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Abstract Background Dark pigmented snow and glacier ice algae on glaciers and ice sheets contribute to accelerating melt. The biological controls on these algae, particularly the role of viruses, remain poorly understood. Giant viruses, classified under the nucleocytoplasmic large DNA viruses (NCLDV) supergroup (phylum Nucleocytoviricota), are diverse and globally distributed. NCLDVs are known to infect eukaryotic cells in marine and freshwater environments, providing a biological control on the algal population in these ecosystems. However, there is very limited information on the diversity and ecosystem function of NCLDVs in terrestrial icy habitats. Results In this study, we investigate for the first time giant viruses and their host connections on ice and snow habitats, such as cryoconite, dark ice, ice core, red and green snow, and genomic assemblies of five cultivated Chlorophyta snow algae. Giant virus marker genes were present in almost all samples; the highest abundances were recovered from red snow and the snow algae genomic assemblies, followed by green snow and dark ice. The variety of active algae and protists in these GrIS habitats containing NCLDV marker genes suggests that infection can occur on a range of eukaryotic hosts. Metagenomic data from red and green snow contained evidence of giant virus metagenome-assembled genomes from the orders Imitervirales, Asfuvirales, and Algavirales. Conclusion Our study highlights NCLDV family signatures in snow and ice samples from the Greenland ice sheet. Giant virus metagenome-assembled genomes (GVMAGs) were found in red snow samples, and related NCLDV marker genes were identified for the first time in snow algal culture genomic assemblies; implying a relationship between the NCLDVs and snow algae. Metatranscriptomic viral genes also aligned with metagenomic sequences, suggesting that NCLDVs are an active component of the microbial community and are potential “top-down” controls of the eukaryotic algal and protistan members. This study reveals the unprecedented presence of a diverse community of NCLDVs in a variety of glacial habitats dominated by algae.

The glacier ice algae Ancylonema alaskanum and Ancylonema nordenskiöldii grow in harsh dynamic environments on bare ice surfaces. In these environments, they contribute to the continuous darkening of the ice surface, which in turn accelerates the ice melt. However, investigation into their adaptation and resilience in these environments is necessary in order to understand their robustness and potential for increasing the intensity of blooms. In this study, it was examined how variations in environmental parameters such as pH, salinity, light and temperature impacted the photophysiology of the glacier ice algae during a bloom in Greenland. Through in situ incubations and pulse-amplitude-modulation (PAM) fluorometric measurements, the photophysiological responses of algal cells were assessed. Results suggest that light intensity significantly influences glacier ice algae photophysiology, with cells exhibiting better performance (in terms of a higher theoretical maximum light coefficient and maximum quantum efficiency) under lower light intensity. Moreover, while light emerges as the primary driver of photophysiology, glacier ice algae demonstrate tolerance to a broad range of pH and temperatures four times higher than those experienced during Greenland’s summer.

The Greenland Ice Sheet is a biome which is mainly microbially driven. Several different niches can be found within the glacial biome for those microbes able to withstand the harsh conditions, e.g., low temperatures, low nutrient conditions, high UV radiation in summer, and contrasting long and dark winters. Eukaryotic algae can form blooms during the summer on the ice surface, interacting with communities of bacteria, fungi, and viruses. Cryoconite holes and snow are also habitats with their own microbial community. Nevertheless, the microbiome of supraglacial habitats remains poorly studied, leading to a lack of representative genomes from these environments. Under-investigated extremophiles, like those living on the Greenland Ice Sheet, may provide an untapped reservoir of chemical diversity that is yet to be discovered. In this study, an inventory of the biosynthetic potential of these organisms is made, through cataloging the presence of biosynthetic gene clusters in their genomes. There were 133 high-quality metagenome-assembled genomes (MAGs) and 28 whole genomes of bacteria obtained from samples of the ice sheet surface, cryoconite, biofilm, and snow using culturing-dependent and -independent approaches. AntiSMASH and BiG-SCAPE were used to mine these genomes and subsequently analyze the resulting predicted gene clusters. Extensive sets of predicted Biosynthetic Gene Clusters (BGCs) were collected from the genome collection, with limited overlap between isolates and MAGs. Additionally, little overlap was found in the biosynthetic potential among different environments, suggesting specialization of organisms in specific habitats. The median number of BGCs per genome was significantly higher for the isolates compared to the MAGs. The most talented producers were found among Proteobacteria. We found evidence for the capacity of these microbes to produce antimicrobials, carotenoid pigments, siderophores, and osmoprotectants, indicating potential survival mechanisms to cope with extreme conditions. The majority of identified BGCs, including those in the most prevalent gene cluster families, have unknown functions, presenting a substantial potential for bioprospecting. This study underscores the diverse biosynthetic potential in Greenland Ice Sheet genomes, revealing insights into survival strategies and highlighting the need for further exploration and characterization of these untapped resources.

