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The influence of mineralogy on the assembly of microbial communities in glacial environments has been difficult to assess due to complications in isolating mineralogy from other variables. Here we assess the abundance and composition of microbial communities that colonized defined minerals incubated for 12 months in two meltwater streams (N and S) emanating from Kaldalónsjökull (Kal), a basalt-hosted glacier in Iceland. The two streams shared similar meltwater geochemistry as well as bedrock and proglacial sediment elemental compositions. Yet genomic DNA and PCR-amplifiable 16S rRNA genes were detected only in Kal S. The amount of recoverable DNA was highest for hematite incubated in Kal S and the composition of 16S rRNA genes recovered from Kal S sediments was most like those recovered from hematite and magnetite, an effect driven largely by similarities in the relative abundance of the putative hydrogenotrophic iron reducer Rhodoferax. We suggest this is attributable to comminution and weathering reactions involving exposed iron silicate minerals that generate and release hydrogen and Fe(III) that can be coupled to support microbial metabolism in Kaldalónsjökull, and possibly other basaltic habitats. The low abundance of cells in Kal N could be due to low availability of Fe(III) or another substrate.

The Subglacial Antarctic Lakes Scientific Access (SALSA) Project accessed Mercer Subglacial Lake using environmentally clean hot-water drilling to examine interactions among ice, water, sediment, rock, microbes and carbon reservoirs within the lake water column and underlying sediments. A ~0.4 m diameter borehole was melted through 1087 m of ice and maintained over ~10 days, allowing observation of ice properties and collection of water and sediment with various tools. Over this period, SALSA collected: 60 L of lake water and 10 L of deep borehole water; microbes >0.2 μm in diameter from in situ filtration of ~100 L of lake water; 10 multicores 0.32–0.49 m long; 1.0 and 1.76 m long gravity cores; three conductivity–temperature–depth profiles of borehole and lake water; five discrete depth current meter measurements in the lake and images of ice, the lake water–ice interface and lake sediments. Temperature and conductivity data showed the hydrodynamic character of water mixing between the borehole and lake after entry. Models simulating melting of the ~6 m thick basal accreted ice layer imply that debris fall-out through the ~15 m water column to the lake sediments from borehole melting had little effect on the stratigraphy of surficial sediment cores.

Subglacial Antarctic aquatic environments are important targets for scientific exploration due to the unique ecosystems they support and their sediments containing palaeoenvironmental records. Directly accessing these environments while preventing forward contamination and demonstrating that it has not been introduced is logistically challenging. The Whillans Ice Stream Subglacial Access Research Drilling (WISSARD) project designed, tested and implemented a microbiologically and chemically clean method of hot-water drilling that was subsequently used to access subglacial aquatic environments. We report microbiological and biogeochemical data collected from the drilling system and underlying water columns during sub-ice explorations beneath the McMurdo and Ross ice shelves and Whillans Ice Stream. Our method reduced microbial concentrations in the drill water to values three orders of magnitude lower than those observed in Whillans Subglacial Lake. Furthermore, the water chemistry and composition of microorganisms in the drill water were distinct from those in the subglacial water cavities. The submicron filtration and ultraviolet irradiation of the water provided drilling conditions that satisfied environmental recommendations made for such activities by national and international committees. Our approach to minimizing forward chemical and microbiological contamination serves as a prototype for future efforts to access subglacial aquatic environments beneath glaciers and ice sheets.

Subglacial Lake Whillans lies below around 800m of Antarctic ice and is isolated from fresh sources of photosynthetic organic matter to sustain life. The diverse microbial ecosystems within the lake and underlying sediments are therefore dependent on a combination of relict, overridden, marine-derived organic matter and mineral-derived energy. Here, we conduct experiments to replicate subglacial erosion involving both gentle and high-energy crushing of Subglacial Lake Whillans sediments and the subsequent addition of anoxic water. We find that substantial quantities of reduced species, including hydrogen, methane, acetate and ammonium and oxidised species such as hydrogen peroxide, sulfate and carbon dioxide are released. We propose that the concomitant presence of both hydrogen and hydrogen peroxide, alongside high concentrations of mineral surface radicals, suggests that the splitting of water on freshly abraded mineral surfaces increases the concentrations of redox pairs from rock-water reactions and could provide a mechanism to augment the energy available to microbial ecosystems. The crushing and wetting of rocks by erosion under glaciers produces chemical compounds that could fuel microbial activity, as demonstrated by experiments on material collected from Subglacial Lake Whillans in Antarctica

Mars Polar Science is an integrated, compelling system that serves as a nearby analogue to numerous other planets, supports human exploration, and habitability. Mars possesses the closest and most easily accessible layered ice deposits outside of Earth, and accessing those layers to read the climate record would be a triumph for planetary science.

E. Gaidos et al, 2020, Après Nous, le Déluge: A Human‐Triggered Jökulhlaup From a Subglacial Lake, Geophysical Research Letters, 47, Citation number, 10.1029/2020GL089876. To view the published open abstract, go to https://doi.org/10.1029/2020GL089876. Deposited by shareyourpaper.org and openaccessbutton.org. We've taken reasonable steps to ensure this content doesn't violate copyright. However, if you think it does you can request a takedown by emailing help@openaccessbutton.org.

