Focus on Cryospheric Ecosystems

The fourteen letters that contributed to this focus issue on cryospheric ecosytems provide an excellent basis for considering the state of the science following a marked increase in research attention since the new millennium. Research letters from the focus issue provide significant insights into the biogeochemical and biological processes associated with snow, glacier ice and glacial sediments. This has been achieved via a significant, empirical effort that has given particular emphasis to glacier surface habitats. However, far less is known about aerobiology, glacial snow covers, supraglacial lakes and sub-ice sedimentary habitats, whose access for sampling and in-situ monitoring remains a great challenge to scientists. Furthermore, the use of models to explore key fluxes, processes and impacts of a changing glacial cryosphere are conspicuous by their absence. As a result, a range of process investigations and modelling studies are required to address the increasing urgency and uncertainty that is associated with understanding the response of cryospheric ecosystems to global change.

Recently deglaciated areas are ideal environments to study soil formation and primary microbial succession where phototrophic microorganisms may play a role as primary producers. The aim of our study was to investigate the cyanobacterial and green algal community composition in three different successional stages of the Damma glacier forefield in the Swiss Alps using 16S rDNA and ITS rDNA clone libraries. Cyanobacterial target sequences varied along the glacier forefield, with the highest cyanobacterial 16S rRNA gene copies found in sparsely vegetated soils. Sequence analysis revealed that the phototrophic communities were distinct in each of the three soil environments. The majority of the cyanobacterial sequences retrieved from barren soils were related to the Oscillatoriales. The diversity in sparsely vegetated soils was low, and sequences closely related to Nostoc sp. dominated. The majority of the algal phylotypes are related to members of the Trebouxiophyceae known to live as symbiotic partners in lichens. We conclude that the community composition appears to shift markedly along the chronosequence, indicating that each soil environment selects for its phototrophic community. When cyanobacteria occur together with eukaryotic microalgae, they form a rich source of organic matter and may be important contributors of carbon in nutrient-deficient deglaciated soils.

Snow and ice algae are cold tolerant algae growing on the surface of snow and ice, and they play an important role in the carbon cycles for glaciers and snowfields in the world. Seasonal and altitudinal variations in seven major taxa of algae (green algae and cyanobacteria) were investigated on the Gulkana glacier in Alaska at six different elevations from May to September in 2001. The snow algal communities and their biomasses changed over time and elevation. Snow algae were rarely observed on the glacier in May although air temperature had been above 0 ° C since the middle of the month and surface snow had melted. In June, algae appeared in the lower areas of the glacier, where the ablation ice surface was exposed. In August, the distribution of algae was extended to the upper parts of the glacier as the snow line was elevated. In September, the glacier surface was finally covered with new winter snow, which terminated algal growth in the season. Mean algal biomass of the study sites continuously increased and reached 6.3 × 10 μl m⁻² in cell volume or 13 mg carbon m⁻² in September. The algal community was dominated by Chlamydomonas nivalis on the snow surface, and by Ancylonema nordenskiöldii and Mesotaenium berggrenii on the ice surface throughout the melting season. Other algae were less abundant and appeared in only a limited area of the glacier. Results in this study suggest that algae on both snow and ice surfaces significantly contribute to the net production of organic carbon on the glacier and substantially affect surface albedo of the snow and ice during the melting season.

Cryoconite is a microbe–mineral aggregate which darkens the ice surface of glaciers. Microbial process and marker gene PCR-dependent measurements reveal active and diverse cryoconite microbial communities on polar glaciers. Here, we provide the first report of a cryoconite metagenome and culture-independent study of alpine cryoconite microbial diversity. We assembled 1.2 Gbp of metagenomic DNA sequenced using an Illumina HiScanSQ from cryoconite holes across the ablation zone of Rotmoosferner in the Austrian Alps. The metagenome revealed a bacterially-dominated community, with Proteobacteria (62% of bacterial-assigned contigs) and Bacteroidetes (14%) considerably more abundant than Cyanobacteria (2.5%). Streptophyte DNA dominated the eukaryotic metagenome. Functional genes linked to N, Fe, S and P cycling illustrated an acquisitive trend and a nitrogen cycle based upon efficient ammonia recycling. A comparison of 32 metagenome datasets revealed a similarity in functional profiles between the cryoconite and metagenomes characterized from other cold microbe–mineral aggregates. Overall, the metagenomic snapshot reveals the cryoconite ecosystem of this alpine glacier as dependent on scavenging carbon and nutrients from allochthonous sources, in particular mosses transported by wind from ice-marginal habitats, consistent with net heterotrophy indicated by productivity measurements. A transition from singular snapshots of cryoconite metagenomes to comparative analyses is advocated.

