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Liquid water has been known to occur beneath the Antarctic ice sheet for more than 40 years, but only recently have these subglacial aqueous environments been recognized as microbial ecosystems that may influence biogeochemical transformations on a global scale. Here we present the first geomicrobiological description of water and surficial sediments obtained from direct sampling of a subglacial Antarctic lake. Subglacial Lake Whillans (SLW) lies beneath approximately 800 m of ice on the lower portion of the Whillans Ice Stream (WIS) in West Antarctica and is part of an extensive and evolving subglacial drainage network. The water column of SLW contained metabolically active microorganisms and was derived primarily from glacial ice melt with solute sources from lithogenic weathering and a minor seawater component. Heterotrophic and autotrophic production data together with small subunit ribosomal RNA gene sequencing and biogeochemical data indicate that SLW is a chemosynthetically driven ecosystem inhabited by a diverse assemblage of bacteria and archaea. Our results confirm that aquatic environments beneath the Antarctic ice sheet support viable microbial ecosystems, corroborating previous reports suggesting that they contain globally relevant pools of carbon and microbes that can mobilize elements from the lithosphere and influence Southern Ocean geochemical and biological systems.

The debris-rich basal ice layers of a high Arctic glacier were shown to contain metabolically diverse microbes that could be cultured oligotrophically at low temperatures (0.3 to 4°C). These organisms included aerobic chemoheterotrophs and anaerobic nitrate reducers, sulfate reducers, and methanogens. Colonies purified from subglacial samples at 4°C appeared to be predominantly psychrophilic. Aerobic chemoheterotrophs were metabolically active in unfrozen basal sediments when they were cultured at 0.3°C in the dark (to simulate nearly in situ conditions), producing 14 CO 2 from radiolabeled sodium acetate with minimal organic amendment (≥38 μM C). In contrast, no activity was observed when samples were cultured at subfreezing temperatures (≤−1.8°C) for 66 days. Electron microscopy of thawed basal ice samples revealed various cell morphologies, including dividing cells. This suggests that the subglacial environment beneath a polythermal glacier provides a viable habitat for life and that microbes may be widespread where the basal ice is temperate and water is present at the base of the glacier and where organic carbon from glacially overridden soils is present. Our observations raise the possibility that in situ microbial production of CO 2 and CH 4 beneath ice masses (e.g., the Northern Hemisphere ice sheets) is an important factor in carbon cycling during glacial periods. Moreover, this terrestrial environment may provide a model for viable habitats for life on Mars, since similar conditions may exist or may have existed in the basal sediments beneath the Martian north polar ice cap.

Measurements made at John Evans Glacier, eastern Ellesmere Island, Nunavut, Canada in 1994 and 1996 provide new insight into the internal hydrology of polythermal glaciers. During the early part of each melt season, supraglacial waters enter the glacier via a crevasse field about 4 km from the terminus and are stored in a subglacial reservoir. Release of the water from the reservoir occurs initially via an artesian fountain on the glacier surface and by upwelling of waters through subglacial sediments at the terminus (event1). Channelization of the subglacial waters then occurs and water is discharged as an outburst flood (event 2), which releases a considerably larger volume of water than event 1. Thus, drainage of the subglacial reservoir follows a cyclical pattern in which discharge oscillations increase in amplitude over time. The cycle may end with complete reservoir drainage. The total volume of water released was much greater in 1994 than in 1996, primarily because the 1994 melt season was longer and warmer than the 1996 season. Interannual differences in the form of the outflow hydrographs, and in the extent and timing of connections between the subglacial reservoir and marginal melt streams, are linked to variations in the size and rate of growth of the subglacial reservoir. This hydrological behaviour may have important implications for the dynamics of polythermal glaciers.