Although subglacial aquatic environments are widespread beneath the Antarctic ice sheet, subglacial biogeochemistry is not well understood, and the contribution of subglacial water to coastal ocean carbon and nutrient cycling remains poorly constrained. The Whillans Subglacial Lake (SLW) ecosystem is upstream from West Antarctica's Gould-Siple Coast ~800 m beneath the surface of the Whillans Ice Stream. SLW hosts an active microbial ecosystem and is part of an active hydrological system that drains into the marine cavity beneath the adjacent Ross Ice Shelf. Here we examine sources and sinks for organic matter in the lake and estimate the freshwater carbon and nutrient delivery from discharges into the coastal embayment. Fluorescence-based characterization of dissolved organic matter revealed microbially driven differences between sediment pore waters and lake water, with an increasing contribution from relict humic-like dissolved organic matter with sediment depth. Mass balance calculations indicated that the pool of dissolved organic carbon in the SLW water column could be produced in 4.8 to 11.9 yr, which is a time frame similar to that of the lakes' fill-drain cycle. Based on these estimates, subglacial lake water discharged at the Siple Coast could supply an average of 5,400% more than the heterotrophic carbon demand within Siple Coast embayments (6.5% for the entire Ross Ice Shelf cavity). Our results suggest that subglacial discharge represents a heretofore unappreciated source of microbially processed dissolved organic carbon and other nutrients to the Southern Ocean.

Aquatic habitats beneath ice masses contain active microbial ecosystems capable of cycling important greenhouse gases, such as methane (CH4). A large methane reservoir is thought to exist beneath the West Antarctic Ice Sheet, but its quantity, source and ultimate fate are poorly understood. For instance, O2 supplied by basal melting should result in conditions favourable for aerobic methane oxidation. Here we use measurements of methane concentrations and stable isotope compositions along with genomic analyses to assess the sources and cycling of methane in Subglacial Lake Whillans (SLW) in West Antarctica. We show that sub-ice-sheet methane is produced through the biological reduction of CO2 using H2. This methane pool is subsequently consumed by aerobic, bacterial methane oxidation at the SLW sediment–water interface. Bacterial oxidation consumes >99% of the methane and represents a significant methane sink, and source of biomass carbon and metabolic energy to the surficial SLW sediments. We conclude that aerobic methanotrophy may mitigate the release of methane to the atmosphere upon subglacial water drainage to ice sheet margins and during periods of deglaciation. Subglacial lakes contain active microbial ecosystems capable of cycling methane. In a subglacial lake in West Antarctica, methane that is produced is subsequently consumed, limiting the potential for methane emissions during lake drainage.

In its polar layered deposits (PLD), Mars possesses a record of its recent climate, analogous to terrestrial ice sheets containing climate records on Earth. Each PLD is greater than 2 ​km thick and contains thousands of layers, each containing information on the climatic and atmospheric state during its deposition, creating a climate archive. With detailed measurements of layer composition, it may be possible to extract age, accumulation rates, atmospheric conditions, and surface activity at the time of deposition, among other important parameters; gaining the information would allow us to “read” the climate record. Because Mars has fewer complicating factors than Earth (e.g. oceans, biology, and human-modified climate), the planet offers a unique opportunity to study the history of a terrestrial planet’s climate, which in turn can teach us about our own planet and the thousands of terrestrial exoplanets waiting to be discovered. During a two-part workshop, the Keck Institute for Space Studies (KISS) hosted 38 Mars scientists and engineers who focused on determining the measurements needed to extract the climate record contained in the PLD. The group converged on four fundamental questions that must be answered with the goal of interpreting the climate record and finding its history based on the climate drivers. The group then proposed numerous measurements in order to answer these questions and detailed a sequence of missions and architecture to complete the measurements. In all, several missions are required, including an orbiter that can characterize the present climate and volatile reservoirs; a static reconnaissance lander capable of characterizing near surface atmospheric processes, annual accumulation, surface properties, and layer formation mechanism in the upper 50 ​cm of the PLD; a network of SmallSat landers focused on meteorology for ground truth of the low-altitude orbiter data; and finally, a second landed platform to access ~500 ​m of layers to measure layer variability through time. This mission architecture, with two landers, would meet the science goals and is designed to save costs compared to a single very capable landed mission. The rationale for this plan is presented below. In this paper we discuss numerous aspects, including our motivation, background of polar science, the climate science that drives polar layer formation, modeling of the atmosphere and climate to create hypotheses for what the layers mean, and terrestrial analogs to climatological studies. Finally, we present a list of measurements and missions required to answer the four major questions and read the climate record. 1. What are present and past fluxes of volatiles, dust, and other materials into and out of the polar regions? 2. How do orbital forcing and exchange with other reservoirs affect those fluxes? 3. What chemical and physical processes form and modify layers? 4. What is the timespan, completeness, and temporal resolution of the climate history recorded in the PLD?