Pigmented microalgae inhabiting snow and ice environments lower the albedo of glacier and ice-sheet surfaces, significantly enhancing surface melt. Our ability to accurately predict their role in glacier and ice-sheet surface mass balance is limited by the current lack of empirical data to constrain their representation in predictive models. Here we present new empirical optical properties for snow and ice algae and incorporate them in a radiative transfer model to investigate their impact on snow and ice surface albedo. We found ice algal cells to be more efficient absorbers than snow algal cells, but their blooms had comparable impact on surface albedo due to the different photic conditions of their habitats. We then used the model to reconstruct the effect of ice algae on bare ice albedo spectra collected at our field site in southern Greenland, where blooms dropped the albedo locally by between 3 and 43%, equivalent to 1–10 L m$^{-2}$ d$^{-1}$ of melted ice. Using the newly parametrized model, future studies could investigate biological albedo reduction and algal quantification from remote hyperspectral and multispectral imagery.

Phytobenthos, major primary producers in freshwater, may be periodically exposed to herbicides through runoff, spray drift, and leaching, but experimental work on their response to herbicides is limited. Outdoor flumes were used to measure the recovery of intact phytobenthic biofilms, following a short-term pulse to a glyphosate-based herbicide (GBH) and chlorotoluron (CLT), singly or as a mixture (GBHC). Two days following the removal of the herbicides, significant declines in biomass and rates of areal gross maximum photosynthesis were recorded in GBH and GBHC-treated flumes. Herbicide pulses negatively impacted the biomass of phytobenthos, measured as chlorophyll a, which in turn may have alleviated resource limitation, evidenced by significant increases in biomass-specific rates of gross maximum photosynthesis. After 4.5 days, biomass concentrations were similar in controls and treatments, indicating biofilm recovery in the GBH and GBHC biofilms, though the proportion of green algae relative to diatoms was significantly higher in GBH and GBHC-impacted biofilms. Significant increases were recorded in the Trophic Diatom Index (TDI), indicating benthic diatom development along different trajectories, following the herbicide pulse. Collectively, these results indicate greater insights into herbicide impacts on phytobenthos may be gained by working with the whole phytobenthic community.

Abstract Blooms of pigmented algae darken the surface of glaciers and ice sheets, thereby enhancing solar energy absorption and amplifying ice and snow melt. The impacts of algal pigment and community composition on surface darkening are still poorly understood. Here, we characterise glacier ice and snow algal pigment signatures on snow and bare ice surfaces and study their role in photophysiology and energy absorption on three glaciers in Southeast Greenland. Purpurogallin and astaxanthin esters dominated the glacier ice and snow algal pigment pools (mass ratios to chlorophyll a of 32 and 56, respectively). Algal biomass and pigments impacted chromophoric dissolved organic matter concentrations. Despite the effective absorption of astaxanthin esters at wavelengths where incoming irradiance peaks, the cellular energy absorption of snow algae was 95% lower than anticipated from their pigmentation, due to pigment packaging. The energy absorption of glacier ice algae was consequently ~ 5 × higher. On bare ice, snow algae may have locally contributed up to 13% to total biological radiative forcing, despite contributing 44% to total biomass. Our results give new insights into the impact of algal community composition on bare ice energy absorption and biomass accumulation during snow melt.

The surface of the Greenland Ice Sheet is darkening, which accelerates its surface melt. The role of glacier ice algae in reducing surface albedo is widely recognised but not well quantified and the feedbacks between the algae and the weathering crust remain poorly understood. In this letter, we summarise recent advances in the study of the biological darkening of the Greenland Ice Sheet and highlight three key research priorities that are required to better understand and forecast algal-driven melt: (i) identifying the controls on glacier ice algal growth and mortality, (ii) quantifying the spatio-temporal variability in glacier ice algal biomass and processes involved in cell redistribution and (iii) determining the albedo feedbacks between algal biomass and weathering crust characteristics. Addressing these key research priorities will allow us to better understand the supraglacial ice-algal system and to develop an integrated model incorporating the algal and physical controls on ice surface albedo.

