The High Arctic glacial ecosystem: new insights from nutrient budgets

This paper describes detailed budgets of water, Cl⁻, dissolved Si and both inorganic and organic forms of nitrogen and phosphorus for two small glacier basins in Arctic Svalbard (Midre Lovénbreen and AustreBrøggerbreen). Rates of nutrient deposition are modest, dominated by inorganic nitrogen and episodically enhanced by extreme events. Hence deposition rates are also variable, ranging from 20 to 72 kg NO₃-N km⁻² a⁻¹ and 10–37 kg NH₄-N km⁻² a⁻¹ over just two consecutive years. Deposition of dissolved organic and particulate forms of nitrogen (DONand PN respectively) also appears significant and therefore requires further investigation (3–8 kg DON-N km⁻² and 7–26 kg PN-N km⁻² during winter – no summer data are available). Evidence for microbially mediated nutrient cycling within the glacial system is clear in the nutrient budgets, as is the release of large phosphorus, Si and organic/particulate nitrogen fluxes by subglacial erosion. The latter is entirely dependent upon the presence of subglacial drainage, promoting silicate mineral dissolution and the erosion of largely unweathered apatite. The large DON and PN fluxes are surprising and may relate to young organic nitrogen associated with microbial life within the glaciers. This is because wide spread assimilation of NH₄⁺ and perhaps even nitrification occurs on the glacier surface, most likely within abundant cryoconite holes. Further microbial activity also occurs at the glacier bed, where denitrification and sulphate reduction is now known to take place. Thus a two component ‘glacial ecosystem’ is proposed that is highly sensitive to climate change.     

Glacier ecosystem response to episodic nitrogen enrichment in Svalbard, European High Arctic

We describe the climatology, hydrology and biogeochemistry of an extreme nitrogen deposition event that occurred in the highly glacierised environment of the European High Arctic during June 1999. Meteorological analysis, three-dimensional air mass trajectories and a 3D transport model show that blocking high pressures over Scandinavia and the rapid advection of western European pollution toward Svalbard were sufficient to cause the most concentrated (1.15 ppm NO₃–N and 1.20 ppm NH₄–N), high magnitude (total 26 mm and up to 2.4 mm h^?1 at 30 m above sea level) nitrogen deposition event on record in this sensitive, high Arctic environment (78.91° N, 11.93° E). Since the event occurred when much of the catchment remained frozen or under snow cover, microbial utilisation of nitrogen within snowpacks and perennially unfrozen subglacial sediments, rather than soils, were mostly responsible for reducing N export. The rainfall event occurred long before the annual subglacial outburst flood and so prolonged (ca. 10 day) water storage at the glacier bed further enhanced the microbial assimilation. When the subglacial outburst eventually occurred, high runoff and concentrations of NO₃ ^? (but not NH₄ ⁺) returned in the downstream rivers. Assimilation accounted for between 53 and 72% of the total inorganic nitrogen deposited during the event, but the annual NO₃ ^? and NH₄ ⁺ runoff yields were still enhanced by up to 5 and 40 times respectively. Episodic atmospheric inputs of reactive nitrogen can therefore directly influence the biogeochemical functioning of High Arctic catchments, even when microbial activity takes place beneath a glacier at a time when terrestrial soil ecosystems remain frozen and unresponsive.     

Culturable Bacteria in Subglacial Sediments and Ice from Two Southern Hemisphere Glaciers

