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Item Biogeochemical and historical drivers of microbial community composition and structure in sediments from Mercer Subglacial Lake, West Antarctica(Springer Science and Business Media LLC, 2023-01) Davis, Christina L.; Venturelli, Ryan A.; Michaud, Alexander B.; Hawkings, Jon R.; Achberger, Amanda M.; Vick-Majors, Trista J.; Rosenheim, Brad E.; Dore, John E.; Steigmeyer, August; Skidmore, Mark L.; Barker, Joel D.; Benning, Liane G.; Siegfried, Matthew R.; Priscu, John C.; Christner, Brent C.; Barbante, Carlo; Bowling, Mark; Burnett, Justin; Campbell, Timothy; Collins, Billy; Dean, Cindy; Duling, Dennis; Fricker, Helen A.; Gagnon, Alan; Gardner, Christopher; Gibson, Dar; Gustafson, Chloe; Harwood, David; Kalin, Jonas; Kasic, Kathy; Kim, Ok-Sun; Krula, Edwin; Leventer, Amy; Li, Wei; Lyons, W. Berry; McGill, Patrick; McManis, James; McPike, David; Mironov, Anatoly; Patterson, Molly; Roberts, Graham; Rot, James; Trainor, Cathy; Tranter, Martyn; Winans, John; Zook, BobIce streams that flow into Ross Ice Shelf are underlain by water-saturated sediments, a dynamic hydrological system, and subglacial lakes that intermittently discharge water downstream across grounding zones of West Antarctic Ice Sheet (WAIS). A 2.06 m composite sediment profile was recently recovered from Mercer Subglacial Lake, a 15 m deep water cavity beneath a 1087 m thick portion of the Mercer Ice Stream. We examined microbial abundances, used 16S rRNA gene amplicon sequencing to assess community structures, and characterized extracellular polymeric substances (EPS) associated with distinct lithologic units in the sediments. Bacterial and archaeal communities in the surficial sediments are more abundant and diverse, with significantly different compositions from those found deeper in the sediment column. The most abundant taxa are related to chemolithoautotrophs capable of oxidizing reduced nitrogen, sulfur, and iron compounds with oxygen, nitrate, or iron. Concentrations of dissolved methane and total organic carbon together with water content in the sediments are the strongest predictors of taxon and community composition. δ¹³C values for EPS (−25 to −30‰) are consistent with the primary source of carbon for biosynthesis originating from legacy marine organic matter. Comparison of communities to those in lake sediments under an adjacent ice stream (Whillans Subglacial Lake) and near its grounding zone provide seminal evidence for a subglacial metacommunity that is biogeochemically and evolutionarily linked through ice sheet dynamics and the transport of microbes, water, and sediments beneath WAIS.Item Enhanced trace element mobilization by Earth’s ice sheets(Proceedings of the National Academy of Sciences, 2020-11) Hawkings, Jon R.; Skidmore, Mark L.; Wadham, Jemma L.; Priscu, John C.; Morton, Peter L.; Hatton, Jade E.; Gardner, Christopher B.; Kohler, Tyler J.; Stibal, Marek; Bagshaw, Elizabeth A.; Steigmeyer, August; Barker, Joel; Dore, John E.; Lyons, W. Berry; Tranter, Martyn; Spencer, Robert G. M.Trace elements sustain biological productivity, yet the significance of trace element mobilization and export in subglacial runoff from ice sheets is poorly constrained at present. Here, we present size-fractionated (0.02, 0.22, and 0.45 µm) concentrations of trace elements in subglacial waters from the Greenland Ice Sheet (GrIS) and the Antarctic Ice Sheet (AIS). Concentrations of immobile trace elements (e.g., Al, Fe, Ti) far exceed global riverine and open ocean mean values and highlight the importance of subglacial aluminosilicate mineral weathering and lack of retention of these species in sediments. Concentrations are higher from the AIS than the GrIS, highlighting the geochemical consequences of prolonged water residence times and hydrological isolation that characterize the former. The enrichment of trace elements (e.g., Co, Fe, Mn, and Zn) in subglacial