Cold-active chemoorganotrophic bacteria from permanently ice-covered Lake Hoare, McMurdo Dry Valleys, Antarctica

Eight strains of chemoorganotrophic bacteria were isolated from the water column of Lake Hoare, McMurdo Dry Valleys, Antarctica, using cold enrichment temperatures. The isolates were Alpha-, Beta-, and Gammaproteobacteria and Actinobacteria spp. All isolates grew at 0 degrees C, and all but one grew at subzero temperatures characteristic of the water column of Lake Hoare. Growth temperature optima varied among isolates, but the majority showed optima near 15 degrees C, indicative of cold-active phenotypes. One isolate was truly psychrophilic, growing optimally around 10 degrees C and not above 20 degrees C. Half of the isolates grew at 2% salt while the other half did not, and all but one isolate grew at 2 atm of O(2). Our isolates are the first prokaryotes from the water column of Lake Hoare to be characterized phylogenetically and physiologically and show that cold-active species of at least two major phyla of Bacteria inhabit Lake Hoare.     

Bacteriohopanepolyols across environmental gradients in Lake Vanda,Antarctica

Bacteriohopanepolyols (BHPs) are bacterial membrane lipids that may be used as biological or environmental biomarkers. Previous studies have described the diversity, distribution, and abundance of BHPs in a variety of modern environments. However, the regulation of BHP production in polar settings is not well understood. Benthic microbial mats from ice‐covered lakes of the McMurdo Dry Valleys, Antarctica provide an opportunity to investigate the sources, physiological roles, and preservation of BHPs in high‐latitude environments. Lake Vanda is one of the most stable lakes on Earth, with microbial communities occupying specific niches along environmental gradients. We describe the influence of mat morphology and local environmental conditions on the diversity and distribution of BHPs and their biological sources in benthic microbial mats from Lake Vanda. The abundance and diversity of C‐2 methylated hopanoids (2‐MeBHP) are of particular interest, given that their stable degradation products, 2‐methylhopanes, are among the oldest and most prevalent taxonomically informative biomarkers preserved in sedimentary rocks. Furthermore, the interpretation of sedimentary 2‐methylhopanes is of great interest to the geobiology community. We identify cyanobacteria as the sole source of 2‐MeBHP in benthic microbial mats from Lake Vanda and assess the hypothesis that 2‐MeBHP are regulated in response to a particular environmental variable, namely solar irradiance.     

Perennially ice-covered Lake Hoare,Antarctica: physical environment,biology and sedimentation

Lake Hoare (77° 38 S, 162° 53 E) is a perennially ice-covered lake at the eastern end of Taylor Valley in southern Victoria Land, Antarctica. The environment of this lake is controlled by the relatively thick ice cover (3–5 m) which eliminates wind generated currents, restricts gas exchange and sediment deposition, and reduces light penetration. The ice cover is in turn largely controlled by the extreme seasonality of Antarctica and local climate. Lake Hoare and other dry valley lakes may be sensitive indicators of short term (< 100 yr) climatic and/or anthropogenic changes in the dry valleys since the onset of intensive exploration over 30 years ago. The time constants for turnover of the water column and lake ice are 50 and 10 years, respectively. The turnover time for atmospheric gases in the lake is 30–60 years. Therefore, the lake environment responds to changes on a 10–100 year timescale. Because the ice cover has a controlling influence on the lake (e.g. light penetration, gas content of water, and sediment deposition), it is probable that small changes in ice ablation, sediment loading on the ice cover, or glacial meltwater (or groundwater) inflow will affect ice cover dynamics and will have a major impact on the lake environment and biota.     