Viable prokaryotes have been detected in basal sediments beneath the few Northern Hemisphere glaciers that have been sampled for microbial communities. However, parallel studies have not previously been conducted in the Southern Hemisphere, and subglacial environments in general are a new and underexplored niche for microbes. Unfrozen subglacial sediments and overlying glacier ice samples collected aseptically from the Fox Glacier and Franz Josef Glacier in the Southern Alps of New Zealand now have been shown to harbor viable microbial populations. Total direct counts of 2–7 × 10⁶ cells g^–1 dry weight sediment were observed, whereas culturable aerobic heterotrophs ranged from 6–9 × 10⁵ colony-forming units g^–1 dry weight. Viable counts in the glacier ice typically were 3–4 orders of magnitude smaller than in sediment. Nitrate-reducing and ferric iron–reducing bacteria were detected in sediment samples from both glaciers, but were few or below detection limits in the ice samples. Nitrogen-fixing bacteria were detected only in the Fox Glacier sediment. Restriction fragment analysis of 16S rDNA amplified from 37 pure cultures of aerobic heterotrophs capable of growth at 4°C yielded 23 distinct groups, of which 11 were identified as -Proteobacteria. 16S rDNA sequences from representatives of these 11 groups were analyzed phylogenetically and shown to cluster with bacteria such as Polaromonas vacuolata and Rhodoferax antarcticus, or with clones obtained from permanently cold environments. Chemical analysis of sediment and ice samples revealed a dilute environment for microbial life. Nevertheless, both the sediment samples and one ice sample demonstrated substantial aerobic mineralization of ¹⁴C-acetate at 8°C, indicating that sufficient nutrients and viable psychrotolerant microbes were present to support metabolism. Unfrozen subglacial sediments may represent a significant global reservoir of biological activity with the potential to influence glacier meltwater chemistry.     

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.     

Methanogenic potential of Arctic and Antarctic subglacial environments with contrasting organic carbon sources

Subglacial environments are largely anoxic, contain organic carbon (OC) overridden by glacier ice during periods of advance, and harbour active microbial communities. This creates favourable conditions for OC degradation via methanogenesis. It has been hypothesized that OC beneath ice sheets is converted to methane (CH₄) and may be released to the atmosphere during retreat. However, there are limited data available to support this contention. Here, we present new data on the abundance, diversity and activity of methanogenic archaea and the amount and character of OC in subglacial sediments from Arctic and Antarctic glacial systems based on different substrate types. We employed long‐term laboratory incubations to quantify the CH₄ production potential in different subglacial settings. Significant numbers of methanogens (up to 7 × 10⁴ cells g^?1) were detected in the samples and clones of Methanomicrobiales and Methanosarcinales were identified in clone libraries. Long lag periods (up to >200 days) were observed before significant CH₄ concentrations were measured. We report order of magnitude differences in rates of CH₄ production (10¹–10⁵ fmol g^?1 d^?1) in different subglacial sediments, reflecting contrasts in the origin of the sediment and the OC character. Hence, we predict that contrasting rates of CH₄ production are likely to occur beneath glaciers and ice sheets that overran different types of substrate. We subsequently estimated the potential for CH₄ production beneath the Laurentide/Inuitian/Cordilleran and Fennoscandian Ice Sheets during a typical 85 ka Quaternary glacial/interglacial cycle. CH₄ production from lacustrine‐derived OC is likely to be an order of magnitude higher (~6.3–27 Pg C) than that from overridden soils (~0.55–0.68 Pg C), possibly due to a difference in lability between lacustrine and soil OC. While representing a fraction of the entire OC pool (~418–610 Pg C), this finding highlights the importance of considering the character of different OC pools when calculating subglacial CH₄ production.     

Comparison of Microbial Community Compositions of Two Subglacial Environments Reveals a Possible Role for Microbes in Chemical Weathering Processes

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.     

Nitrogen dynamics in two antarctic streams

The many glacier meltwater streams of southern Victoria Land flow through catchments where life forms are almost entirely microbial. Allochthonous inputs of nitrogen from two study streams near McMurdo Sound were derived mostly from the melting glaciers (ca. 100–200 mg N m^–3) with some originating from N₂-fixation by heterocystous cyanobacteria (max. 939 mg N m^–2 year^–1). Thirty to fifty per cent of the glacier derived N was dissolved organic N and a major proportion of this was identified as urea N which was utilised by the rich algal and cyanobacterial mats in the streams. A nutrient budget for Fryxell Stream was estimated, quantifying uptake of urea-N and dissolved inorganic N and the release of dissolved organic (non urea) and particulate N by the stream communities. An index of in-stream nitrogen processing, the Net Uptake Length Constant in these streams was compared with that from temperate climates and was found to be similar. Despite the influence of low temperatures on microbial activity (mean daily water temperature = 5 °C) nutrient removal rates from these antarctic streams are high because of the large standing stock of microbial biomass there.     