Geologic, chemical and isotopic evidence indicate that Earth has experienced numerous intervals of widespread glaciation throughout its history, with roughly 11% of present day Earth’s land surface covered in ice. Despite the pervasive nature of glacial ice both today and in Earth’s past and the potential contribution of these systems to global biogeochemical cycles, the composition and phylogenetic structure of an active microbial community in subglacial systems has yet to be described. Here, using RNA-based approaches, we demonstrate the presence of active and endogenous archaeal, bacterial and eukaryal assemblages in cold (0–1 °C) subglacial sediments sampled from Robertson Glacier, Alberta, Canada. Patterns in the phylogenetic structure and composition of subglacial sediment small subunit (SSU) ribosomal RNA (rRNA) assemblages indicate greater diversity and evenness than in glacial surface environments, possibly due to facilitative or competitive interactions among populations in the subglacial environment. The combination of phylogenetically more even and more diverse assemblages in the subglacial environment suggests minimal niche overlap and optimization to capture a wider spectrum of the limited nutrients and chemical energy made available from weathering of bedrock minerals. The prevalence of SSU rRNA affiliated with lithoautotrophic bacteria, autotrophic methane producing archaea and heterotrophic eukarya in the subglacial environment is consistent with this hypothesis and suggests an active contribution to the global carbon cycle. Collectively, our findings demonstrate that subglacial environments harbor endogenous active ecosystems that have the potential to impact global biogeochemical cycles over extended periods of time.

In this paper we examine the degree to which Michigan’s property value assessment growth cap has eroded the tax base and created substantial differences in effective tax rates among residential properties within the City of Detroit. While the analysis focuses on a specific city with significant tax base erosion challenges, it is relevant to other cities in Michigan and across the nation, particularly in states that impose assessment growth limits. Using quantile regression techniques, we examine how an assessment growth cap alters effective tax rate distributions within and across property value groups. Results show that the cap creates a wide range of effective tax rates across properties of similar value (horizontal inequity), and similar tax payments for properties of differing values (vertical inequity).

Polycrystalline ice, as found in glaciers and the ice sheets of Antarctica, is a low porosity porous media consisting of a complicated and dynamic pore structure of liquid-filled intercrystalline veins within a solid ice matrix. In this work, Nuclear Magnetic Resonance measurements of relaxation rates and molecular diffusion, useful for probing pore structure and transport dynamics in porous systems, were used to physically characterize the unfrozen vein network structure in ice and its response to the presence of metabolic products produced by V3519-10, a cold tolerant microorganism isolated from the Vostok ice core. Recent research has found microorganisms that can remain viable and even metabolically active within icy environments at sub-zero temperatures. One potential mechanism of survival for V3519-10 is secretion of an extracellular ice binding protein that binds to the prism face of ice crystals and inhibits ice recrystallization, a coarsening process resulting in crystal growth with ice aging. Understanding the impact of ice binding activity on the bulk vein network structure in ice is important to modeling of frozen geophysical systems and in development of ice interacting proteins for biotechnology applications, such as cryopreservation of cell lines, and manufacturing processes in food sciences. Here, we present the first observations of recrystallization inhibition in low porosity ice containing V3519-10 extracellular protein extract as measured with Nuclear Magnetic Resonance and Magnetic Resonance Imaging.

Cryopreservation of microorganisms in ancient glacial ice is possible if lethal levels of macromolecular damage are not incurred and cellular integrity is not compromised via intracellular ice formation or recrystallization. Previously, a bacterium (isolate 3519-10) recovered from a depth of 3,519 m below the surface in the Vostok ice core was shown to secrete an ice-binding protein (IBP) that inhibits the recrystallization of ice. To explore the advantage that IBPs confer to ice-entrapped cells, experiments were designed to examine the expression of 3519-10's IBP gene and protein at different temperatures, assess the effect of the IBP on bacterial viability in ice, and determine how the IBP influences the physical structure of the ice. Total RNA isolated from cultures grown between 4 and 25°C and analyzed by reverse transcription-PCR indicated constitutive expression of the IBP gene. Sodium dodecyl sulfate-polyacrylamide gel electrophoretic analysis of 3519-10's extracellular proteins revealed a polypeptide of the predicted size of the 54-kDa IBP at all temperatures tested. In the presence of 100 μg mL(-1) of extracellular protein from 3519-10, the survival of Escherichia coli was increased by greater than 100-fold after 5 freeze-thaw cycles. Microscopic analysis of ice formed in the presence of the IBP indicated that per square millimeter field of view, there were ~5 times as many crystals as in ice formed in the presence of washed 3519-10 cells and non-IBP producing bacteria, and ~10 times as many crystals as in filtered deionized water. Presumably, the effect that the IBP has on bacterial viability and ice crystal structure is due to its activity as an inhibitor of ice recrystallization. A myriad of molecular adaptations are likely to play a role in bacterial persistence under frozen conditions, but the ability of 3519-10's IBP to control ice crystal structure, and thus the liquid vein network within the ice, may provide one explanation for its successful survival deep within the Antarctic ice sheet for thousands of years.

