with the results of determination of the isotopic composition of organic and mineral carbon (δ¹³C) and hydrogen sulfide and sulfate (δ³⁴S) and suggest considerable seasonal variations in the activity of the microbial community in the water bodies investigated.__________Translated from Mikrobiologiya, Vol. 74, No. 4, 2005, pp. 552–561.Original Russian Text Copyright © 2005 by Savvichev, Rusanov, Rogozin, Zakharova, Lunina, Bryantseva, Yusupov, Pimenov, Degermendzhi, Ivanov.     

Some generalizations based on stratification and vertical mixing in meromictic Lake Shira,Russia, in the period 2002–2009

In a brackish, temperate, 24-m-deep Lake Shira, the profiles of salinity, temperature, oxygen and sulfide concentrations were measured on a seasonal basis from 2002 to 2009. The lake was shown to be meromictic with autumnal overturn restricted to mixolimnion. The depth of mixolimnion and position of oxic–anoxic interface varied annually. The spring mixing processes contribute to the formation of mixolimnion in autumn. The exceptionally windy spring of 2007 caused the deepening of mixolimnion in the winter of 2008. The winter position of oxic–anoxic interface was affected by the position of lower boundary of mixolimnion in all winters. The salinity in the winter mixolimnion increased compared with the autumn because of freezing out of salts from the upper water layers meters during ice formation and their dissolution in water below. The profiles of salinity and temperature were simulated by the mathematical 1-D model of temperature and salinity conditions taking into account ice formation. The simulated profiles generally coincided with the measured ones. The coincidence implies that simplified one-dimensional model can be applied to roughly describe salinity and density profiles and mixing behavior of Lake Shira.     

Numerical modeling of vertical stratification of Lake Shira in summer

A one-dimensional numerical model and a two-dimensional numerical model of the hydrodynamic and thermal structure of Lake Shira during summer have been developed, with several original physical and numerical features. These models are well suited to simulate the formation and dynamics of vertical stratification and provide a basis for an ecological water-quality model of the lake. They allow for the quantification of the vertical mixing processes that govern not only the thermal structure but also the nutrient exchange, and more generally, the exchange of dissolved and particulate matter between different parts of the lake. The outcome of the calculations has been compared with the field data on vertical temperature and salinity distributions in Lake Shira. Lake Shira is meromictic and exhibits very stable annual stratification. The stratification is so stable because of the high salinity of the water. If the water in Lake Shira were fresh and other parameters (depth, volume, and meteorology) were the same, as now, the lake would be mixed in autumn. Using the newly developed models and using common meteorological parameters, we conclude that Lake Shira will remain stratified in autumn as long as the average salinity is higher than 3‰.     

Meromixis and Seasonal Dynamics of Vertical Structure of Lake Uchum (South Siberia)

The seasonal dynamics of the vertical structure of small saline Lake Uchum, located in the steppe arid zone of the south of Siberia (Krasnoyarsk krai), has been studied in detail for the first time. This lake is a meromictic water body. We have revealed a heterogeneous vertical distribution of plankton organisms and a dense population of purple sulfuric bacteria in the redox zone. The taxonomic composition and seasonal dynamics of phyto- and zooplankton are described. Presumably, the meromixis of Lake Uchum is due to the inflow of fresh water to the surface of the saline water body during the rise of its level in the early 20th century, similarly to lakes Shira and Shunet located nearby. The processes of salt displacement into the solution during the formation of ice, as well as the precipitation of salts in the winter, also contribute to the maintenance of permanent stratification. The information on the current state of the lake can be useful for reconstructing the climate by bottom sediments, as well as for creating a model of water quality and investigating the therapeutic properties of lake mud.