Biogeochemical and historical drivers of microbial community composition and structure in sediments from Mercer Subglacial Lake, West Antarctica.

Ice 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.

Bioavailability of dissolved organic matter varies with anthropogenic landcover in the Upper Mississippi River Basin.

Anthropogenic conversion of forests and wetlands to agricultural and urban landcovers impacts dissolved organic matter (DOM) within streams draining these catchments. Research on how landcover conversion impacts DOM molecular level composition and bioavailability, however, is lacking. In the Upper Mississippi River Basin (UMRB), water from low-order streams and rivers draining one of three dominant landcovers (forest, agriculture, urban) was incubated for 28 days to determine bioavailable DOC (BDOC) concentrations and changes in DOM composition. The BDOC concentration averaged 0.49 ± 0.30 mg L-1 across all samples and was significantly higher in streams draining urban catchments (0.72 ± 0.34 mg L-1) compared to streams draining agricultural (0.28 ± 0.15 mg L-1) and forested (0.47 ± 0.17 mg L-1) catchments. Percent BDOC was significantly greater in urban (10% ± 4.4%) streams compared to forested streams (5.6% ± 3.2%), corresponding with greater relative abundances of aliphatic and N-containing aliphatic compounds in urban streams. Aliphatic compound relative abundance decreased across all landcovers during the bioincubation (average -4.1% ± 10%), whereas polyphenolics and condensed aromatics increased in relative abundance across all landcovers (average of +1.4% ± 5.9% and +1.8% ± 10%, respectively). Overall, the conversion of forested to urban landcover had a larger impact on stream DOM bioavailability in the UMRB compared to conversion to agricultural landcover. Future research examining the impacts of anthropogenic landcover conversion on stream DOM composition and bioavailability needs to be expanded to a range of spatial scales and to different ecotones, especially with continued landcover alterations.

Catchment characteristics and seasonality control the composition of microbial assemblages exported from three outlet glaciers of the Greenland Ice Sheet.

Glacial meltwater drains into proglacial rivers where it interacts with the surrounding landscape, collecting microbial cells as it travels downstream. Characterizing the composition of the resulting microbial assemblages in transport can inform us about intra-annual changes in meltwater flowpaths beneath the glacier as well as hydrological connectivity with proglacial areas. Here, we investigated how the structure of suspended microbial assemblages evolves over the course of a melt season for three proglacial catchments of the Greenland Ice Sheet (GrIS), reasoning that differences in glacier size and the proportion of glacierized versus non-glacierized catchment areas will influence both the identity and relative abundance of microbial taxa in transport. Streamwater samples were taken at the same time each day over a period of 3 weeks (summer 2018) to identify temporal patterns in microbial assemblages for three outlet glaciers of the GrIS, which differed in glacier size (smallest to largest; Russell, Leverett, and Isunnguata Sermia [IS]) and their glacierized: proglacial catchment area ratio (Leverett, 76; Isunnguata Sermia, 25; Russell, 2). DNA was extracted from samples, and 16S rRNA gene amplicons sequenced to characterize the structure of assemblages. We found that microbial diversity was significantly greater in Isunnguata Sermia and Russell Glacier rivers compared to Leverett Glacier, the latter of which having the smallest relative proglacial catchment area. Furthermore, the microbial diversity of the former two catchments continued to increase over monitored period, presumably due to increasing hydrologic connectivity with proglacial habitats. Meanwhile, diversity decreased over the monitored period in Leverett, which may have resulted from the evolution of an efficient subglacial drainage system. Linear discriminant analysis further revealed that bacteria characteristic to soils were disproportionately represented in the Isunnguata Sermia river, while putative methylotrophs were disproportionately abundant in Russell Glacier. Meanwhile, taxa typical for glacierized habitats (i.e., Rhodoferax and Polaromonas) dominated in the Leverett Glacier river. Our findings suggest that the proportion of deglaciated catchment area is more influential to suspended microbial assemblage structure than absolute glacier size, and improve our understanding of hydrological flowpaths, particulate entrainment, and transport.

Benthic Dissolved Silicon and Iron Cycling at Glaciated Patagonian Fjord Heads.