Distant glacial areas are interconnected by a complex system of fractures and water channels which run in the glacier interior and characterize the englacial realm. Water can slowly freeze in these channels where the slow freezing excludes air bubbles giving the ice a clear aspect. This ice is uplifted to the surface ablation zone by glacial movements and can therefore be observed in the form of clear surface ice bands. We employed an indirect method to sample englacial water by coring these ice bands. We were able, for the first time, to compare microbial communities sampled from clear (i.e. frozen englacial water bands) and cloudy ice (i.e. meteoric ice) through 16S rRNA gene sequencing. Although microbial communities were primarily shaped and structured by their spatial distribution on the glacier, ice type was a clear secondary factor. One area of the glacier, in particular, presented significant microbial community clear/cloudy ice differences. Although the clear ice and supraglacial communities showed typical cold-adapted glacial communities, the cloudy ice had a less defined glacial community and ubiquitous environmental organisms. These results highlight the role of englacial channels in the microbial dispersion within the glacier and, possibly, in the shaping of glacial microbial communities.

Volatile organic compounds (VOCs) are emitted by organisms for a range of physiological and ecological reasons. They play an important role in biosphere–atmosphere interactions and contribute to the formation of atmospheric secondary aerosols. The Greenland ice sheet is home to a variety of microbial communities, including highly abundant glacier ice algae, yet nothing is known about the VOCs emitted by glacial communities. For the first time, we present VOC emissions from supraglacial habitats colonized by active microbial communities on the southern Greenland ice sheet during July 2020. Emissions of C5–C30 compounds from bare ice, cryoconite holes, and red snow were collected using a push–pull chamber active sampling system. A total of 92 compounds were detected, yielding mean total VOC emission rates of 3.97 ± 0.70 μg m–2 h–1 from bare ice surfaces (n = 31), 1.63 ± 0.13 μg m–2 h–1 from cryoconite holes (n = 4), and 0.92 ± 0.08 μg m–2 h–1 from red snow (n = 2). No correlations were found between VOC emissions and ice surface algal counts, but a weak positive correlation (r = 0.43, p = 0.015, n = 31) between VOC emission rates from bare ice surfaces and incoming shortwave radiation was found. We propose that this may be due to the stress that high solar irradiance causes in bare ice microbial communities. Acetophenone, benzaldehyde, and phenylmaleic anhydride, all of which have reported antifungal activity, accounted for 51.1 ± 11.7% of emissions from bare ice surfaces, indicating a potential defense strategy against fungal infections. Greenland ice sheet microbial habitats are, hence, potential sources of VOCs that may play a role in supraglacial microbial interactions, as well as local atmospheric chemistry, and merit future research efforts.

The Programme for Monitoring of the Greenland Ice Sheet (PROMICE) provides surface meteorological and glaciological measurements from widespread on-ice automatic weather stations since mid-2007. In this study, we use 105 PROMICE ice-ablation time series to identify the timing of seasonal bare-ice onset preceded by snow cover conditions. From this collection, we find a bare-ice albedo at ice-ablation onset (here called bare-ice-onset albedo) of 0.565 ± 0.109 that has no apparent spatial dependence among 20 sites across Greenland. We then apply this snow-to-ice albedo transition value to measure the variations in daily Greenland bare-ice area in Sentinel-3 optical satellite imagery covering the extremely low and high respective melt years of 2018 and 2019. Daily Greenland bare-ice area peaked at 153 489 km² in 2019, 1.9 times larger than in 2018 (80 220 km²), equating to 9.0% (in 2019) and 4.7% (in 2018) of the ice sheet area.

Microbial communities and nutrient dynamics in glaciers and ice sheets continuously change as the hydrological conditions within and on the ice change. Glaciers and ice sheets can be considered bioreactors as microbiomes transform nutrients that enter these icy systems and alter the meltwater chemistry. Global warming is increasing meltwater discharge, affecting nutrient and cell export, and altering proglacial systems. In this review, we integrate the current understanding of glacial hydrology, microbial activity, and nutrient and carbon dynamics to highlight their interdependence and variability on daily and seasonal time scales, as well as their impact on proglacial environments.