Phytoplankton population dynamics in perennially ice-covered Lake Fryxell, Antarctica

Phytoplankton were collected over five austral summers (1987–88through 1991–92) to examine seasonal and annual fluctuationsin species composition and biovolume in Lake Fryxell, a perenniallyice-covered lake located in the Dry Valleys of South VictoriaLand, Antarctica. Lake Fryxell has perennial gradients of salinity,dissolved oxygen and nutrients. We found that algal speciesdiversity was low (56 taxa were collected), confirming the resultsof previous short-term studies. The phytoplankton consistedprimarily of cryptophyte and chlorophyte flagellates, and filamentouscyanobacteria. The presence of filamentous cyanobacteria, whichhave not been reported as abundant in this lake by previousworkers, may represent a significant ecological change. Eachaustral summer, one dominant species contributed >70% ofthe total biovolume; Chroomonas lacustris was dominant in 1987–88,while Cryptomonas sp. dominated the phytoplankton in the remaining4 years. No species succession occurred during the austral summer.Some common taxa were vertically stratified (Oscillatoria limnetica,Phormidium angustissimum, Pyramimonas sp., Oscillatoria sp.),while others showed no distinct vertical stratification (Chlamydomonassubcaudata, Cryptomonas sp.). The stratification of the phytoplanktonreflects the gradients of nutrients and light, and the stabilityof the water column.     

Phytoplankton dynamics in the stratified water column of Lake Bonney,Antarctica

Phytoplankton populations in perennially ice-covered Lake Bonney, Antarctica grow in a unique non-turbulent environment. The absence of turbulence generated by winds or major streams, combined with strong vertical gradients in temperature and nutrients, create vertically stratified environmental conditions that support three discrete phytoplankton populations in the east lobe of this lake. Phytoplankton biomass and photosynthesis were measured in the east lobe of Lake Bonney during the winter-spring transicion (September) to mid-summer (January). During this period, irradiance beneath the ice increased from 0.03 to 1.9 mol quanta m⁻² d⁻¹. Chlorophylla concentrations ranged from 0.03 to 3.8 μl⁻¹ within the trophogenic zone (just beneath the permanent ice cover to 20 m) and photosynthesis ranged from below detection to 3.2 μg Cl⁻¹ d⁻¹. Our results indicate: (1) phytoplankton photosynthesis began in late winter (before 9 September, our earliest sampling date); (2) maxima for phytoplankton biomass and production developed sequentially in time from the top to the bottom of the trophogenic zone, following the seasoral increase in irradiance; and (3) the highest photosynthetic efficiencies occurred in early spring, then decreased over the remainder of the phytoplankton growth season. The spring decrease in photosynthetic rates for shallower phytoplankton appeared to be related to nutrient availability, while photosynthesis in the deeper populations was solely lightdependent.     

The geochemistry of methane in Lake Fryxell,an amictic,permanently ice-covered,antarctic lake

The abundance and distribution of dissolved CH₄ were determined from 1987–1990 in Lake Fryxell, Antarctica, an amictic, permanently ice-covered lake in which solute movement is controlled by diffusion. CH₄ concentrations were < 1 υM in the upper oxic waters, but increased below the oxycline to 936 μM at 18 m. Sediment CH₄ was 1100 μmol (1 sed)⁻¹ in the 0–5 cm zone. Upward flux from the sediment was the source of the CH₄, NH₄ ⁺, and DOC in the water column; CH₄ was 27% of the DOC+CH₄ carbon at 18 m. Incubations with surficial sediments indicated that H¹⁴CO₃ ⁻ reduction was 0.4 μmol (1 sed)⁻¹ day⁻¹ or 4× the rate of acetate fermentation to CH₄. There was no measurable CH₄ production in the water column. However, depth profiles of CH₄, NH₄, and DIC normalized to bottom water concentrations demonstrated that a significant CH₄ sink was evident in the anoxic, sulfate-containing zone of the water column (10–18 m). The δ¹³CH₄ in this zone decreased from −72 % at 18 m to −76% at 12 m, indicating that the consumption mechanism did not result in an isotopic enrichment of ¹³CH₄. In contrast, δ¹³CH₄ increased to −55 % at 9 m due to aerobic oxidation, though this was a minor aspect of the CH₄ cycle. The water column CH₄ profile was modeled by coupling diffusive flux with a first order consumption term; the best-fit rate constant for anaerobic CH₄ consumption was 0.012 yr⁻¹. On a total carbon basis, CH₄ consumption in the anoxic water column exerted a major effect on the flux of carbonaceous material from the underlying sediments and serves to exemplify the importance of CH₄ to carbon cycling in Lake Fryxell.     