Microbial diversity of an Antarctic subglacial community and high-resolution replicate sampling inform hydrological connectivity in a polar desert

Antarctic subglacial environments host microbial ecosystems and are proving to be geochemically and biologically diverse. The Taylor Glacier, Antarctica, periodically expels iron-rich brine through a conduit sourced from a deep subglacial aquifer, creating a dramatic red surface feature known as Blood Falls. We used Illumina MiSeq sequencing to describe the core microbiome of this subglacial brine and identified previously undetected but abundant groups including the candidate bacterial phylum Atribacteria and archaeal phylum Pacearchaeota. Our work represents the first microbial characterization of samples collected from within a glacier using a melt probe, and the only Antarctic subglacial aquatic environment that, to date, has been sampled twice. A comparative analysis showed the brine community to be stable at the operational taxonomic unit level of 99% identity over a decade. Higher resolution sequencing enabled deconvolution of the microbiome of subglacial brine from mixtures of materials collected at the glacier surface. Diversity patterns between this brine and samples from the surrounding landscape provide insight into the hydrological connectivity of subglacial fluids to the surface polar desert environment. Understanding subice brines collected on the surfaces of thick ice covers has implications for analyses of expelled materials that may be sampled on icy extraterrestrial worlds.     

Microbial community structure and ecology of subglacial sediments in two polythermal Svalbard glaciers characterized by epifluorescence microscopy and PLFA

Biological and physico-chemical characteristics of subglacial sediments were studied in Svalbard. Sediment from close proglacial and supraglacial environments was used for a comparison. Viable bacteria, cyanobacteria and microalgae were detected in subglacial sediments from two polythermal glaciers using epifluorescence microscopy and phospholipid fatty acid (PLFA) analyses. The subglacial samples were generally of higher pH values, coarser texture and lower water content, organic matter, organic carbon, and nitrogen compared to proglacial and supraglacial sediments). Bacterial counts of 1.6 × 10⁷ cells mg^{− 1} OM (organic matter) were found. Cyanobacteria and algae were also of low abundance [4.2 cells mg^{− 1} DW (dry weight)]. Cyanobacteria comprised the major proportion of the autophotothrophic assemblages of subglacial soils. Deglaciated soils were similar to subglacial sediment in physico-chemical properties and microbial structure and numbers, unlike soil from vegetated sites or cryoconite sediment. In subglacial and deglaciated soil, relatively low diversity of microorganisms and low substrate availability was detected by PLFA analyses. Good accordance in microbial community structure assessments between epifluorescence microscopy and PLFA analyses was found. Our results suggest that the subglacial microbial populations can be divided into two groups: autochthonous microorganims (chemoheterotrophic bacteria) and allochthonous that retain the ability to proliferate and give rise to active population when conditions become favorable. Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Microbial Life beneath a High Arctic Glacier

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 ¹⁴CO₂ 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 CO₂ and CH₄ 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.     

Relative Incidence of Ascomycetous Yeasts in Arctic Coastal Environments

Previous studies of fungi in polar environments have revealed a prevalence of basidiomycetous yeasts in soil and in subglacial environments of polythermal glaciers. Ascomycetous yeasts have rarely been reported from extremely cold natural environments, even though they are known contaminants of frozen foods. Using media with low water activity, we have isolated various yeast species from the subglacial ice of four glaciers from the coastal Arctic environment of Kongsfjorden, Spitzbergen, including Debaryomyces hansenii and Pichia guillermondii, with counts reaching 10⁴ CFU L⁻¹. Together with the basidiomycetes Cryptococcus liquefaciens and Rhodotorula mucilaginosa, these yeasts represent the stable core of the subglacial yeast communities. Other glacial ascomycetous species isolated included Candida parapsilosis and a putative new species that resembles Candida pseudorugosa. The archiascomycete Protomyces inouyei has seldom been detected anywhere in the world but was here recovered from ice in a glacier cave. The glacier meltwater contained only D. hansenii, whereas the seawater contained D. hansenii, Debaryomyces maramus, Pichia guilliermondii, what appears to represent a novel species resembling Candida galli and Metschnikowia bicuspidata. Only P. guilliermondii was isolated from sea ice, while snow/ice in the fjord tidal zone included C. parapsilosis, D. hansenii, P. guilliermondii and Metschnikowia zobellii. All of these isolated strains were characterized as psychrotolerant and xero/halotolerant, with the exception of P. inouyei.     