Arctic snowpacks are often considered as chemical reactors for a variety of chemicals deposited through wet and dry events, but are overlooked as potential sites for microbial metabolism of reactive nitrogen species. The fate of deposited species is critical since warming leads to the transfer of contaminants to snowmelt-fed ecosystems. Here, we examined the role of microorganisms and the potential pathways involved in nitrogen cycling in the snow. Next generation sequencing data were used to follow functional gene abundances and a 16S rRNA (ribosomal ribonucleic acid) gene microarray was used to follow shifts in microbial community structure during a two-month spring-time field study at a high Arctic site, Svalbard, Norway (79° N). We showed that despite the low temperatures and limited water supply, microbial communities inhabiting the snow cover demonstrated dynamic shifts in their functional potential to follow several different pathways of the nitrogen cycle. In addition, microbial specific phylogenetic probes tracked different nitrogen species over time. For example, probes for Roseomonas tracked nitrate concentrations closely and probes for Caulobacter tracked ammonium concentrations after a delay of one week. Nitrogen cycling was also shown to be a dominant process at the base of the snowpack.

During the austral summers of 2004 and 2009, we sampled a supraglacial stream on the Cotton Glacier, Antarctica. The stream dissolved organic matter (DOM) was low (44–48 μM C) and lacked detectable humic fluorescence signatures. Analysis of the excitation emissions matrices (EEMs) indicated that amino-acid fluorophores dominated, consistent with DOM of microbial origin, with little humic-like fluorescence. In most aquatic ecosystems, humic DOM attenuates harmful UV radiation and its absence may represent an additional stressor influencing the microbial community. Nonetheless, the stream contained an active microbial assemblage with bacterial cell abundances from 2.94 × 10⁴ to 4.97 × 10⁵ cells ml⁻¹, and bacterial production ranging from 58.8 to 293.2 ng C l⁻¹ d⁻¹. Chlorophyll-a concentrations ranged from 0.3 to 0.53 μg l⁻¹ indicating that algal phototrophs were the probable source of the DOM. Microbial isolates produced a rainbow of pigment colors, suggesting adaptation to stress, and were similar to those from other cryogenic systems (Proteobacteria and Bacteroidetes lineages). Supraglacial streams provide an example of contemporary microbial processes on the glacier surface and a natural laboratory for studying microbial adaptation to the absence of humics.

We compared planktonic primary and secondary production across twenty meltwater ponds on the surface of the McMurdo Ice Shelf in January 2007, including some ponds with basal brines created by meromictic stratification. Primary production ranged from 1.07 to 65.72 mgC m⁻³ h⁻¹ in surface waters. In stratified ponds primary production was always more than ten times higher in basal brines than in the corresponding mixolimnion. Regression tree analysis (r² = 0.80) identified inorganic nitrogen (as ${\mathrm{NH}}_{4}^{+}$) as the main factor limiting planktonic primary production. However, there was also evidence of inorganic carbon co-limitation of photosynthesis in some of the more oligotrophic waters. Neither C nor N limited carbon fixation at [NH₄–N] > 50 mg m⁻³, with photoinhibition the factor most likely limiting photosynthesis under such conditions. Primary production was the only factor significantly correlated to bacterial production and the relationship (r² = 0.56) was non-linear. Nitrogen limitation and tight coupling of planktonic primary and bacterial production is surprising in these ponds, as all have large pools of dissolved organic carbon (1.2–260 g m⁻³) and organic nitrogen (all >130 mg m⁻³). The dissolved pools of organic carbon and nitrogen appear to be recalcitrant and bacterial production to be constrained by limited release of labile organics from phytoplankton.

The aggregation of surface debris particles on melting glaciers into larger units (cryoconite) provides microenvironments for various microorganisms and metabolic processes. Here we investigate the microbial community on the surface of Aldegondabreen, a valley glacier in Svalbard which is supplied with carbon and nutrients from different sources across its surface, including colonies of seabirds. We used a combination of geochemical analysis (of surface debris, ice and meltwater), quantitative polymerase chain reactions (targeting the 16S ribosomal ribonucleic acid and amoA genes), pyrosequencing and multivariate statistical analysis to suggest possible factors driving the ecology of prokaryotic microbes on the surface of Aldegondabreen and their potential role in nitrogen cycling. The combination of high nutrient input with subsidy from the bird colonies, supraglacial meltwater flow and the presence of fine, clay-like particles supports the formation of centimetre-scale cryoconite aggregates in some areas of the glacier surface. We show that a diverse microbial community is present, dominated by the cyanobacteria, Proteobacteria, Bacteroidetes, and Actinobacteria, that are well-known in supraglacial environments. Importantly, ammonia-oxidizing archaea were detected in the aggregates for the first time on an Arctic glacier.