Wet-based regions of glaciers and ice sheets are now recognized to host unique and diverse microbial communities capable of influencing global biogeochemical cycles. However, the isolated nature of subglacial environments poses limitations upon the supply of protons for chemical weathering and energy sources (electron donors/acceptors) to support in situ microbial communities. A less well recognized source of these substrates is the release of gases from mineral structures, pore spaces or fluid inclusions and the generation of gases from the breakage of mineral bonds during the mechanical breakdown of rocks by moving ice. Here, we investigate the potential release of H2, CO2, CO, and short chain hydrocarbons, particularly CH4, by glacial erosion at rates relevant to chemical weathering and microbial activity beneath glaciers. A wide range of magmatic, metamorphic, and sedimentary rocks, and subglacial sediments from glaciated catchments in Greenland, Norway and Canada were ground in the laboratory to varying grain sizes and the release of gases was measured. The volume of gas released increased as the grain size of the ground sediments decreased. The results of these laboratory experiments were used to estimate rates of catchment-scale gas release based upon estimates of long term abrasion rates at each glacier. H2 generation was calculated to be sufficient to potentially support previously estimated rates of methanogenesis in the upper centimeters of subglacial sediment at a gneissic catchment in Greenland and a sedimentary catchment in Canada. Sufficient CO2 could be released by grinding to drive as much as 20% of subglacial chemical weathering at a metamorphic catchment in Svalbard, with potential implications for the inferred quantity of CO2 drawn-down from the atmosphere by glacial weathering. Rates of CH4 generation from grinding bedrock has the potential to be greater than subglacial microbial generation in a sedimentary catchment in Canada with carbon rich bedrock, suggesting a potentially important source of CH4 for methanotrophic microorganisms. We conclude that mechanical erosion beneath a range of glaciers generates significant quantities of gases which have the potential to enhance chemical weathering and/or support subglacial microbial communities in the deep icy biosphere.

A possible analog for saline lakes on planetary ice bodies lurks beneath a Canadian Arctic ice cap., Subglacial lakes are unique environments that, despite the extreme dark and cold conditions, have been shown to host microbial life. Many subglacial lakes have been discovered beneath the ice sheets of Antarctica and Greenland, but no spatially isolated water body has been documented as hypersaline. We use radio-echo sounding measurements to identify two subglacial lakes situated in bedrock troughs near the ice divide of Devon Ice Cap, Canadian Arctic. Modeled basal ice temperatures in the lake area are no higher than −10.5°C, suggesting that these lakes consist of hypersaline water. This implication of hypersalinity is in agreement with the surrounding geology, which indicates that the subglacial lakes are situated within an evaporite-rich sediment unit containing a bedded salt sequence, which likely act as the solute source for the brine. Our results reveal the first evidence for subglacial lakes in the Canadian Arctic and the first hypersaline subglacial lakes reported to date. We conclude that these previously unknown hypersaline subglacial lakes may represent significant and largely isolated microbial habitats, and are compelling analogs for potential ice-covered brine lakes and lenses on planetary bodies across the solar system.

Ice shelves surround ∼ 75% of Antarctica's coastline and are highly sensitive to climate change; several have recently collapsed and others are predicted to in the near future. Marine waters beneath ice shelves harbor active ecosystems, while adjacent seas can be important areas of bottom water formation. Despite their oceanographic significance, logistical constraints have resulted in few opportunities to directly sample sub-ice shelf cavities. Here, we present the first data on microbial diversity and biogeochemistry beneath the McMurdo Ice Shelf (MIS) near Ross Island, Antarctica. Physicochemical profiles obtained via a 56 m deep borehole through the MIS revealed three vertically layered water masses (Antarctic Surface Water [AASW], Ice Shelf Water [ISW], and modified High Salinity Shelf Water [mHSSW]). Metabolically active, moderately diverse (Shannon diversity from 2.06 to 5.74) microbial communities were detected in the AASW and mHSSW. Heterotrophic bacterial production and dissolved organic matter concentrations were higher (12–37% and 24%, respectively) in mHSSW relative to AASW. Chemoautotrophic production was 5.3 nmol C L−1 d−1 and 6.0 nmol C L−1 d−1 in the AASW and mHSSW, respectively. Phytoplankton cells were more abundant and larger in the mHSSW sample relative to the AASW, which indicates sinking of phytoplankton produced in surface waters and, together with southerly flowing currents (0.09–0.16 m s−1), horizontal advection of phytoplankton from McMurdo Sound. Advected phytoplankton carbon together with in situ chemoautotrophic production provide important sources of organic matter and other reduced compounds to support ecosystem processes in the dark waters in the ice shelf cavity.

Due to solute impurities and freezing-point depression in polycrystalline ice, a complicated and dynamic network of liquid water forms within the solid ice matrix at the boundaries between ice crystal grains. Impurity concentrations, temperature and pressure influence this network structure and impact physical, transport and rheological properties of ice. However, the nature of this internal network structure is not fully understood. Here we utilize nuclear magnetic resonance (NMR) measurements of diffusion and magnetic relaxation to study the geometry and interconnectivity of the liquid-filled network in laboratory ice, formed from a 7 g L−1 NaCl solution, and its evolution due to recrystallization processes. Additionally, we apply these NMR measurements to observe the impact on ice microstructure of an ice-binding protein (IBP) excreted by the V3519-10 organism (Flavobacteriaceae family) isolated from the Vostok ice core in Antarctica. Recrystallization inhibition was observed as a function of IBP concentration. This work demonstrates the utility of advanced NMR techniques for applications to ice microstructure and has broader implications for understanding geophysical properties of cryospheric systems.