Chemical weathering and solute export by meltwater in a maritime Antarctic glacier basin

Solute yields, laboratory dissolution data and both chemical and isotopic markers of rock weathering reactions are used to characterise the biogeochemistry of glacial meltwaters draining a maritime Antarctic glacier. We find that delayed flowpaths through ice-marginal talus and moraine sediments are critical for the acquisition of solute from rock minerals because delayed flowpaths through subglacial sediments are absent beneath this small, cold-based glacier. Here the mechanisms of weathering are similar to those reported in subglacial environments, and include sub-oxic conditions in the early summer and increasingly oxic conditions thereafter. Up to 85% of the NO₃ ^? and 65% of the SO₄ ^2? are most likely produced by bacterially mediated reactions in these ice marginal sediments. However, reactive pyrite phases are sparse in the host rocks, limiting the export of Fe, SO₄ ^2? and cations that may be removed by weathering once pyrite oxidation has taken place. This means that dissolution of Ca²⁺ and Na⁺ from carbonate and silicate minerals dominate, producing moderate cationic denudation yields from Tuva Glacier (163 Σ*meq⁺ m^?2 a^?1) compared to a global range of values (94–4,200 Σ*meq⁺ km^?2 a^?1). Overall, crustally derived cations represent 42% of the total cationic flux, the rest being accounted for by snowpack sources.     

Comparison of microbial community compositions of two subglacial environments reveals a possible role for microbes in chemical weathering processes

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.     

Indicators for nitrogen status and leaching in subtropical forest ecosystems,South China

The deposition of nitrogen (N) is high in subtropical forest in South China and it is expected to increase further in the coming decades. To assess effects of increasing deposition on N cycling, we investigated the current N status of two selected 40–45-year-old masson pine-dominated Chinese subtropical forest stands at Tieshanping (TSP, near Chongqing City) and Caijiatang (CJT in Shaoshan, Hunan province), and explored the applicability of several indicators for N status and leaching, suggested for temperate and boreal forest ecosystems. Current atmospheric N deposition to the systems is from 25 to 49 kg ha⁻¹ year⁻¹. The concentration of total N in the upper 15 cm of the soil is from as low as 0.05% in the B₂ horizon to as high as 0.53% in the O/A horizon. The concentration of organic carbon (C) varies from 0.74 (B₂) to 9.54% (O/A). Pools of N in the upper 15 cm of the soils range from 1460 to 2290 kg N ha⁻¹, where 25–55% of the N pool is in the O/A horizon (upper 3 cm of the soil). Due to a lack of a well-developed continuous O horizon (forest floor), the C/N ratio of this layer cannot be used as an indicator for the N status, as is commonly done in temperate and boreal forests. The net N mineralization rate (mg N g⁻¹ C year⁻¹) in individual horizons correlates significantly with the C/N ratio, which is from as high as 18.2 in the O/A horizon to as low as 11.2 in the B₂ horizon. The N₂O emission flux from soil is significantly correlated with the KCl extractable NH₄⁺–N in the O/A horizon and with the net nitrification in the upper 15 cm of the soil. However, the spatial and temporal variation of the N₂O emission rate is high and rates are small and often difficult to detect in the field. The soil flux density of mineral N, defined as the sum of the throughfall N input rate and the rate of in situ net N mineralization in the upper 15 cm of the soil, i.e., the combination of deposition input and the N status of the system, explains the NO₃⁻ leaching potential at 30 cm soil depth best. The seasonality of stream water N concentration at TSP and CJT is climatic and hydrologically controlled, with highest values commonly occurring in the wet growing season and lowest in the dry dormant season. This is different from temperate forest ecosystems, where N saturation is indicated by elevated NO₃⁻ leaching in stream water during summer.     

