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Key Points Active, diverse microorganisms have been detected in all cryospheric habitats on Earth, and they are as abundant as those living in freshwater habitats. Numerous intracellular and extracellular adaptations enable microorganisms to thrive at temperatures below 0 °C, and thus to survive and grow in liquid inclusions within porous ice matrices. The many different types of cryospheric habitats pose distinct habitability challenges to resident microorganisms. Communities specialized to each type of ice environment use different strategies to fulfil their energy and growth requirements, using either sunlight, or inorganic or organic compounds as energy sources. Today, the use of next-generation sequencing approaches enables more detailed insights into microbial community composition and function, and allows the tracking of structural and functional differences and changes at a high resolution. Climate change is altering the cryosphere; it is expected that this will lead to shifts in the distribution, composition and activity of cold-adapted microorganisms. In this Review, Boetius et al . summarize our current knowledge of the microbial ecology of Earth's frozen realms, including sea ice and glacial habitats. They describe the diversity of niches, the composition of microbial communities at these sites and their biogeochemical activities. The Earth's cryosphere comprises those regions that are cold enough for water to turn into ice. Recent findings show that the icy realms of polar oceans, glaciers and ice sheets are inhabited by microorganisms of all three domains of life, and that temperatures below 0 °C are an integral force in the diversification of microbial life. Cold-adapted microorganisms maintain key ecological functions in icy habitats: where sunlight penetrates the ice, photoautotrophy is the basis for complex food webs, whereas in dark subglacial habitats, chemoautotrophy reigns. This Review summarizes current knowledge of the microbial ecology of frozen waters, including the diversity of niches, the composition of microbial communities at these sites and their biogeochemical activities.

There has been active debate over microbial life in Antarctic subglacial lakes owing to a paucity of direct observations from beneath the ice sheet and concerns about contamination in the samples that do exist; here the authors present the first geomicrobiological description of pristine water and surficial sediments from Subglacial Lake Whillans, and show that the lake water contains a diverse microbial community, many members of which are closely related to chemolithoautotrophic bacteria and archaea. Abundant microbes in subglacial Lake Whillans Whether there is microbial life in subglacial lakes in the Antarctic has been a matter of controversy, as early results were compromised when it was discovered that contamination may have occurred during drilling. Discovered less than a decade ago using satellite data, Lake Whillans lies beneath some 800 metres 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. In the first study to sample Antarctic subglacial waters directly, analysis of sediments obtained by the WISSARD drilling program shows that Lake Whillans' water contains more than 3,900 different types of bacteria and archaea, including one closely related to the nitrite oxidizing betaproteobacterium ' Candidatus Nitrotoga arctica', which comprised 13% of the sequence data. The lake waters contain a diverse range of metabolically active microorganisms, many of which seem to gain nutrients from the melting ice and from the rock and sediment beneath the ice. Liquid water has been known to occur beneath the Antarctic ice sheet for more than 40 years 1 , but only recently have these subglacial aqueous environments been recognized as microbial ecosystems that may influence biogeochemical transformations on a global scale 2 , 3 , 4 . 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 5 . 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 2 , 4 that can mobilize elements from the lithosphere 6 and influence Southern Ocean geochemical and biological systems 7 .

Knowledge of past ice sheet configurations is useful for informing projections of future ice sheet dynamics and for calibrating ice sheet models. The topology of grounding line retreat in the Ross Sea sector of Antarctica has been much debated, but it has generally been assumed that the modern ice sheet is as small as it has been for more than 100 000 years (Conway et al., 1999; Lee et al., 2017; Lowry et al., 2019; McKay et al., 2016; Scherer et al., 1998). Recent findings suggest that the West Antarctic Ice Sheet (WAIS) grounding line retreated beyond its current location earlier in the Holocene and subsequently readvanced to reach its modern position (Bradley et al., 2015; Kingslake et al., 2018). Here, we further constrain the post-LGM (Last Glacial Maximum) grounding line retreat and readvance in the Ross Sea sector using a two-phase model of radiocarbon input and decay in subglacial sediments from six sub-ice sampling locations. In addition, we reinterpret high basal temperature gradients, measured previously at three sites in this region (Engelhardt, 2004), which we explain as resulting from recent ice shelf re-grounding accompanying grounding line readvance. At one location – Whillans Subglacial Lake (SLW) – for which a sediment porewater chemistry profile is known, we estimate the grounding line readvance by simulating ionic diffusion. Collectively, our analyses indicate that the grounding line retreated over SLW 4300-2500+1500 years ago, and over sites on Whillans Ice Stream (WIS), Kamb Ice Stream (KIS), and Bindschadler Ice Stream (BIS) 4700-2300+1500, 1800-700+2700, and 1700-600+2800 years ago, respectively. The grounding line only recently readvanced back over those sites 1100-100+200, 1500-200+500, 1000-300+200, and 800±100 years ago for SLW, WIS, KIS, and BIS, respectively. The timing of grounding line retreat coincided with a warm period in the mid-Holocene to late Holocene. Conversely, grounding line readvance is coincident with cooling climate in the last 1000–2000 years. Our estimates for the timing of grounding line retreat and readvance are also consistent with relatively low carbon-to-nitrogen ratios measured in our subglacial sediment samples (suggesting a marine source of organic matter) and with the lack of grounding zone wedges in front of modern grounding lines. Based on these results, we propose that the Siple Coast grounding line motions in the mid-Holocene to late Holocene were primarily driven by relatively modest changes in regional climate, rather than by ice sheet dynamics and glacioisostatic rebound, as hypothesized previously (Kingslake et al., 2018).