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 ((14) C-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.

The search for life on Mars is a priority for NASA’s Science Mission Directorate, a pivotal question of the Astrobiology Program, and the ultimate goal of the Mars Exploration Program (MEPAG, 2008). Also, assessment of the presence or absence of life on Mars is a prerequisite for human exploration in that it will allow mitigation of potential threats to planetary protection. Nevertheless, the Viking missions remain the first, and only, attempt to search for life on the planet (or anywhere else beyond the confines of Earth). Since Viking, and specially in the last decade, robotic missions to Mars have focused on characterizing the physical, chemical, and geological environment as a necessary step to an efficient and systematic search for life. These missions have provided an unprecedented amount of data, and our knowledge of Mars has grown vastly, with respect to its current conditions and geological evolution. In the wake of 15 years of extraordinary efforts, and at a time when the next Decadal Survey is being drafted with recommendations and plans for the exploration of the Solar System, it would seem that the search for life on Mars would be a high priority. However, based on a preliminary disclosure of the main points included in the Decadal Survey during the Astrobiology Science Conference (AbSciCon, April 26–29, 2010, League City, Texas), in situ life-detection missions to Mars will not be recommended. Instead, a Mars Sample Return mission (2018–2022) will play a preeminent role as the only mission in the next 10 years, excluding the Mars Science Laboratory (2011), with the specific task of searching for evidence of past life on Mars. This programmatic strategy is in line with the general consensus reached after the Viking missions, which was that conditions on, or near, the surface of Mars are prohibitive for microbial activity. This assumption has been reinforced with each follow-up mission and is largely based on the fact that Viking, and more recently Phoenix, failed to detect any organic compounds in the surface soils. The constantly low temperatures and hyper-arid conditions on the surface, together with high doses of radiation and the formation of reactive oxidants in the atmosphere, conspire against an extant martian biota in surface soils. Given these observations, the search for extant life has received little support and has not been considered a programmatic priority, and it is on the verge of being removed altogether from the next Decadal Survey. We argue that the strategy for Mars exploration should center on the search for extant life. By extant life, we mean life that is active today or was active during the recent geological past and is now dormant. As we discuss below, the immediate strategy for Mars exploration cannot focus only on past life based on the results of the Viking missions, particularly given that recent analyses call for a re-evaluation of some of these results. It also cannot be based on the assumption that the surface of Mars is uniformly prohibitive for extant life, since research conducted in the past 30 years in extreme environments on Earth has shown that life is possible under extremes of cold and dryness. For these reasons, we recommend a new strategy for the next decade of robotic investigations on Mars, one in which the in situ search for extant life is the first priority. Given the most updated knowledge we have about Mars’ environmental evolution, we call for a long-term architecture of the Mars Exploration Program that is organized around three main goals in the following order of priority: (1) the search for extant life; (2) the search for past life; and (3) sample return. We argue that this is the most efficient approach by which to address, with a high degree of certainty, the question as to whether life exists on Mars.

[1] The basal regions of continental ice sheets are gaps in our current understanding of the Earth’s biosphere and biogeochemical cycles. We draw on existing and new chemical data sets for subglacial meltwaters to provide the first comprehensive assessment of sub‐ice sheet biogeochemical weathering. We show that size of the ice mass is a critical control on the balance of chemical weathering processes and that microbial activity is ubiquitous in driving dissolution. Carbonate dissolution fueled by sulfide oxidation and microbial CO2 dominate beneath small valley glaciers. Prolonged meltwater residence times and greater isolation characteristic of ice sheets lead to the development of anoxia and enhanced silicate dissolution due to calcite saturation. We show that sub‐ice sheet environments are highly geochemically reactive and should be considered in regional and global solute budgets. For example, calculated solute fluxes from Antarctica (72–130 t yr −1 ) are the same order of magnitude as those from some of the world’s largest rivers and rates of chemical weathering (10–17 t km −2 yr −1 ) are high for the annual specific discharge (2.3–4.1 × 10 −3 m). Our model of chemical weathering dynamics provides important information on subglacial biodiversity and global biogeochemical cycles and may be used to design strategies for the first sampling of Antarctic Subglacial Lakes and other sub‐ice sheet environments for the next decade.