Abstract Melting of the Greenland Ice Sheet is a leading cause of land-ice mass loss and cryosphere-attributed sea level rise. Blooms of pigmented glacier ice algae lower ice albedo and accelerate surface melting in the ice sheet’s southwest sector. Although glacier ice algae cause up to 13% of the surface melting in this region, the controls on bloom development remain poorly understood. Here we show a direct link between mineral phosphorus in surface ice and glacier ice algae biomass through the quantification of solid and fluid phase phosphorus reservoirs in surface habitats across the southwest ablation zone of the ice sheet. We demonstrate that nutrients from mineral dust likely drive glacier ice algal growth, and thereby identify mineral dust as a secondary control on ice sheet melting.

This study provides the first contemporaneous measurements of the concentration and speciation of dissolved nitrogen (N) in snow, meltwater and ice during the onset of the ablation season at a site within the Dark Zone of the Greenland Ice Sheet. The decaying, partially leached snow pack near S6 in south-west Greenland produced meltwater with relatively constant nitrate (NO3−) concentrations, approximating the snow pack average (1.1 µM). By contrast, ammonium (NH4+) (0–∼ 4 µM) and dissolved organic nitrogen (DON) (0–∼ 3 µM) concentrations were more variable, and sometimes higher than the average snow pack concentrations of 1.0 and 0.4 µM, respectively. This variability could be the result of microbial uptake and production within the melting snow pack. We observed pooled meltwater at the snow-ice interface that appeared to scavenge DON (∼1–8 µM) and possibly NO3− (∼1–2 µM) from the underlying ice, whose initial surface was a continuum of superimposed ice and weathering crust from the previous season. The shallow meteoric ice (∼10 cm–1 m) beneath the snow pack had high concentrations of DON and NH4+(6.5 and 2.6 µM, respectively), while NO3− concentrations were similar to the snow pack (1.1 µM). The absence of NH4+ in the snowmelt traversing the snow-ice interface may also point to microbial activity occurring at this boundary layer. Melt modelling indicated the presence of liquid meltwater at the snow-ice interface and that at least 10 cm of the surface ice below the snow pack was at 0°C. Solar radiation transmitted through the thin snow pack to the ice surface is absorbed by pigmented glacier algae causing melt of the surrounding ice, allowing the possibly of photosynthesis to begin under the thinning snowpack in these micro-melt environments. Hence, we conjecture that glacier algal blooms can commence before the snow pack has completely melted, occuring at a time when meltwaters are enhanced in nutrients scavenged from the snowpack, superimposed ice and the remnants of the weathering crust from the previous year.

Bacteriophages and their hosts are involved in a constant evolutionary arms race that should lead to divergence between phage genes over time. Here, the authors recruit metagenomic reads to virus reference genomes and genome fragments in samples from cryoconite holes and show that phages with near-identical core genomes maintain diversity by possession of numerous flexible gene modules, where homologous genes present in the pan-genome interchange to create new phage variants.

The Arctic environment is particularly affected by global warming, and a clear trend of the ice retreat is observed worldwide. In proglacial systems, the newly exposed terrain represents different environmental and nutrient conditions compared to later soil stages. Therefore, proglacial systems show several environmental gradients along the soil succession where microorganisms are active protagonists of the soil and carbon pool formation through nitrogen fixation and rock weathering. We studied the microbial succession of three Arctic proglacial systems located in Svalbard (Midtre Lovénbreen), Sweden (Storglaciären), and Greenland (foreland close to Kangerlussuaq). We analyzed 65 whole shotgun metagenomic soil samples for a total of more than 400 Gb of sequencing data. Microbial succession showed common trends typical of proglacial systems with increasing diversity observed along the forefield chronosequence. Microbial trends were explained by the distance from the ice edge in the Midtre Lovénbreen and Storglaciären forefields and by total nitrogen (TN) and total organic carbon (TOC) in the Greenland proglacial system. Furthermore, we focused specifically on genes associated with nitrogen fixation and biotic rock weathering processes, such as nitrogenase genes, obcA genes, and genes involved in cyanide and siderophore synthesis and transport. Whereas we confirmed the presence of these genes in known nitrogen-fixing and/or rock weathering organisms (e.g., Nostoc, Burkholderia), in this study, we also detected organisms that, even if often found in soil and proglacial systems, have never been related to nitrogen-fixing or rock weathering processes before (e.g., Fimbriiglobus, Streptomyces). The different genera showed different gene trends within and among the studied systems, indicating a community constituted by a plurality of organisms involved in nitrogen fixation and biotic rock weathering, and where the latter were driven by different organisms at different soil succession stages.