The oldest evidence of eukaryotes in the fossil record comes from a recurrent assemblage of morphologically differentiated late Paleoproterozoic to early Mesoproterozoic microfossils. Although widely distributed, the principal constituents of this Tappania-Dictyosphaera-Valeria assemblage have not hitherto been recognized on Laurentia. We have recovered all three taxa from a shallow-water shale succession in the early Mesoproterozoic Greyson Formation (Belt Supergroup, Montana, USA). An exceptionally preserved population of Tappania substantially expands the morphological range of this developmentally complex organism, suggesting phylogenetic placement within, or immediately adjacent to, crown-group eukaryotes. Correspondence with Tappania-bearing biotas from China, India, Australia, and Siberia demonstrates an open-ocean connection to the intracratonic Belt Basin and, along with broadly co-occurring macrofossils Grypania and Horodyskia, supports the recognition of a globally expressed biozone. The Greyson Formation, along with contiguous strata in Glacier National Park, is unique in preserving all currently confirmed taxa of early eukaryotic and macroscopic fossils.

Glacial comminution of bedrock generates fresh mineral surfaces capable of sustaining chemotrophic microbial communities under the dark conditions that pervade subglacial habitats. Geochemical and isotopic evidence suggests that pyrite oxidation is a dominant weathering process generating protons that drive mineral dissolution in many subglacial systems. Here, we provide evidence correlating pyrite oxidation with chemosynthetic primary productivity and carbonate dissolution in subglacial sediments sampled from Robertson Glacier (RG), Alberta, Canada. Quantification and sequencing of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) transcripts suggest that populations closely affiliated with Sideroxydans lithotrophicus , an iron sulfide-oxidizing autotrophic bacterium, are abundant constituents of microbial communities at RG. Microcosm experiments indicate sulfate production during biological assimilation of radiolabeled bicarbonate. Geochemical analyses of subglacial meltwater indicate that increases in sulfate levels are associated with increased calcite and dolomite dissolution. Collectively, these data suggest a role for biological pyrite oxidation in driving primary productivity and mineral dissolution in a subglacial environment and provide the first rate estimate for bicarbonate assimilation in these ecosystems. Evidence for lithotrophic primary production in this contemporary subglacial environment provides a plausible mechanism to explain how subglacial communities could be sustained in near-isolation from the atmosphere during glacial-interglacial cycles.

Sandstone and carbonate core samples were challenged with a two-phase supercritical CO2 and brine mixture to investigate the effects of chemical processes on the physical properties of these rocks during injection of CO2. The experiments were monitored in real-time for pressure, temperature, and volumetric rate discharge. Pore geometry and connectivity were characterized before and after each experimental challenge using magnetic resonance (MR) imaging and two-dimensional MR relaxation correlations. Quartz arenite sandstone cores were largely unaffected by the challenge with no measurable change in effective permeability at moderate and high temperatures (~50 °C and ~95 °C) or brine concentrations (~1 g/L and ~10 g/L). In contrast, a carbonate core sample showed evidence of significant dissolution leading to a six-fold increase in effective permeability. MR images and relaxation measurements revealed a marked increase in the volume and connectivity of pre-existing pore networks in the carbonate core. We infer that the increase in permeability in the carbonate core was enhanced by focused dissolution in the existing pore and fracture networks that enhanced fast-flow paths through the core.

While bacterial communities from McMurdo Dry Valley soils have been studied using molecular techniques, data from permafrost are particularly scarce given the logistical difficulties of sampling. This study examined the molecular diversity and culturability of bacteria in permafrost from the Taylor Valley (TV), Antarctica. A 16S rRNA gene clone library was constructed to assess bacterial diversity, while a clone library of the RNA polymerase beta subunit ( rpoB ) gene was constructed to examine amino acid composition of an essential protein-coding gene. The 16S rRNA gene clone library was dominated by Acidobacteria from Gp6 and Gemmatimonadetes . The rpoB gene clone library (created with primers designed in this study) was also dominated by Acidobacteria . The ability of sequence analyses to garner additional information about organisms represented by TV sequences was explored. Specifically, optimum growth temperature was estimated from the stem GC content of the 16S rRNA gene, while potential cold adaptations within translated rpoB sequences were assessed. These analyses were benchmarked using known psychrophiles and mesophiles. Bioinformatic analyses suggested that many TV sequences could represent organisms capable of activity at low temperatures. Plate counts confirmed that c . 103 cells per gram permafrost remained viable and were culturable, while laboratory respiration assays demonstrated that microbial activity occurred at −5 °C and peaked at 15 °C.

Rock glaciers are periglacial alpine landforms that are found in many locations worldwide. Whereas well-developed models of deformation are established for traditional alpine glaciers, rock glacier deformation is poorly understood. Geophysical data from Lone Peak Rock Glacier (LPRG), southwest Montana, USA, are paired with lidar bare-earth 1 m digital elevation model (DEM) analysis to explore potential genetic relationships between internal composition, structure and regularly spaced arcuate transverse ridges expressed at the rock glacier surface. The internal composition of LPRG is heterogeneous, with frozen debris and clean ice overlain by an unconsolidated talus mantle. Upslope-dipping, clearly distinguished reflectors in the ground-penetrating radar (GPR) longitudinal survey at LPRG correspond to transverse ridges. The spacing and slope of individual features at the surface and in the subsurface were measured and compared and are found to be similar. The structures observed at LPRG and other rock glaciers are similar to structures detected in glaciotectonically altered sediment, ice-cored moraines and other rock glacier settings. This finding suggests that transverse ridges on rock glaciers may be used as geomorphic indicators of internal deformation. This study contributes to the body of research on the application of GPR to rock glaciers, and is the first to directly pair and analyze individual surface topographic features with internal structures.