Recently, new data have been acquired on the chemical composition of waters within the glacial till beneath the Kamb Ice Stream (KIS, formerly Ice Stream C) and the Bindschadler Ice Stream (BIS, formerly Ice Stream D), two of the arteries of ice loss from the Antarctic Ice Sheet (AIS). We compare this water chemistry with basally stored subglacial waters from Arctic and Alpine glaciers and the only other measurements of sub-ice sheet chemistry to date, that emerging from beneath Law Dome near Casey Station, SW Antarctica, during a jokulhlaup. We find that the chemistries pose significant questions about the hydrology and biogeochemistry of ice sheet beds. Copyright © 2010 John Wiley & Sons, Ltd.

Subglacial environments, particularly those that lie beneath polar ice sheets, are beginning to be recognized as an important part of Earth's biosphere. However, except for indirect indications of microbial assemblages in subglacial Lake Vostok, Antarctica, no sub-ice sheet environments have been shown to support microbial ecosystems. Here we report 16S rRNA gene and isolate diversity in sediments collected from beneath the Kamb Ice Stream, West Antarctic Ice Sheet and stored for 15 months at 4 degrees C. This is the first report of microbes in samples from the sediment environment beneath the Antarctic Ice Sheet. The cells were abundant ( approximately 10(7) cells g(-1)) but displayed low diversity (only five phylotypes), likely as a result of enrichment during storage. Isolates were cold tolerant and the 16S rRNA gene diversity was a simplified version of that found in subglacial alpine and Arctic sediments and water. Although in situ cell abundance and the extent of wet sediments beneath the Antarctic ice sheet can only be roughly extrapolated on the basis of this sample, it is clear that the subglacial ecosystem contains a significant and previously unrecognized pool of microbial cells and associated organic carbon that could potentially have significant implications for global geochemical processes.

In 2006, we sampled the anoxic bottom waters of a volcanic lake beneath the Vatnajokull ice cap (Iceland). The sample contained 5 x 10(5) cells per ml, and whole-cell fluorescent in situ hybridization (FISH) and PCR with domain-specific probes showed these to be essentially all bacteria, with no detectable archaea. Pyrosequencing of the V6 hypervariable region of the 16S ribosomal RNA gene, Sanger sequencing of a clone library and FISH-based enumeration of four major phylotypes revealed that the assemblage was dominated by a few groups of putative chemotrophic bacteria whose closest cultivated relatives use sulfide, sulfur or hydrogen as electron donors, and oxygen, sulfate or CO(2) as electron acceptors. Hundreds of other phylotypes are present at lower abundance in our V6 tag libraries and a rarefaction analysis indicates that sampling did not reach saturation, but FISH data limit the remaining biome to

[1] Glacial meltwaters draining shield bedrock under the Greenland Ice Sheet (GIS) contain

Biological ice nucleators (IN) function as catalysts for freezing at relatively warm temperatures (warmer than −10 °C). We examined the concentration (per volume of liquid) and nature of IN in precipitation collected from Montana and Louisiana, the Alps and Pyrenees (France), Ross Island (Antarctica), and Yukon (Canada). The temperature of detectable ice-nucleating activity for more than half of the samples was ≥ −5 °C based on immersion freezing testing. Digestion of the samples with lysozyme (i.e., to hydrolyze bacterial cell walls) led to reductions in the frequency of freezing (0–100%); heat treatment greatly reduced (95% average) or completely eliminated ice nucleation at the measured conditions in every sample. These behaviors were consistent with the activity being bacterial and/or proteinaceous in origin. Statistical analysis revealed seasonal similarities between warm-temperature ice-nucleating activities in snow samples collected over 7 months in Montana. Multiple regression was used to construct models with biogeochemical data [major ions, total organic carbon (TOC), particle, and cell concentration] that were accurate in predicting the concentration of microbial cells and biological IN in precipitation based on the concentration of TOC, Ca 2+ , and NH 4 + , or TOC, cells, Ca 2+ , NH 4 + , K + , PO 4 3− , SO 4 2− , Cl − , and HCO 3 − . Our results indicate that biological IN are ubiquitous in precipitation and that for some geographic locations the activity and concentration of these particles is related to the season and precipitation chemistry. Thus, our research suggests that biological IN are widespread in the atmosphere and may affect meteorological processes that lead to precipitation.