Glacier algae residing within the surface ice of glaciers and ice sheets play globally significant roles in biogeochemical cycling, albedo feedbacks, and melt of the world’s cryosphere. Here, we present an assessment of the macro-nutrient stoichiometry of glacier algal assemblages from the southwestern Greenland Ice Sheet (GrIS) margin, where widespread glacier algal blooms proliferate during summer melt seasons. Samples taken during the mid-2019 ablation season revealed overall lower cellular carbon (C), nitrogen (N), and phosphorus (P) content than predicted by standard microalgal cellular content:biovolume relationships, and elevated C:N and C:P ratios in all cases, with an overall estimated C:N:P of 1,997:73:1. We interpret lower cellular macro-nutrient content and elevated C:N and C:P ratios to reflect adaptation of glacier algal assemblages to their characteristic oligotrophic surface ice environment. Such lower macro-nutrient requirements would aid the proliferation of blooms across the nutrient poor cryosphere in a warming world. Up-scaling of our observations indicated the potential for glacier algal assemblages to accumulate ∼ 29 kg C km2 and ∼ 1.2 kg N km2 within our marginal surface ice location by the mid-ablation period (early August), confirming previous modeling estimates. While the long-term fate of glacier algal autochthonous production within surface ice remains unconstrained, data presented here provide insight into the possible quality of dissolved organic matter that may be released by assemblages into the surface ice environment.

Proglacial environments are ideal for studying the development of soils through the changes of rocks exposed by glacier retreat to weathering and microbial processes. Carbon (C) and nitrogen (N) contents as well as soil pH and soil elemental compositions are thought to be dominant factors structuring the bacterial, archaeal and fungal communities in the early stages of soil ecosystem formation. However, the functional linkages between C and N contents, soil composition and microbial community structures remain poorly understood. Here, we describe a multivariate analysis of geochemical properties and associated microbial community structures between a moraine and a glaciofluvial outwash in the proglacial area of a High Arctic glacier (Longyearbreen, Svalbard). Our results reveal distinct differences in developmental stages and heterogeneity between the moraine and the glaciofluvial outwash. We observed significant relationships between C and N contents, δ13Corg and δ15N isotopic ratios, weathering and microbial abundance and community structures. We suggest that the observed differences in microbial and geochemical parameters between the moraine and the glaciofluvial outwash are primarily a result of geomorphological variations of the proglacial terrain.

Glaciers retreating in response to climate warming are progressively exposing primary mineral substrates to surface conditions. As primary production is constrained by nitrogen (N) availability in these emerging ecosystems, improving our understanding of how N accumulates with soil formation is of critical concern. In this study, we quantified how the distribution and speciation of N, as well as rates of free-living biological N fixation (BNF), change along a 2000-year chronosequence of soil development in a High Arctic glacier forefield. Our results show the soil N pool increases with time since exposure and that the rate at which it accumulates is influenced by soil texture. Further, all N increases were organically bound in soils which had been ice-free for 0–50 years. This is indicative of N limitation and should promote BNF. Using the acetylene reduction assay technique, we demonstrated that microbially mediated inputs of N only occurred in soils which had been ice-free for 0 and 3 years, and that potential rates of BNF declined with increased N availability. Thus, BNF only supports N accumulation in young soils. When considering that glacier forefields are projected to become more expansive, this study has implications for understanding how ice-free ecosystems will become productive over time.