The Arctic net ecosystem exchange (NEE) of CO₂ between the land surface and the atmosphere is influenced by the timing of snow onset and melt. The objective of this study was to examine whether uncertainty in model estimates of NEE could be reduced by representing the influence of snow on NEE using remote sensing observations of snow cover area (SCA). Observations of NEE and time-lapse images of SCA were collected over four locations at a low Arctic site (Daring Lake, NWT) in May–June 2010. Analysis of these observations indicated that SCA influences NEE, and that good agreement exists between SCA derived from time-lapse images, Landsat and MODIS. MODIS SCA was therefore incorporated into the vegetation photosynthesis respiration model (VPRM). VPRM was calibrated using observations collected in 2005 at Daring Lake. Estimates of NEE were then generated over Daring Lake and Ivotuk, Alaska (2004–2007) using VPRM formulations with and without explicit representations of the influence of SCA on respiration and/or photosynthesis. Model performance was assessed by comparing VPRM output against unfilled eddy covariance observations from Daring Lake and Ivotuk (2004–2007). The uncertainty in VPRM estimates of NEE was reduced when respiration was estimated as a function of air temperature when SCA ≤ 50% and as a function of soil temperature when SCA > 50%.

Cryoconite holes are ice-bound habitats that can act as refuges for aquatic and terrestrial microorganisms on glacier surfaces. In the McMurdo Dry Valleys of Antarctica, these holes are often capped by an ice lid that prevents the exchange of material and gases with the surrounding atmosphere and aquatic environment. Diatoms have been documented in cryoconite holes, and recent findings suggest that these habitats may harbour a distinctive diatom flora compared to the surrounding aquatic environments. In this study, we examined diatom community composition in cryoconite holes and environmental correlates across three glaciers in Taylor Valley, Antarctica. The diatom communities were dominated by two genera, Muelleria and Diadesmis, both of which had high viability and could have been seeded from the surrounding ephemeral streams. The location of the cryoconite hole within the valley was a key determinant of community composition. A diatom species richness gradient was observed that corresponded to distance inland from the coast and co-varied with species richness in streams within the same lake basin. Cryoconite holes that were adjacent to streams with higher diversity displayed greater species richness. However, physical factors, such as the ability to withstand freeze–thaw conditions and to colonize coarse sediments, acted as additional selective filters and influenced diatom diversity, viability and community composition.

Dissolved humic material (HDOM) is ubiquitous to all natural waters and its source material influences its chemical structure, reactivity, and bioavailability. While terrestrially derived HDOM reference materials distributed by the International Humic Substances Society (IHSS) have been readily available to engineering and scientific communities, a microbially derived reference HDOM was not, despite the well-characterized differences in the chemistry and reactivity of HDOM derived from terrestrial versus microbial sources. To address this gap, we collected a microbial reference fulvic acid from Pony Lake (PLFA) for distribution through the IHSS. Pony Lake is a saline coastal pond on Ross Island, Antarctica, where the landscape is devoid of terrestrial plants. Sample collection occurred over a 17-day period in the summer season at Pony Lake. During this time, the dissolved organic carbon (DOC) concentrations increased nearly two-fold, and the fulvic acid fraction (collected using the XAD-8 method) accounted for 14.6% of the DOC. During the re-concentration and desalting procedures we isolated two other chemically distinct fulvic acid fractions: (1) PLFA-2, which was high in carbohydrates and (2) PLFA-CER, which was high in nitrogen. The chemical characteristics (elemental analysis, optical characterization with UV–vis and fluorescence spectroscopy, and ¹³C NMR spectroscopy) of the three fulvic acid fractions helped to explain their behavior during isolation.

The surfaces of glaciers are extreme ecosystems dominated by microbial communities. Viruses are found in abundance here, with a high frequency of bacteria displaying visible virus infection. In this study, viral and bacterial production was measured in Arctic cryoconite holes to address the control that viruses play in these highly truncated ecosystems. Mean bacterial carbon production in the sediments of cryoconite holes was found to be 57.8 ± 12.9 ng C g⁻¹ dry wt. h⁻¹, which predicted a mean of 1.89–5.41 × 10⁶ cells g⁻¹ dry wt. h⁻¹ based on a range of conversion factors. Relative to this, virus production was found to be high, up to 8.98 × 10⁷ virus like particles g⁻¹ dry wt. h⁻¹ were produced, which is comparable to virus production in sediments around the globe. The virus burst size was assessed by transmission electron microscopy and found to be amongst the lowest recorded in the literature (mean 2.4). Hence, to account for the measured virus production, the viral induced bacterial mortality was calculated to be more than capable of accounting for the mortality of all bacterial production. The data presented here, therefore, suggests that viral induced mortality is a dominant process for the release and recycling of carbon and nutrients in supraglacial ecosystems.