Glacier meltwater supplies silicon (Si) and iron (Fe) sourced from weathered bedrock to downstream ecosystems. However, the extent to which these nutrients reach the ocean is regulated by the nature of the benthic cycling of dissolved Si and Fe within fjord systems, given the rapid deposition of reactive particulate fractions at fjord heads. Here, we examine the benthic cycling of the two nutrients at four Patagonian fjord heads through geochemical analyses of sediment pore waters, including Si and Fe isotopes (δ30Si and δ56Fe), and reaction-transport modeling for Si. A high diffusive flux of dissolved Fe from the fjord sediments (up to 0.02 mmol m-2 day-1) compared to open ocean sediments (typically <0.001 mmol m-2 day-1) is supported by both reductive and non-reductive dissolution of glacially-sourced reactive Fe phases, as reflected by the range of pore water δ56Fe (-2.7 to +0.8‰). In contrast, the diffusive flux of dissolved Si from the fjord sediments (0.02-0.05 mmol m-2 day-1) is relatively low (typical ocean values are >0.1 mmol m-2 day-1). High pore water δ30Si (up to +3.3‰) observed near the Fe(II)-Fe(III) redox boundary is likely associated with the removal of dissolved Si by Fe(III) mineral phases, which, together with high sedimentation rates, contribute to the low diffusive flux of Si at the sampled sites. Our results suggest that early diagenesis promotes the release of dissolved Fe, yet suppresses the release of dissolved Si at glaciated fjord heads, which has significant implications for understanding the downstream transport of these nutrients along fjord systems.

Globally elevated chemical weathering rates beneath glaciers.

Physical erosion and chemical weathering rates beneath glaciers are expected to increase in a warming climate with enhanced melting but are poorly constrained. We present a global dataset of cations in meltwaters of 77 glaciers, including new data from 19 Asian glaciers. Our study shows that contemporary cation denudation rates (CDRs) beneath glaciers (2174 ± 977 Σ*meq+ m-2 year-1) are ~3 times higher than two decades ago, up to 10 times higher than ice sheet catchments (~150-2000 Σ*meq+ m-2 year-1), up to 50 times higher than whole ice sheet means (~30-45 Σ*meq+ m-2 year-1) and ~4 times higher than major non-glacial riverine means (~500 Σ*meq+ m-2 year-1). Glacial CDRs are positively correlated with air temperature, suggesting glacial chemical weathering yields are likely to increase in future. Our findings highlight that chemical weathering beneath glaciers is more intense than many other terrestrial systems and may become increasingly important for regional biogeochemical cycles.

Enhanced trace element mobilization by Earth's ice sheets.

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.

Patterns in Microbial Assemblages Exported From the Meltwater of Arctic and Sub-Arctic Glaciers.

Meltwater streams connect the glacial cryosphere with downstream ecosystems. Dissolved and particulate matter exported from glacial ecosystems originates from contrasting supraglacial and subglacial environments, and exported microbial cells have the potential to serve as ecological and hydrological indicators for glacial ecosystem processes. Here, we compare exported microbial assemblages from the meltwater of 24 glaciers from six (sub)Arctic regions - the southwestern Greenland Ice Sheet, Qeqertarsuaq (Disko Island) in west Greenland, Iceland, Svalbard, western Norway, and southeast Alaska - differing in their lithology, catchment size, and climatic characteristics, to investigate spatial and environmental factors structuring exported meltwater assemblages. We found that 16S rRNA gene sequences of all samples were dominated by the phyla Proteobacteria, Bacteroidetes, and Actinobacteria, with Verrucomicrobia also common in Greenland localities. Clustered OTUs were largely composed of aerobic and anaerobic heterotrophs capable of degrading a wide variety of carbon substrates. A small number of OTUs dominated all assemblages, with the most abundant being from the genera Polaromonas, Methylophilus, and Nitrotoga. However, 16-32% of a region's OTUs were unique to that region, and rare taxa revealed unique metabolic potentials and reflected differences between regions, such as the elevated relative abundances of sulfur oxidizers Sulfuricurvum sp. and Thiobacillus sp. at Svalbard sites. Meltwater alpha diversity showed a pronounced decrease with increasing latitude, and multivariate analyses of assemblages revealed significant regional clusters. Distance-based redundancy and correlation analyses further resolved associations between whole assemblages and individual OTUs with variables primarily corresponding with the sampled regions. Interestingly, some OTUs indicating specific metabolic processes were not strongly associated with corresponding meltwater characteristics (e.g., nitrification and inorganic nitrogen concentrations). Thus, while exported assemblage structure appears regionally specific, and probably reflects differences in dominant hydrological flowpaths, OTUs can also serve as indicators for more localized microbially mediated processes not captured by the traditional characterization of bulk meltwater hydrochemistry. These results collectively promote a better understanding of microbial distributions across the Arctic, as well as linkages between the terrestrial cryosphere habitats and downstream ecosystems.