High-elevation ground-layer plant community composition across environmental gradients in spruce-fir forests

We examined the influence of vegetation structure and soil chemistry on post-adelgid, ground-layer plant communities in high-elevation forests of the southern Appalachian Mountains. Specifically, we hypothesized that post-disturbance community composition and diversity would vary along a gradient of soil acidity and other soil characteristics influenced by acid deposition. Ground-layer vegetation and soils were sampled on 60 randomly located nested vegetation plots in the spruce-fir (Picea abies) zone of Great Smoky Mountains National Park, TN and NC, USA. To capture a range of deposition levels, plot placement was stratified based on modeled acid deposition classes. Ordination and multiple regression results showed that ground-layer composition and diversity were negatively associated with acidity of the A horizon and the presence of ericaceous shrubs (i.e., Rhododendron spp.). A strong correspondence between soil acidity and ericaceous shrub cover was also observed, suggesting that soil acidity may be, in conjunction with overstory disturbance resulting from chronic acid deposition and adelgid induced mortality, an important driver of ericaceous shrub thicket expansion. Slow-decaying, acidic ericaceous litter may also induce a positive feedback resulting in enhanced acidification.     

Bacterial and Archaeal Diversity in Sediments of West Lake Bonney,McMurdo Dry Valleys,Antarctica

Bacterial and archaeal diversity was examined in a sediment core from Lake Bonney, Antarctica. Members of the Archaea showed both low abundance and diversity, whereas bacterial diversity was moderately high and some phyla were fairly abundant, even in geologically old samples. Microbial diversity correlated with sample texture and differed in silty and coarse samples.     

Stromatolite records of environmental change in perennially ice-covered Lake Joyce,McMurdo Dry Valleys,Antarctica

Calcite-rich columnar stromatolites grew in perennially ice-covered Lake Joyce in the McMurdo Dry Valleys, Antarctica, during a period of environmental change associated with rising lake level. Stromatolite calcite contains carbon and oxygen isotope records of changes to microbial activity in response to variable light environments and water chemistry through time. The stromatolites grew synchronously with correlative calcite zones. The innermost (oldest) calcite zone has a wide range of δ¹³C_calcite values consistent with variable photosynthetic effects on local DIC ¹³C/¹²C. Subsequent calcite zones preserve a progressive enrichment in δ¹³C_calcite values of approximately + 2.6‰ through time, with δ¹³C_calcite values becoming less variable. This enrichment likely records the removal of ¹²C by photosynthesis from the DIC reservoir over decades, with photosynthetic effects decreasing as light levels became lower and more consistent through time. Mean δ¹⁸O_calcite values of the innermost calcified zone were at least 1‰ lower than those of the other calcified zones (t test p-level < 0.001). The significant difference in δ¹⁸O_calcite values between the innermost and other calcified zones could be a product of mixing source waters with different isotopic values associated with the initiation of lake stratification associated with rising lake level. Overall, Lake Joyce stromatolites record significant lateral variability in relative photosynthetic rate and long-lived lake water stratification with microbial modification of the DIC pool. Such processes provide criteria for interpreting microbial activity within polar paleolake deposits and may shed light on variability in lake environments associated with changing climate in the McMurdo Dry Valleys.     