Bare soils in the area of retreating glaciers are ideal environments to study the role of microorganisms in the early soil formation and in processes of mineral weathering. The aim of our study was to investigate whether the source of carbohydrate would influence the patterns of organic acids exuded by fungal species. Three pioneering fungus species, isolated from fine granitic sediments in front of the Damma glacier from the central Swiss Alps, have previously been found to have the capability to exude organic acids and dissolve granite powder. In batch experiments, various carbohydrates, including glucose, cellulose, pectin, pollen, and cell remnants of cyanobacteria, fungi, and algae, were applied as carbohydrate sources and the patterns of exuded organic acids recorded. The results showed that two fungi, the zygomycete fungus Mucor hiemalis and the ascomycete fungus Penicillium chrysogenum, released a significantly higher amount of organic acids in dependence on specific carbohydrate sources. Pollen and algae as carbohydrate sources triggered significantly the exudation of malate in M. hiemalis, and pollen and cellulose that of oxalate in P. chrysogenum. We conclude that the occurrence of complex carbohydrate sources in nutrient-deficient deglaciated soils may positively influence the exudation of organic acids of fungi. In particular, pollen and remnants of other microorganisms can trigger the exudation of organic acids of fungi in order to promote the weathering of minerals and to make nutrients available that would otherwise be trapped in that cryospheric environment.

Cryoconite is a dark-coloured surface dust deposited on glaciers that consists of wind-blown mineral particles, as well as organic matter derived from microbes living on glaciers. In this paper, we analyse the Sr and Nd isotopic ratios of four mineral fractions (i.e., the saline, carbonate, phosphate, and silicate mineral fractions), as well as the organic fraction, of cryoconite samples obtained from six Asian glaciers (the Altai, Pamir, Tien Shan, Qillian Shan, and Himalayan regions), and discuss their geographical variations in terms of the geological origins of the mineral particles and the biogeochemical processes on the glaciers. The silicate mineral fraction showed lower Sr and higher Nd ratios for the glaciers located to the north (Altai, ⁸⁷Sr/⁸⁶Sr: 0.713 490–0.715 284, εNd(0): −6.4 to −5.6), while higher Sr and lower Nd ratios for the glaciers located to the south (Himalayas, ⁸⁷Sr/⁸⁶Sr: 0.740 121–0.742 088, εNd(0): −16.4 to −15.7); the ratios were similar to those of desert sand, loess, and river sediments in the respective regions of the glaciers. This result suggests that the silicate minerals within the cryoconites were derived from different sources depending on the geographical locations of the glaciers. The isotopic ratios of the saline, carbonate, and phosphate mineral fractions were distinct from those of the silicate fraction, and were similar to those of evaporites and apatite deposits from the Asian deserts, but also varied geographically, indicating that they are likely to reflect their geological origin. The Sr isotopic ratios of the organic fraction were similar to those of the saline and carbonate fractions from glaciers in the central area (Tien Shan and Qillian Shan), but were higher than those of the saline and carbonate fractions, and lower than the phosphate mineral fraction, in the northern and southern areas. The ratios of organic fraction may be determined from the mixing ratio of calcium sources incorporated by microbes on the glaciers.

Glacier ecosystems are a significant source of bioavailable, yet ancient dissolved organic carbon (DOC). Characterizing DOC in Mendenhall Glacier outflow (southeast Alaska) we document a seasonal persistence to the radiocarbon-depleted signature of DOC, highlighting ancient DOC as a ubiquitous feature of glacier outflow. We observed no systematic depletion in Δ¹⁴C-DOC with increasing discharge during the melt season that would suggest mobilization of an aged subglacial carbon store. However, DOC concentration, δ¹³C-DOC, Δ¹⁴C-DOC and fluorescence signatures appear to have been influenced by runoff from vegetated hillslopes above the glacier during onset and senescence of melt. In the peak glacier melt period, the Δ¹⁴C-DOC of stream samples at the outflow (−181.7 to −355.3‰) was comparable to the Δ¹⁴C-DOC for snow samples from the accumulation zone (−207.2 to −390.9‰), suggesting that ancient DOC from the glacier surface is exported in glacier runoff. The pre-aged DOC in glacier snow and runoff is consistent with contributions from fossil fuel combustion sources similar to those documented previously in ice cores and thus provides evidence for anthropogenic perturbation of the carbon cycle. Overall, our results emphasize the need to further characterize DOC inputs to glacier ecosystems, particularly in light of predicted changes in glacier mass and runoff in the coming century.