The Influence of Glacial Cover on Riverine Silicon and Iron Exports in Chilean Patagonia.

Glaciated environments have been highlighted as important sources of bioavailable nutrients, with inputs of glacial meltwater potentially influencing productivity in downstream ecosystems. However, it is currently unclear how riverine nutrient concentrations vary across a spectrum of glacial cover, making it challenging to accurately predict how terrestrial fluxes will change with continued glacial retreat. Using 40 rivers in Chilean Patagonia as a unique natural laboratory, we investigate how glacial cover affects riverine Si and Fe concentrations, and infer how exports of these bioessential nutrients may change in the future. Dissolved Si (as silicic acid) and soluble Fe (<0.02 µm) concentrations were relatively low in glacier-fed rivers, whereas concentrations of colloidal-nanoparticulate (0.02-0.45 µm) Si and Fe increased significantly as a function of glacial cover. These colloidal-nanoparticulate phases were predominately composed of aluminosilicates and Fe-oxyhydroxides, highlighting the need for size-fractionated analyses and further research to quantify the lability of colloidal-nanoparticulate species. We also demonstrate the importance of reactive particulate (>0.45 µm) phases of both Si and Fe, which are not typically accounted for in terrestrial nutrient budgets but can dominate riverine exports. Dissolved Si and soluble Fe yield estimates showed no trend with glacial cover, suggesting no significant change in total exports with continued glacial retreat. However, yields of colloidal-nanoparticulate and reactive sediment-bound Si and Fe were an order of magnitude greater in highly glaciated catchments and showed significant positive correlations with glacial cover. As such, regional-scale exports of these phases are likely to decrease as glacial cover disappears across Chilean Patagonia, with potential implications for downstream ecosystems.

Greenland melt drives continuous export of methane from the ice-sheet bed.

Ice sheets are currently ignored in global methane budgets1,2. Although ice sheets have been proposed to contain large reserves of methane that may contribute to a rise in atmospheric methane concentration if released during periods of rapid ice retreat3,4, no data exist on the current methane footprint of ice sheets. Here we find that subglacially produced methane is rapidly driven to the ice margin by the efficient drainage system of a subglacial catchment of the Greenland ice sheet. We report the continuous export of methane-supersaturated waters (CH4(aq)) from the ice-sheet bed during the melt season. Pulses of high CH4(aq) concentration coincide with supraglacially forced subglacial flushing events, confirming a subglacial source and highlighting the influence of melt on methane export. Sustained methane fluxes over the melt season are indicative of subglacial methane reserves that exceed methane export, with an estimated 6.3 tonnes (discharge-weighted mean; range from 2.4 to 11 tonnes) of CH4(aq) transported laterally from the ice-sheet bed. Stable-isotope analyses reveal a microbial origin for methane, probably from a mixture of inorganic and ancient organic carbon buried beneath the ice. We show that subglacial hydrology is crucial for controlling methane fluxes from the ice sheet, with efficient drainage limiting the extent of methane oxidation5 to about 17 per cent of methane exported. Atmospheric evasion is the main methane sink once runoff reaches the ice margin, with estimated diffusive fluxes (4.4 to 28 millimoles of CH4 per square metre per day) rivalling that of major world rivers6. Overall, our results indicate that ice sheets overlie extensive, biologically active methanogenic wetlands and that high rates of methane export to the atmosphere can occur via efficient subglacial drainage pathways. Our findings suggest that such environments have been previously underappreciated and should be considered in Earth's methane budget.

The silicon cycle impacted by past ice sheets.

Globally averaged riverine silicon (Si) concentrations and isotope composition (δ30Si) may be affected by the expansion and retreat of large ice sheets during glacial-interglacial cycles. Here we provide evidence of this based on the δ30Si composition of meltwater runoff from a Greenland Ice Sheet catchment. Glacier runoff has the lightest δ30Si measured in running waters (-0.25 ± 0.12‰), significantly lower than nonglacial rivers (1.25 ± 0.68‰), such that the overall decline in glacial runoff since the Last Glacial Maximum (LGM) may explain 0.06-0.17‰ of the observed ocean δ30Si rise (0.5-1.0‰). A marine sediment core proximal to Iceland provides further evidence for transient, low-δ30Si meltwater pulses during glacial termination. Diatom Si uptake during the LGM was likely similar to present day due to an expanded Si inventory, which raises the possibility of a feedback between ice sheet expansion, enhanced Si export to the ocean and reduced CO2 concentration in the atmosphere, because of the importance of diatoms in the biological carbon pump.