Two ∼4 m vertical sequences of basal ice were collected from tunnels dug into the northern lateral margin of Taylor Glacier, McMurdo Dry Valleys, Antarctica. In both cases the basal sequences exhibit two contrasting ice facies groups; clean (debris-free) and banded dispersed (debris-rich). Debris-rich ices exhibit elevated CO2 and depleted O2 concentrations compared to the clean facies. Bacterial cell numbers, respiration rates, and nutrient concentrations are highest in debris-rich layers. Together, our geochemical and biological data indicate that microbial heterotrophic respiration is likely occurring in situ within the basal ice matrix at ambient temperatures near −15°C. This implies that the basal ice zone of polar glaciers and larger ice sheets is a viable subglacial microbial habitat and active biome of significant volume that has not previously been considered.

Microorganisms in subglacial environments drive the chemical weathering of bedrock; however, the influence of bedrock mineralogy on the composition and activity of microbial assemblages in such environments is poorly understood. Here, using a combination of in situ mineral incubation and DNA fingerprinting techniques, we demonstrate that pyrite is the dominant mineralogical control on subglacial bacterial community structure and composition. In addition, we show that the abundance of Fe in the incubated minerals influences the development of mineral-associated biomass. The ubiquitous nature of pyrite in many common bedrock types and high SO 4 2– concentrations in most glacial meltwaters suggest that pyrite may be a dominant lithogenic control on microbial communities in many subglacial systems. Mineral-based energy may therefore serve a fundamental role in sustaining subglacial microbial populations and enabling their persistence over glacial-interglacial time scales.

Measurement of gases entrapped in clean ice from basal portions of the Taylor Glacier, Antarctica, revealed that CO2 ranged from 229 to 328 ppmv and O2 was near 20% of the gas volume. In contrast, vertically adjacent sections of the sediment laden basal ice contained much higher concentrations of CO2 (60,000 to 325,000 ppmv), whereas O2 represented 4 to 18% of the total gas volume. The deviation in gas composition from atmospheric values occurred concurrently with increased microbial cell concentrations in the basal ice profile, suggesting that in situ microbial processes (i.e., aerobic respiration) may have altered the entrapped gas composition. Molecular characterization of 16S rRNA genes amplified from samples of the basal ice indicated a low diversity of bacteria, and most of the sequences characterized (87%) were affiliated with the phylum, Firmicutes. The most abundant phylotypes in libraries from ice horizons with elevated CO2 and depleted O2 concentrations were related to the genus Paenisporosarcina, and 28 isolates from this genus were obtained by enrichment culturing. Metabolic experiments with Paenisporosarcina sp. TG14 revealed its capacity to conduct macromolecular synthesis when frozen in water derived from melted basal ice samples and incubated at −15 °C. The results support the hypothesis that the basal ice of glaciers and ice sheets are cryospheric habitats harboring bacteria with the physiological capacity to remain metabolically active and biogeochemically cycle elements within the subglacial environment.

The Whillans Ice Stream Subglacial Access Research Drilling (WISSARD) project will test the overarching hypothesis that an active hydrological system exists beneath a West Antarctic ice stream that exerts a major control on ice dynamics, and the metabolic and phylogenetic diversity of the microbial community in subglacial water and sediment. WISSARD will explore Subglacial Lake Whillans (SLW, unofficial name) and its outflow toward the grounding line where it is thought to enter the Ross Ice Shelf seawater cavity. Introducing microbial contamination to the subglacial environment during drilling operations could compromise environmental stewardship and the science objectives of the project, consequently we developed a set of tools and procedures to directly address these issues. WISSARD hot water drilling efforts will include a custom water treatment system designed to remove micron and sub-micron sized particles (biotic and abiotic), irradiate the drilling water with germicidal ultraviolet (UV) radiation, and pasteurize the water to reduce the viability of persisting microbial contamination. Our clean access protocols also include methods to reduce microbial contamination on the surfaces of cables/hoses and down-borehole equipment using germicidal UV exposure and chemical disinfection. This paper presents experimental data showing that our protocols will meet expectations established by international agreement between participating Antarctic nations.

The abundance of water at the base of glaciers and polar ice sheets forms lacustrine features and habitats in the saturated sediments of subglacial hydrological systems. Nutrients and energy sources may be made available through mineralization of stored organic matter or through glacial processes (e.g., bedrock comminution) that provide redox couples for microbial life. The logistical challenges of accessing subglacial environments has limited direct observations to a small number of locations, but microorganisms and associated microbial activities have been found in all subglacial environments examined to date (i.e., basal ice and sediment cores, subglacial lakes, and subglacial outflows at glacial margins). Molecular and biogeochemical data imply that the microbial clades common in subglacial environments are utilizing reduced iron, sulfur, and nitrogen compounds as energy sources to fuel primary production at the glacial bed. Here, we review the latest information on the diversity of subglacial environments and discuss how interactions between physical and biogeochemical processes affect microbial ecosystems and processes at the glacier bed.