Metabolic potential and survival strategies of microbial communities across extreme temperature gradients on Deception Island volcano,Antarctica

Active volcanoes in Antarctica have remarkable temperature and geochemical gradients that could select for a wide variety of microbial adaptive mechanisms and metabolic pathways. Deception Island is a stratovolcano flooded by the sea, resulting in contrasting ecosystems such as permanent glaciers and active fumaroles, which creates steep gradients that have been shown to affect microbial diversity. In this study, we used shotgun metagenomics and metagenome-assembled genomes to explore the metabolic potentials and survival strategies of microbial communities along an extreme temperature gradient in fumarole and glacier sediments on Deception Island. We observed that communities from a 98 °C fumarole were significantly enriched in genes related to hyperthermophilic (e.g. reverse gyrase, GroEL/GroES and thermosome) and oxidative stress responses, as well as genes related to sulfate reduction, ammonification and carbon fixation. Communities from <80 °C fumaroles possessed more genes related osmotic, cold- and heat-shock responses, and diverse metabolic potentials, such as those related to sulfur oxidation and denitrification, while glacier communities showed abundant metabolic potentials mainly related to heterotrophy. Through the reconstruction of genomes, we were able to reveal the metabolic potentials and different survival strategies of underrepresented taxonomic groups, especially those related to Nanoarchaeota, Pyrodictiaceae and thermophilic ammonia-oxidizing archaeal lineages.     

Biodiversity of methanogenic and other archaea in the permanently frozen Lake Fryxell, Antarctica

Archaea were detected in molecular diversity studies of the permanently frozen Lake Fryxell, Antarctica. Two clusters of methanogens were detected in the sediments, and another cluster of possibly methanotrophic Euryarchaeota was detected in the anoxic water column just above the sediments. One crenarchaeote was detected in water just below the oxycline. The Archaea present in Lake Fryxell are likely involved in the major biogeochemical cycles that occur there.     

Growth of elaborate microbial pinnacles in Lake Vanda,Antarctica

Microbial pinnacles in ice‐covered Lake Vanda, McMurdo Dry Valleys, Antarctica, extend from the base of the ice to more than 50 m water depth. The distribution of microbial communities, their photosynthetic potential, and pinnacle morphology affects the local accumulation of biomass, which in turn shapes pinnacle morphology. This feedback, plus environmental stability, promotes the growth of elaborate microbial structures. In Lake Vanda, all mats sampled from greater than 10 m water depth contained pinnacles with a gradation in size from <1‐mm‐tall tufts to pinnacles that were centimeters tall. Small pinnacles were cuspate, whereas larger ones had variable morphology. The largest pinnacles were up to ~30 cm tall and had cylindrical bases and cuspate tops. Pinnacle biomass was dominated by cyanobacteria from the morphological and genomic groups Leptolyngbya, Phormidium, and Tychonema. The photosynthetic potential of these cyanobacterial communities was high to depths of several millimeters into the mat based on PAM fluorometry, and sufficient light for photosynthesis penetrated ~5 mm into pinnacles. The distribution of photosynthetic potential and its correlation to pinnacle morphology suggests a working model for pinnacle growth. First, small tufts initiate from random irregularities in prostrate mat. Some tufts grow into pinnacles over the course of ~3 years. As pinnacles increase in size and age, their interiors become colonized by a more diverse community of cyanobacteria with high photosynthetic potential. Biomass accumulation within this subsurface community causes pinnacles to swell, expanding laminae thickness and creating distinctive cylindrical bases and cuspate tops. This change in shape suggests that pinnacle morphology emerges from a specific distribution of biomass accumulation that depends on multiple microbial communities fixing carbon in different parts of pinnacles. Similarly, complex patterns of biomass accumulation may be reflected in the morphology of elaborate ancient stromatolites.     