Prokaryotic assemblages in suspended and subglacial sediments within a glacierized catchment on Qeqertarsuaq (Disko Island), west Greenland.

Microbes transported by glacial meltwater streams are thought to be a product of passive dispersal from both supra- and subglacial sources, though studies investigating the origins of these assemblages are scarce. Here, we conducted a survey within a large catchment containing multiple glaciers on Qeqertarsuaq (Disko Island), west Greenland, to investigate whether meltwater-exported microbial assemblages in suspended sediments differ between glacial meltwater streams, and if they reflect corresponding bulk subglacial and extraglacial sediment communities. Using 16S rRNA gene amplicon sequencing, we found proglacial stream assemblages substantially differ from one another, despite their close spatial proximity. Furthermore, proglacial stream assemblages were composed of greater proportions of Cyanobacteria compared to bulk subglacial sediment communities, dominated by Betaproteobacteria, demonstrating large contributions of meltwater and microbial cells from supraglacial habitats. Corresponding physico-chemical characteristics of meltwater suggest that streams draining smaller glaciers had more equal contributions of both supra- and subglacial inputs compared with the main catchment outlet, aligning with observed changes in assemblage structure, such as the decreased proportion of Cyanobacteria. These results suggest that glacier size and hydrological drainage systems may influence the structure of exported microbial assemblages, and collectively provide insights into their formation and fate in thiscurrent age of deglaciation.

Ice sheets as a missing source of silica to the polar oceans.

Ice sheets play a more important role in the global silicon cycle than previously appreciated. Input of dissolved and amorphous particulate silica into natural waters stimulates the growth of diatoms. Here we measure dissolved and amorphous silica in Greenland Ice Sheet meltwaters and icebergs, demonstrating the potential for high ice sheet export. Our dissolved and amorphous silica flux is 0.20 (0.06-0.79) Tmol year-1, ∼50% of the input from Arctic rivers. Amorphous silica comprises >95% of this flux and is highly soluble in sea water, as indicated by a significant increase in dissolved silica across a fjord salinity gradient. Retreating palaeo ice sheets were therefore likely responsible for high dissolved and amorphous silica fluxes into the ocean during the last deglaciation, reaching values of ∼5.5 Tmol year-1, similar to the estimated export from palaeo rivers. These elevated silica fluxes may explain high diatom productivity observed during the last glacial-interglacial period.

Meltwater export of prokaryotic cells from the Greenland ice sheet.

Microorganisms are flushed from the Greenland Ice Sheet (GrIS) where they may contribute towards the nutrient cycling and community compositions of downstream ecosystems. We investigate meltwater microbial assemblages as they exit the GrIS from a large outlet glacier, and as they enter a downstream river delta during the record melt year of 2012. Prokaryotic abundance, flux and community composition was studied, and factors affecting community structures were statistically considered. The mean concentration of cells exiting the ice sheet was 8.30 × 104 cells mL-1 and we estimate that ∼1.02 × 1021 cells were transported to the downstream fjord in 2012, equivalent to 30.95 Mg of carbon. Prokaryotic microbial assemblages were dominated by Proteobacteria, Bacteroidetes, and Actinobacteria. Cell concentrations and community compositions were stable throughout the sample period, and were statistically similar at both sample sites. Based on our observations, we argue that the subglacial environment is the primary source of the river-transported microbiota, and that cell export from the GrIS is dependent on discharge. We hypothesise that the release of subglacial microbiota to downstream ecosystems will increase as freshwater flux from the GrIS rises in a warming world.

Ice sheets as a significant source of highly reactive nanoparticulate iron to the oceans.

The Greenland and Antarctic Ice Sheets cover ~ 10% of global land surface, but are rarely considered as active components of the global iron cycle. The ocean waters around both ice sheets harbour highly productive coastal ecosystems, many of which are iron limited. Measurements of iron concentrations in subglacial runoff from a large Greenland Ice Sheet catchment reveal the potential for globally significant export of labile iron fractions to the near-coastal euphotic zone. We estimate that the flux of bioavailable iron associated with glacial runoff is 0.40-2.54 Tg per year in Greenland and 0.06-0.17 Tg per year in Antarctica. Iron fluxes are dominated by a highly reactive and potentially bioavailable nanoparticulate suspended sediment fraction, similar to that identified in Antarctic icebergs. Estimates of labile iron fluxes in meltwater are comparable with aeolian dust fluxes to the oceans surrounding Greenland and Antarctica, and are similarly expected to increase in a warming climate with enhanced melting.