meltwaters compared with seawater and typical riverine systems, together with the likely sensitivity to future ice sheet melting, suggests that their export in glacial runoff is likely to be important for biological productivity. For example, our dissolved Fe concentration (20,900 nM) and associated flux values (1.4 Gmol y−1) from AIS to the Fe-deplete Southern Ocean exceed most previous estimates by an order of magnitude. The ultimate fate of these micronutrients will depend on the reactivity of the dominant colloidal size fraction (likely controlled by nanoparticulate Al and Fe oxyhydroxide minerals) and estuarine processing. We contend that ice sheets create highly geochemically reactive particulates in subglacial environments, which play a key role in trace elemental cycles, with potentially important consequences for global carbon cycling.Item Environmentally clean access to Antarctic subglacial aquatic environments(2020-10) Michaud, Alexander B.; Vick-Majors, Trista J.; Achberger, Amanda M.; Skidmore, Mark L.; Christner, Brent C.; Tranter, Martyn; Priscu, John C.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.Item Biogeochemical Connectivity Between Freshwater Ecosystems beneath the West Antarctic Ice Sheet and the Sub‐Ice Marine Environment(2020-03) Vick‐Majors, Trista J.; Michaud, Alexander B.; Skidmore, Mark L.; Turetta, Clara; Barbante, Carlo; Christner, Brent C.; Dore, John E.; Christianson, Knut; Mitchell, Andrew C.; Achberger, Amanda M.; Mikucki, Jill A.; Priscu, John C.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.Item Microbial oxidation as a methane sink beneath the West Antarctic Ice Sheet(2017-07) Michaud, Alexander B.; Dore, John E.; Achberger, Amanda M.; Christner, Brent C.; Mitchell, Andrew C.; Skidmore, Mark L.; Vick-Majors, Trista J.; Priscu, John C.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.Item Physiological Ecology of Microorganisms in Subglacial Lake Whillans(2016-10) Vick-Majors, Trista J.; Mitchell, Andrew C.; Achberger, Amanda M.; Christner, Brent C.; Dore, John E.; Michaud, Alexander B.; Mikucki, Jill A.; Purcell, Alicia M.; Skidmore, Mark L.; Priscu, John C.Subglacial microbial habitats are widespread in glaciated regions of our planet. Some of these environments have been isolated from the atmosphere and from sunlight for many thousands of years. Consequently, ecosystem processes must rely on energy gained from the oxidation of inorganic substrates or detrital organic matter. Subglacial Lake Whillans (SLW) is one of more than 400 subglacial lakes known to exist under the Antarctic ice sheet; however, little is known about microbial physiology and energetics in these systems. When it was sampled through its 800 m thick ice cover in 2013, the SLW water column was shallow (~2 m deep), oxygenated, and possessed sufficient concentrations of C, N, and P substrates to support microbial growth. Here, we use a combination of physiological assays and models to assess the energetics of microbial life in SLW. In general, SLW microorganisms grew slowly in this energy-limited environment. Heterotrophic cellular carbon turnover times, calculated from (3)H-thymidine and (3)H-leucine incorporation rates, were long (60 to 500 days) while cellular doubling times averaged 196 days. Inferred growth rates (average ~0.006 d(-1)) obtained from the same incubations were at least an order of magnitude lower than those measured in Antarctic surface lakes and oligotrophic areas of the ocean. Low growth efficiency (8%) indicated that heterotrophic populations in SLW partition a majority of their carbon demand to cellular maintenance rather than growth. Chemoautotrophic CO2-fixation exceeded heterotrophic organic C-demand by a factor of ~1.5. Aerobic respiratory activity associated with heterotrophic and chemoautotrophic metabolism surpassed the estimated supply of oxygen to SLW, implying that microbial activity could deplete the oxygenated waters, resulting