Diversity and distribution of sulfate-reducing bacteria in permanently frozen Lake Fryxell, McMurdo Dry Valleys, Antarctica

The permanently frozen freshwater Lake Fryxell, located in the Dry Valleys of Antarctica, exhibits an ideal geochemistry for microbial sulfate reduction. To investigate the population of sulfate-reducing bacteria in Lake Fryxell, both 16S rRNA gene and metabolic primer sets targeting the dsrA gene for the dissimilatory sulfite reductase alpha subunit were employed to analyze environmental DNA obtained from the water column and sediments of Lake Fryxell. In addition, enrichment cultures of sulfate-reducing bacteria established at 4 degrees C from Lake Fryxell water were also screened using the dsrA primer set. The sequence information obtained showed that a diverse group of sulfate-reducing prokaryotes of the domain Bacteria inhabit Lake Fryxell. With one exception, the enrichment culture sequences were not represented within the environmental sequences. Sequence data were compared with the geochemical profile of Lake Fryxell to identify possible connections between the diversity of sulfate-reducing bacteria and limnological conditions. Several clone groups were highly localized with respect to lake depth and, therefore, experienced specific physiochemical conditions. However, all sulfate-reducing bacteria inhabiting Lake Fryxell must function under the constantly cold conditions characteristic of this extreme environment.     

Seasonal cycle of the microbial plankton in Crooked Lake,Antarctica

Summary Changes in the abundance of the components of the microbial plankton between July 1990 and March 1991 in Crooked Lake, one of the largest and deepest freshwater lakes in Antarctica, are described. Chlorophyll a concentration is low (0.2–0.4g·1^–1) and there is no discernable spring increase. The phytoplankton is largely dominated by flagellates. Bacterioplankton exhibits a seasonal pattern of abundance ranging from 1.0 × 10⁸·1^–1 in July to 3.25 × 10⁸·1^–1 in September. Changes in bacterial abundance probably relate to temperature and grazing by heterotrophic and mixotrophic flagellates. Total flagellated protozoan concentrations ranged between 25–136 × 10²·l^–1. Autotrophic and heterotrophic flagellate abundances were coupled and peaks in their abundance oscillated with peaks in bacterioplankton concentration. Four species of ciliated protozoa, dominated by oligotrichs, particularly the plastidic Strombidium, inhabit the lake. The plankton is characterised by the presence of floes which act as loci for bacteria, flagellates and amoebae and feeding sites for the ciliates and the two sparce metazoan components of the plankton. Crooked Lake is extremely oligotrophic but nonetheless supports a plankton community with a low species diversity and simple trophodynamics.     

Microbial Phototrophic, Heterotrophic, and Diazotrophic Activities Associated with Aggregates in the Permanent Ice Cover of Lake Bonney, Antarctica

Abstract The McMurdo Dry Valley lakes, Antarctica, one of the Earth's southernmost ecosystems containing liquid water, harbor some of the most environmentally extreme (cold, nutrient-deprived) conditions on the planet. Lake Bonney has a permanent ice cover that supports a unique microbial habitat, provided by soil particles blown onto the lake surface from the surrounding, ice-free valley floor. During continuous sunlight summers (Nov.-Feb.), the dark soil particles are heated by solar radiation and melt their way into the ice matrix. Layers and patches of aggregates and liquid water are formed. Aggregates contain a complex cyanobacterial-bacterial community, concurrently conducting photosynthesis (CO2 fixation), nitrogen (N2) fixation, decomposition, and biogeochemical zonation needed to complete essential nutrient cycles. Aggregate-associated CO2- and N2-fixation rates were low and confined to liquid water (i.e., no detectable activities in the ice phase). CO2 fixation was mediated by cyanobacteria; both cyanobacteria and eubacteria appeared responsible for N2 fixation. CO2 fixation was stimulated primarily by nitrogen (NO3-), but also by phosphorus (PO43-). PO43- and iron (FeCl3 + EDTA) enrichment stimulated of N2 fixation. Microautoradiographic and physiological studies indicate a morphologically and metabolically diverse microbial community, exhibiting different cell-specific photosynthetic and heterotrophic activities. The microbial community is involved in physical (particle aggregation) and chemical (establishing redox gradients) modification of a nutrient- and organic matter-enriched microbial "oasis," embedded in the desertlike (i.e., nutrient depleted) lake ice cover. Aggregate-associated production and nutrient cycling represent microbial self-sustenance in a microenvironment supporting "life at the edge," as it is known on Earth.     

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).