Nature Geoscience 8, 851–855 (2015); published online 29 October 2015; corrected after print 29 October 2015. In the print version of this Letter originally published, the published online date was incorrectly stated as 21 September 2015; the correct published online date is 29 October 2015. This has been corrected in all online versions of the Letter.

Viable microbes have been detected beneath several geographically distant glaciers underlain by different lithologies, but comparisons of their microbial communities have not previously been made. This study compared the microbial community compositions of samples from two glaciers overlying differing bedrock. Bulk meltwater chemistry indicates that sulfide oxidation and carbonate dissolution account for 90% of the solute flux from Bench Glacier, Alaska, whereas gypsum/anhydrite and carbonate dissolution accounts for the majority of the flux from John Evans Glacier, Ellesmere Island, Nunavut, Canada. The microbial communities were examined using two techniques: clone libraries and dot blot hybridization of 16S rRNA genes. Two hundred twenty-seven clones containing amplified 16S rRNA genes were prepared from subglacial samples, and the gene sequences were analyzed phylogenetically. Although some phylogenetic groups, including the Betaproteobacteria , were abundant in clone libraries from both glaciers, other well-represented groups were found at only one glacier. Group-specific oligonucleotide probes were developed for two phylogenetic clusters that were of particular interest because of their abundance or inferred biochemical capabilities. These probes were used in quantitative dot blot hybridization assays with a range of samples from the two glaciers. In addition to shared phyla at both glaciers, each glacier also harbored a subglacial microbial population that correlated with the observed aqueous geochemistry. These results are consistent with the hypothesis that microbial activity is an important contributor to the solute flux from glaciers.

We describe a viable microbial community in a subglacial lake within the Grímsvötn volcanic caldera, Iceland. We used a hot water drill to penetrate the 300-m ice shelf and retrieved lake water and volcanic tephra sediments. We also acquired samples of borehole water before and after penetration to the lake, overlying glacial ice and snow, and water from a nearby subaerial geothermal lake for comparative analyses. Lake water is at the freezing point and fresh (total dissolved solids = 260 mg L(-1)). Detectable numbers of cells were found in samples of the lake water column and tephra sediments: 2 x 10(4) ml(-1) and 4 x 10(7) g(-1), respectively. Plate counts document abundant cold-adapted cultivable organisms in the lake water, but not in the borehole (before penetration) or glacial ice. Denaturing gradient gel electrophoresis (DGGE) of 16S rRNA gene fragments amplified from genomic DNA extracted from Grímsvötn samples indicates that the lake community is distinct from the assemblages of organisms in borehole water (before penetration) and the overlying ice and snow. Sequencing of selected DGGE bands revealed that many sequences are highly similar to known psychrophilic organisms or cloned DNA from other cold environments. Significant uptake of 14C-labeled bicarbonate occurred in dark, low-temperature incubations of lake water samples, indicating the presence of autotrophs. Acetylene reduction assays under similar incubation conditions showed no significant nitrogen fixation potential by lake water samples. This may be a consequence of the inhibition of diazotrophy by nitrogen in the lake.

Solute fluxes from a glacier-covered alpine catchment are partitioned into components derived from sea-salt, acid aerosol, dissolution of atmospheric CO2, and crustal weathering. The bulk of solute is crustally derived. Coupled sulfide oxidation and carbonate dissolution (SO-CD) and carbonation of carbonate minerals generate approximately equal amounts of solute. Chemical denudation constitutes