Subglacial Lake Whillans (SLW) is located beneath ∼800 m of ice on the Whillans Ice Stream in West Antarctica and was sampled in January of 2013, providing the first opportunity to directly examine water and sediments from an Antarctic subglacial lake. To minimize the introduction of surface contaminants to SLW during its exploration, an access borehole was created using a microbiologically clean hot water drill designed to reduce the number and viability of microorganisms in the drilling water. Analysis of 16S rRNA genes (rDNA) amplified from samples of the drilling and borehole water allowed an evaluation of the efficacy of this approach and enabled a confident assessment of the SLW ecosystem inhabitants. Based on an analysis of 16S rDNA and rRNA (i.e., reverse-transcribed rRNA molecules) data, the SLW community was found to be bacterially dominated and compositionally distinct from the assemblages identified in the drill system. The abundance of bacteria (e.g., Candidatus Nitrotoga, Sideroxydans, Thiobacillus, and Albidiferax) and archaea (Candidatus Nitrosoarchaeum) related to chemolithoautotrophs was consistent with the oxidation of reduced iron, sulfur, and nitrogen compounds having important roles as pathways for primary production in this permanently dark ecosystem. Further, the prevalence of Methylobacter in surficial lake sediments combined with the detection of methanogenic taxa in the deepest sediment horizons analyzed (34–36 cm) supported the hypothesis that methane cycling occurs beneath the West Antarctic Ice Sheet. Large ratios of rRNA to rDNA were observed for several operational taxonomic units abundant in the water column and sediments (e.g., Albidiferax, Methylobacter, Candidatus Nitrotoga, Sideroxydans, and Smithella), suggesting a potentially active role for these taxa in the SLW ecosystem. Our findings are consistent with chemosynthetic microorganisms serving as the ecological foundation in this dark subsurface environment, providing new organic matter that sustains a microbial ecosystem beneath the West Antarctic Ice Sheet.

Geochemical data indicate that protons released during pyrite (FeS 2 ) oxidation are important drivers of mineral weathering in oxic and anoxic zones of many aquatic environments, including those beneath glaciers. Oxidation of FeS 2 under oxic, circumneutral conditions proceeds through the metastable intermediate thiosulfate (S 2 O 3 2− ), which represents an electron donor capable of supporting microbial metabolism. Subglacial meltwaters sampled from Robertson Glacier (RG), Canada, over a seasonal melt cycle revealed concentrations of S 2 O 3 2− that were typically below the limit of detection, despite the presence of available pyrite and concentrations of the FeS 2 oxidation product sulfate (SO 4 2− ) several orders of magnitude higher than those of S 2 O 3 2− . Here we report on the physiological and genomic characterization of the chemolithoautotrophic facultative anaerobe Thiobacillus sp. strain RG5 isolated from the subglacial environment at RG. The RG5 genome encodes genes involved with pathways for the complete oxidation of S 2 O 3 2− , CO 2 fixation, and aerobic and anaerobic respiration with nitrite or nitrate. Growth experiments indicated that the energy required to synthesize a cell under oxygen- or nitrate-reducing conditions with S 2 O 3 2− as the electron donor was lower at 5.1°C than 14.4°C, indicating that this organism is cold adapted. RG sediment-associated transcripts of soxB , which encodes a component of the S 2 O 3 2− -oxidizing complex, were closely affiliated with soxB from RG5. Collectively, these results suggest an active sulfur cycle in the subglacial environment at RG mediated in part by populations closely affiliated with RG5. The consumption of S 2 O 3 2− by RG5-like populations may accelerate abiotic FeS 2 oxidation, thereby enhancing mineral weathering in the subglacial environment.

Summary The habitability of icy environments may be limited by low temperature, low nutrient concentrations, high solute concentrations and the physical ice matrix. The basal ice of ice sheets and glaciers contains sediments that may be a source of nutrients for microbial activity. Here we quantify microbial respiration and active cell populations of Antarctic glacial isolates Paenisporosarcina sp. B5 and Chryseobacterium sp. V3519-10 in laboratory ices with abundant nutrients at temperatures from −4°C to −33°C. At all temperatures, initial high rates of metabolism were followed by lower rates suggestive of a non-reproductive metabolic state such as maintenance or dormancy. Metabolism was sustained by viable cells as quantified via culturability, CTC reduction and LIVE/DEAD staining. Respiration rates based on active cell populations did not correspond to rates representative of reproductive growth from the literature, but suggested lower levels of metabolism. Our data demonstrated that bacteria actively respired acetate in polycrystalline ice with abundant nutrients despite low temperatures and the physical ice matrix. Our results suggest that the debris-rich basal ice that exists at temperatures just below the freezing point and underlies portions of both the Greenland and Antarctic ice sheets represents a significant potential habitat for metabolically active microbial communities.

Subglacial sediments sampled from beneath Robertson Glacier (RG), Alberta, Canada, were shown to harbor diverse assemblages of potential nitrifiers, nitrate reducers, and diazotrophs, as assessed by amoA , narG , and nifH gene biomarker diversity. Although archaeal amoA genes were detected, they were less abundant and less diverse than bacterial amoA , suggesting that bacteria are the predominant nitrifiers in RG sediments. Maximum nitrification and nitrate reduction rates in microcosms incubated at 4°C were 280 and 18.5 nmol of N per g of dry weight sediment per day, respectively, indicating the potential for these processes to occur in situ . Geochemical analyses of subglacial sediment pore waters and bulk subglacial meltwaters revealed low concentrations of inorganic and organic nitrogen compounds. These data, when coupled with a C/N atomic ratio of dissolved organic matter in subglacial pore waters of ∼210, indicate that the sediment communities are N limited. This may reflect the combined biological activities of organic N mineralization, nitrification, and nitrate reduction. Despite evidence of N limitation and the detection of nifH , we were unable to detect biological nitrogen fixation activity in subglacial sediments. Collectively, the results presented here suggest a role for nitrification and nitrate reduction in sustaining microbial life in subglacial environments. Considering that ice currently covers 11% of the terrestrial landmass and has covered significantly greater portions of Earth at times in the past, the demonstration of nitrification and nitrate reduction in subglacial environments furthers our understanding of the potential for these environments to contribute to global biogeochemical cycles on glacial-interglacial timescales.