Novel opportunity for understanding origin and evolution of life: perspectives on the exploration of subglacial environment of Lake Vostok, Antarctica

The year 2012 began with reports on the completion of drilling over the largest subglacial lake of Antarctica, Lake Vostok. It has been sealed from free exchange with the atmosphere for ～420,000 years, making it a potentially rich and largely unexplored storehouse of genetic information for solving evolutionary queries. Indirect evidence suggests that many impediments exist in the subglacial environment, such as high oxygen tension, low inorganic and organic nutrient concentrations, etc., whereas analysis of glacial and accretion ice from Vostok also implies that the elemental requirements for microbial growth could be satisfied in the lake by many possible electron donors (like H₂, Fe²⁺ and NO ₂ ^? ). Recent reports on the recovery of the longest ever ice core of ～3,769 m open up a new window for answering many questions about microbiological life in extremes, evolution, and adaptations. This review attempts to present an updated understanding on the potential significance of subglacial environments in unraveling the mystery of evolution of life.     

Redox Fluctuations Frame Microbial Community Impacts on N-cycling Rates in a Humid Tropical Forest Soil

Fluctuating soil redox regimes may facilitate the co-occurrence of microbial nitrogen transformations with significantly different sensitivities to soil oxygen availability. In an upland humid tropical forest, we explored the impact of fluctuating redox regimes on gross nitrogen cycling rates and microbial community composition. Our results suggest that the rapidly fluctuating redox conditions that characterize these upland soils allow anoxic and oxic N processing to co-occur. Gross nitrogen mineralization was insensitive to soil redox fluctuations. In contrast, nitrifiers in this soil were directly affected by low redox periods, yet retained some activity even after 3–6 weeks of anoxia. Dissimilatory nitrate reduction to ammonium (DNRA) was less sensitive to oxygen exposure than expected, indicating that the organisms mediating this reductive process were also tolerant of unfavorable (oxic) conditions. Denitrification was a stronger sink for NO₃⁻ in consistently anoxic soils than in variable redox soils. Microbial biomass and community composition were maintained with redox fluctuation, but biomass decreased and composition changed under static oxic and anoxic soil regimes. Bacterial community structure was significantly correlated with rates of nitrification, denitrification and DNRA, suggesting that redox-control of soil microbial community structure was an important determinant of soil N-cycling rates. Specific nitrogen cycling functional groups in this environment (such as nitrifiers, DNRA organisms, and denitrifiers) appear to have adapted to nutrient resources that are spatially and temporally variable. In soils where oxygen is frequently depleted and re-supplied, characteristics of microbial tolerance and resilience can frame N cycling patterns.     

The prokaryotic community of subglacial bottom sediments of Antarctic Lake Untersee: Detection by cultural and direct microscopic techniques

The heterotrophic mesophilic microbial component was studied in microbial communities of the samples of frozen regolith collected from the glacier near Lake Untersee collected in 2011 during the joint Russian-American expedition to central Dronning Maud Land (Eastern Antarctica). Cultural techniques revealed high bacterial numbers in the samples. For enumeration of viable cells, the most probable numbers (MPN) method proved more efficient than plating on agar media. Fluorescent in situ hybridization with the relevant oligonucleotide probes revealed members of the groups Eubacteria (Actinobacteria, Firmicutes) and Archaea. The application of the methods of cell resuscitation, such as the use of diluted media and prevention of oxidative stress, did not result in a significant increase in the numbers of viable cells retrieved from subglacial sediment samples. Our previous investigations demonstrated the necessity for special procedures for efficient reactivation of the cells from microbial communities of replace with buried soil and permafrost samples collected in the Arctic zone. The differing responses to the special resuscitation procedures may reflect the differences in the physiological and morphological state of bacterial cells in microbial communities subject to continuous or periodic low temperatures and dehydration.