An active microbial assemblage cycles sulfur in a sulfate-rich, ancient marine brine beneath Taylor Glacier, an outlet glacier of the East Antarctic Ice Sheet, with Fe(III) serving as the terminal electron acceptor. Isotopic measurements of sulfate, water, carbonate, and ferrous iron and functional gene analyses of adenosine 5′-phosphosulfate reductase imply that a microbial consortium facilitates a catalytic sulfur cycle. These metabolic pathways result from a limited organic carbon supply because of the absence of contemporary photosynthesis, yielding a subglacial ferrous brine that is anoxic but not sulfidic. Coupled biogeochemical processes below the glacier enable subglacial microbes to grow in extended isolation, demonstrating how analogous organic-starved systems, such as Neoproterozoic oceans, accumulated Fe(II) despite the presence of an active sulfur cycle.

Taylor Glacier hosts an active englacial hydrologic system that feeds Blood Falls, a supraglacial outflow of iron-rich subglacial brine at the terminus, despite mean annual air temperatures of −17°C and limited surface melt. Taylor Glacier is an outlet glacier of the East Antarctic ice sheet that terminates in Lake Bonney, McMurdo Dry Valleys. To image and map the brine feeding Blood Falls, we used radio echo sounding to delineate a subhorizontal zone of englacial brine upstream from Blood Falls and elongated in the ice flow direction. We estimate volumetric brine content in excess of 13% within 2 m of the central axis of this zone, and likely much higher at its center. Brine content decreases, but remains detectable, up to 45 m away along some transects. Hence, we infer a network of subparallel basal crevasses allowing injection of pressurized subglacial brine into the ice. Subglacial brine is routed towards Blood Falls by hydraulic potential gradients associated with deeply incised supraglacial valleys. The brine remains liquid within the subglacial and englacial environments through latent heat of freezing coupled with elevated salt content. Our findings suggest that cold glaciers could support freshwater hydrologic systems through localized warming by latent heat alone.

Antarctic subice environments are diverse, underexplored microbial habitats. Here, we describe the ecophysiology and annotated genome of a Marinobacter strain isolated from a cold, saline, iron-rich subglacial outflow of the Taylor Glacier, Antarctica. This strain (BF04_CF4) grows fastest at neutral pH (range 6-10), is psychrophilic (range: 0°C-20°C), moderately halophilic (range: 0.8%-15% NaCl) and hosts genes encoding potential low temperature and high salt adaptations. The predicted proteome suggests it utilizes fewer charged amino acids than a mesophilic Marinobacter strain. BF04_CF4 has increased concentrations of membrane unsaturated fatty acids including palmitoleic (33%) and oleic (27.5%) acids that may help maintain cell membrane fluidity at low temperatures. The genome encodes proteins for compatible solute biosynthesis and transport, which are known to be important for growth in saline environments. Physiological verification of predicted metabolic functions demonstrate BF04_CF4 is capable of denitrification and may facilitate iron oxidation. Our data indicate that strain BF04_CF4 represents a new Marinobacter species, Marinobacter gelidimuriae sp. nov., that appears well suited for the subglacial environment it was isolated from. Marinobacter species have been isolated from other cold, saline environments in the McMurdo Dry Valleys and permanently cold environments globally suggesting that this lineage is cosmopolitan and ecologically relevant in icy brines.

Previous studies of the lakes of the McMurdo Dry Valleys have attempted to constrain lake level history, and results suggest the lakes have undergone hundreds of meters of lake level change within the last 20 000 years. Past studies have utilized the interpretation of geologic deposits, lake chemistry, and ice sheet history to deduce lake level history; however a substantial amount of disagreement remains between the findings, indicating a need for further investigation using new techniques. This study utilizes a regional airborne resistivity survey to provide novel insight into the paleohydrology of the region. Mean resistivity maps revealed an extensive brine beneath the Lake Fryxell basin, which is interpreted as a legacy groundwater signal from higher lake levels in the past. Resistivity data suggest that active permafrost formation has been ongoing since the onset of lake drainage and that as recently as 1500–4000 years BP, lake levels were over 60 m higher than present. This coincides with a warmer-than-modern paleoclimate throughout the Holocene inferred by the nearby Taylor Dome ice core record. Our results indicate Mid to Late Holocene lake level high stands, which runs counter to previous research finding a colder and drier era with little hydrologic activity throughout the last 5000 years.

The water of the ice-covered lakes of the McMurdo Dry Valleys is derived primarily from glacial melt streams and to a lesser extent permafrost seeps and subglacial outflow. The result is a mixture of radiocarbon ages that reflect both the end-member water source and the biogeochemical processing of waters as they migrate to the lake-water column. Samples were collected from various locations within perennially ice-covered Antarctic lakes and the streams that feed them, and they were analyzed for radiocarbon abundance of organic and inorganic carbon. Stream gradient and length were shown to affect the degree of equilibration of water with the modern atmosphere prior to entering the lakes. Stream microbial mats assimilate inorganic carbon flowing over them. Seasonal ice-free ‘moat’ water dissolved inorganic carbon (DIC) is largely dependent on the amount of meltwater input from streams (modern) vs. that from direct glaciers input (old). Under the ice cover, 14C ages of lake-water DIC and organic matter are dependent on lake history, composition, and quantity of particulate matter fallout. Bottom waters of the west lobe of Lake Bonney have a DIC age of ~ 27,000 14C yr before present, which we believe are the most radiocarbon-deficient lake waters on Earth. Comparison of the radiocarbon profiles in the two lobes of Lake Bonney, along with previously published geochemical data, provides a new chronology of the evolution of these two waterbodies and shows that currently deep saline water is being displaced over the sill separating them.