Summary Metabolic activity, but not growth, has been observed in ice at temperatures from −5°C to −32°C. To improve understanding of metabolism in ice, we simultaneously examined various aspects of metabolism (14C-acetate utilization, macromolecule syntheses and viability via reduction of CTC) of the glacial isolates Sporosarcina sp. B5 and Chryseobacterium sp. V3519-10 during incubation in nutrient-rich ice and brine at −5°C for 50 days. Measured rates of acetate utilization and macromolecule syntheses were high in the first 20 days suggesting adjustment to the lower temperatures and higher salt concentrations of both the liquid vein network in the ice and the brine. Following this adjustment, reproductive growth of both organisms was evident in brine, and suggested for Sporosarcina sp. B5 in ice by increases in cell numbers and biomass. Chryseobacterium sp. V3519-10 cells incubated in ice remained active. These data indicate that neither low temperature nor high salt concentrations prohibit growth in ice, but some other aspect of living within ice slows growth to within the detection limits of current methodologies. These results imply that microbial growth is plausible in natural ice systems with comparable temperatures and sufficient nutrients, such as debris-rich basal ices of glaciers and ice masses.

Boyd, E. S., Skidmore, M., Mitchell, A. C., Bakermans, C., Peters, J. W. (2010). Methanogenesis in subglacial sediemtns. Environmental Microbiology Reports, 2 (5), 685-692

Microbes live under glaciers that have persisted for millions of years, without a clear energy supply. Analyses of crushed rocks suggest that interactions of glaciers with the rocks beneath can produce enough H2 to support methanogenic bacteria. Substantial parts of the beds of glaciers, ice sheets and ice caps are at the pressure melting point1. The resulting water harbours diverse subglacial microbial ecosystems2,3 capable of affecting global biogeochemical cycles4,5. Such subglacial habitats may have acted as refugia during Neoproterozoic glaciations6. However, it is unclear how life in subglacial environments could be supported during glaciations lasting millions of years because energy from overridden organic carbon would become increasingly depleted7,8. Here we investigate the potential for abiogenic H2 produced during rock comminution to provide a continual source of energy to support subglacial life. We collected a range of silicate rocks representative of subglacial environments in Greenland, Canada, Norway and Antarctica and crushed them with a sledgehammer and ball mill to varying surface areas. Under an inert atmosphere in the laboratory, we added water, and measured H2 production with time. H2 was produced at 0 °C in all silicate–water experiments, probably through the reaction of water with mineral surface silica radicals formed during rock comminution. H2 production increased with increasing temperature or decreasing silicate rock grain size. Sufficient H2 was produced to support previously measured rates of methanogenesis under a Greenland glacier. We conclude that abiogenic H2 generation from glacial bedrock comminution could have supported life and biodiversity in subglacial refugia during past extended global glaciations.

Laboratory experiments on reagent-grade calcium carbonate and carbonate rich glacial sediments demonstrate previously unreported kinetic fractionation of carbon isotopes during the initial hydrolysis and early stages of carbonate dissolution driven by atmospheric CO2. There is preferential dissolution of Ca 12 CO3 during hydrolysis, resulting in 13 C-DIC values that are significantly lighter isotopically than the bulk carbonate. The fractionation factor for this kinetic isotopic effect is defined as carb. carb is greater on average for glacial sediments (17.4‰) than for calcium carbonate (7.8‰) for the 63 m size fraction, a sediment concentration o f5gL 1 and closed system conditions at 5°C. This difference is most likely due to the preferential dissolution of highly reactive ultra-fine particles with damaged surfaces that are common in subglacial sediments. The kinetic isotopic fractionation has a greater impact on 13 C-DIC at higher CaCO3: water ratios and is significant during at least the firs t6ho fcarbonate dissolution driven by atmospheric CO2 at sediment concentrations o f5gL 1 . Atmospheric CO2 dissolving into solution following carbonate hydrolysis does not exhibit any significant equilibrium isotopic fractionation for at least 6 h after the start of the experiment at 5°C. This is considerably longer than previously reported in the literature. Thus, kinetic fractionation processes will likely dominate the 13 C-DIC signal in natural environments where rock:water contact times are short 6 -24 h (e.g., glacial systems, headwaters in fluvial catchments) and there is an excess of carbonate in the sediments. It will be difficult apply conventional isotope mass balance techniques in these types of environment to identify microbial CO2 signatures in DIC from 13 C-DIC data. Copyright © 2004