in anoxia. We used thermodynamic calculations to examine the biogeochemical and energetic consequences of environmentally imposed switching between aerobic and anaerobic metabolisms in the SLW water column. Heterotrophic metabolisms utilizing acetate and formate as electron donors yielded less energy than chemolithotrophic metabolisms when calculated in terms of energy density, which supports experimental results that showed chemoautotrophic activity in excess of heterotrophic activity. The microbial communities of subglacial lake ecosystems provide important natural laboratories to study the physiological and biogeochemical behavior of microorganisms inhabiting cold, dark environments.Item Microbial Community Structure of Subglacial Lake Whillans, West Antarctica(2016-09) Achberger, Amanda M.; Christner, Brent C.; Michaud, Alexander B.; Priscu, John C.; Skidmore, Mark L.; Vick-Majors, Trista J.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.Item Diversity, abundance, and potential activity of nitrifying and nitrate-reducing microbial assemblages in a subglacial ecosystem(2011-05) Boyd, Eric S.; Lange, Rachel K.; Mitchell, Andrew C.; Havig, Jeff R.; Lafreniere, M. J.; Shock, Everett L.; Peters, John W.; Skidmore, Mark L.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.Item Microbial sulfur transformations in Subglacial Lake Whillans sediments(2014-11) Purcell, Alicia M.; Mikucki, Jill A.; Achberger, Amanda M.; Alekhina, Irina A.; Barbante, Carlo; Christner, Brent C.; Ghosh, Dhritiman; Michaud, Alexander B.; Mitchell, Andrew C.; Priscu, John C.; Scherer, Reed; Skidmore, Mark L.; Vick-Majors, Trista J.Diverse microbial assemblages inhabit subglacial aquatic environments. While few of these environments have been sampled, data reveal that subglacial organisms gain energy for growth from reduced minerals containing nitrogen, iron, and sulfur. Here we investigate the role of microbially mediated sulfur transformations in sediments from Subglacial Lake Whillans (SLW), Antarctica, by examining key genes involved in dissimilatory sulfur oxidation and reduction. The presence of sulfur transformation genes throughout the top 34 cm of SLW sediments changes with depth. SLW surficial sediments were dominated by genes related to known sulfur-oxidizing chemoautotrophs. Sequences encoding the adenosine-5′-phosphosulfate (APS) reductase gene, involved in both dissimilatory sulfate reduction and sulfur oxidation, were present in all samples and clustered into 16 distinct operational taxonomic units. The majority of APS reductase sequences (74%) clustered with known sulfur oxidizers including those within the “Sideroxydans” and Thiobacillus genera. Reverse-acting dissimilatory sulfite reductase (rDSR) and 16S rRNA gene sequences further support dominance of “Sideroxydans” and Thiobacillus phylotypes in the top 2 cm of SLW sediments. The SLW microbial community has the genetic potential for sulfate reduction which is supported by experimentally measured low rates (1.4 pmol cm-3d-1) of biologically mediated sulfate reduction and the presence of APS reductase and DSR gene sequences related to Desulfobacteraceae and Desulfotomaculum. Our results also infer the presence of sulfur oxidation, which can be a significant energetic pathway for chemosynthetic biosynthesis in SLW sediments. The water in SLW ultimately flows into the Ross Sea where intermediates from subglacial sulfur transformations can influence the flux of solutes to the Southern Ocean.Item Biogeochemistry and microbial diversity in the marine cavity beneath the McMurdo Ice Shelf, Antarctica(2016-03) Vick-Majors, Trista J.; Achberger, Amanda M.; Santibanez, Pamela A.; Dore, John E.; Hodson, Timothy; Michaud, Alexander B.; Christner, Brent C.; Mikucki, Jill A.; Skidmore, Mark L.; Powell, Ross; Adkins, W. Peyton; Barbante, Carlo; Mitchell, Andrew C.; Scherer, Reed; Priscu, John C.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.