The relationship between subglacial chemical weathering processes and the Sr isotope composition of runoff from Robertson Glacier, Alberta, Canada, is investigated. This glacier rests on predominantly carbonate bedrock of Upper Devonian age, but silicate minerals are also present. The provenance of solute in meltwaters is found to vary systematically with solute concentration and, by inference, subglacial water residence time. In dilute waters, the principal process of solute acquisition is calcite dissolution fueled by protons derived from the dissolution of CO 2 and subsequent dissociation of carbonic acid. At higher solute concentrations, dolomite dissolution coupled to sulfide oxidation is more important. Sr concentration is found to increase with total solute concentration in two separate meltwater streams draining from the glacier, but 87 Sr/ 86 Sr only increases in the eastern melt stream. Carbonate and K-feldspar sources are shown to dominate the Sr content of the western stream, irrespective of concentration. They also dominate the Sr content of the eastern stream at low and intermediate concentrations, but at higher concentrations, muscovite (with high 87 Sr/ 86 Sr) is also an important Sr source. This reflects the outcrop of muscovite-bearing lithologies in the catchment of the eastern stream and an increase in the rate of weathering of K-silicates relative to that of carbonates as more concentrated solutions approach saturation with respect to carbonates. Nonstoichiometric release of 87 Sr/ 86 Sr and preferential release of Sr over K from freshly ground K-silicate surfaces may also occur. This may help to explain the radiogenic nature of runoff from distributed subglacial drainage systems, which are characterized by long water:rock contact times and water flow through environments in which crushing and grinding of bedrock are active processes. Although the exchangeable Sr in tills has higher 87 Sr/ 86 Sr than local carbonate bedrock, only the more concentrated meltwaters from the eastern stream display similarly high values. The most dilute waters, which probably transport the bulk of the dissolved Sr flux from the glacier, have 87 Sr/ 86 Sr characteristic of local carbonate bedrock. Thus, the results suggest that although enhanced weathering of silicate minerals containing radiogenic Sr (such as muscovite) does occur in glaciated carbonate terrains, it is unlikely to contribute to any enhanced flux of radiogenic Sr from glaciated continental surfaces to the oceans.

This paper describes the physical and chemical properties of the snow- pack on John Evans Glacier, Ellesmere Island, Canadian Arctic Archipelago, and investigates the controls on snowpack solute concentrations and atmospheric deposition. The snowpack contains three layers that are traceable across the whole glacier. These represent fall accumulation that has been metamorphosed to depth hoar, winter accumulation mixed with snow reworked by wind from the underlying depth hoar, and spring accumulation mixed with wind-reworked snow. The seasonal cycle in snow chemistry closely reflects changes in the composition of the atmospheric aerosol at Alert, with some modification of NO3− concentrations by post-depositional processes. Mean water-weighted solute concentrations in the snowpack are largely independent of accumulation, while atmospheric deposition tends to increase with accumulation. This suggests that, for most species, wet deposition is the dominant depositional process throughout the year. However, concentrations of Ca2+ and K+ increase with both accumulation and elevation, implying an enhanced input from dry deposition of soil dust above 800 m elevation. Concentrations of SO42− are inversely related to accumulation, especially in the winter layer, suggesting a significant input from non-precipitating events, such as dry deposition or riming, during this period of very limited snowfall.

This chapter focuses on glaciers and ice sheets, sea ice, and ice shelves of the polar regions, i.e., those latitudes above the Arctic and Antarctic Circles where glaciers and ice sheets cover a significant proportion of the land mass and where large expanses of the surface waters of the Arctic and Southern Oceans undergo an annual cycle of freezing and melting. This chapter further introduces sea ice as a microbial habitat and summarizes from some of the aforementioned reviews what is known to date about the abundance, activity, diversity, and ecology of prokaryotic sea-ice microorganisms. It provides a brief outline of the role of microorganisms in biogeochemical cycling of elements in sea ice. The majority of bacteria isolated from sea ice are pigmented and highly cold adapted, with some able to form gas vesicles. Possible cold-adaptation strategies revealed by whole-genome sequence analysis also include the production of cryoprotective osmolytes and exopolymers. Polar ice shelves are thick masses of ice floating on the ocean. They are formed through glacial ice and ice sheets pushing onto the sea or long-term accumulations of sea ice. Analysis of ice-shelf heterotrophic bacteria and microbial eukaryotes suggests phylogenetic affiliation with taxa from diverse environments and climatic zones ranging from Antarctica and other cryosphere habitats to temperate ecozones. Microbial investigations on polar glaciers, ice sheets, and ice shelves are still largely in their infancy, with sea-ice research being somewhat more established.

Highlights • MR techniques can quantitatively characterize ice liquid vein network structure. • MR techniques can non-invasively monitor ice recrystallization processes with time. • Bacterial ice binding protein results in persistent small ice crystal structure. • MR is a new method to quantify ice structure during biotechnological processes. • Results demonstrate potential of MR in biotechnology process monitoring., Liquid water present in polycrystalline ice at the interstices between ice crystals results in a network of liquid-filled veins and nodes within a solid ice matrix, making ice a low porosity porous media. Here we used nuclear magnetic resonance (NMR) relaxation and time dependent self-diffusion measurements developed for porous media applications to monitor three dimensional changes to the vein network in ices with and without a bacterial ice binding protein (IBP). Shorter effective diffusion distances were detected as a function of increased irreversible ice binding activity, indicating inhibition of ice recrystallization and persistent small crystal structure. The modification of ice structure by the IBP demonstrates a potential mechanism for the microorganism to enhance survivability in ice. These results highlight the potential of NMR techniques in evaluation of the impact of IBPs on vein network structure and recrystallization processes; information useful for continued development of ice-interacting proteins for biotechnology applications.