Cryoconite holes are miniature freshwater aquatic ecosystems that harbor a relatively diverse microbial community. This microbial community can withstand the extreme conditions of the supraglacial environment, including fluctuating temperatures, extreme and varying geochemical conditions and limited nutrients. We analyzed the physiological capabilities of microbial isolates from cryoconite holes from Antarctica, Greenland, and Svalbard in selected environmental conditions: extreme pH, salinity, freeze-thaw and limited carbon sources, to identify their physiological limits. The results suggest that heterotrophic microorganisms in cryoconite holes are well adapted to fast-changing environmental conditions, by surviving multiple freeze-thaw cycles, a wide range of salinity and pH conditions and scavenging a variety of organic substrates. Under oxic and anoxic conditions, the communities grew well in temperatures up to 30°C, although in anoxic conditions the community was more successful at colder temperatures (0.2°C). The most abundant cultivable microorganisms were facultative anaerobic bacteria and yeasts. They grew in salinities up to 10% and in pH ranging from 4 to 10.5 (Antarctica), 2.5 to 10 (Svalbard), and 3 to 10 (Greenland). Their growth was sustained on at least 58 single carbon sources and there was no decrease in viability for some isolates after up to 100 consecutive freeze-thaw cycles. The elevated viability of the anaerobic community in the lowest temperatures indicates they might be key players in winter conditions or in early melt seasons, when the oxygen is potentially depleted due to limited flow of meltwater. Consequently, facultative anaerobic heterotrophs are likely important players in the reactivation of the community after the polar night. This detailed physiological investigation shows that despite inhabiting a freshwater environment, cryoconite microorganisms are able to withstand conditions not typically encountered in freshwater environments (namely high salinities or extreme pH), making them physiologically more similar to arid soil communities. The results also point to a possible resilience of the most abundant microorganisms of cryoconite holes in the face of rapid change regardless of the location.

Snow packs cover large expanses of Earth’s land surface, making them integral components of the cryosphere in terms of past climate and atmospheric proxies, surface albedo regulators, insulators for other Arctic environments and habitats for diverse microbial communities such as algae, bacteria and fungi. Yet, most of our current understanding of snow pack environments, specifically microbial activity and community interaction, is limited to the main microbial growing season during spring ablation. At present, little is known about microbial activity and its influence on nutrient cycling during the subfreezing temperatures and 24-h darkness of the polar winter. Here, we examined microbial dynamics in a simulated cold (−5°C), dark snow pack to determine polar winter season microbial activity and its dependence on critical nutrients. Snow collected from Ny-Ålesund, Svalbard was incubated in the dark over a 5-week period with four different nutrient additions, including glacial mineral particles, dissolved inorganic nitrogen (DIN), dissolved inorganic phosphorus (DIP) and a combined treatment of DIN plus DIP. Data indicate a consumption of dissolved inorganic nutrients, particularly DIN, by heterotrophic communities, suggesting a potential nitrogen limitation, contradictory to phosphorus limitations found in most aquatic environments. 16S amplicon sequencing also reveal a clear difference in microbial community composition in the particulate mineral treatment compared to dissolved nutrient treatments and controls, suggesting that certain species of heterotrophs living within the snow pack are more likely to associate with particulates. Particulate phosphorus analyses indicate a potential ability of heterotrophic communities to access particulate sources of phosphorous, possibly explaining the lack of phosphorus limitation. These findings have importance for understanding microbial activity during the polar winter season and its potential influences on the abundance and bioavailability of nutrients released to surface ice and downstream environments during the ablation season.

Bacteria inhabiting non-polar glaciers are exposed to large variations in temperature, which significantly affects the fluidity of bacterial cell membranes. In order to maintain normal functions of the cell membranes, psychrophilic bacteria adapt by changing the composition of cell membrane fatty acids. However, information on the exact pattern of cell membrane adaptability in non-polar low-temperature habitats is scarce. In the present study, 42 bacterial strains were isolated from the Ghulmet, Ghulkin, and Hopar glaciers of the Hunza Valley in the Karakoram Mountain Range, Pakistan and their cell membrane fatty acid distributions studied, using gas chromatography/mass spectrometry (GC-MS) for the analysis of fatty acid methyl esters (FAMEs) liberated by acid-catalyzed methanolysis. Furthermore, Gram-negative and Gram-positive groups were grown under different temperature settings (5, 15, 25, and 35°C) in order to determine the effect of temperature on cell membrane (CM) fatty acid distribution. The analyses identified the major groups of cell membrane fatty acids (FA) as straight-chain monounsaturated fatty acids (n-MUFAs) and branched fatty acids (br-FAs), accounting for more than 70% of the fatty acids analyzed. The distribution of br-FAs and n-FAs in bacterial cell membranes was significantly affected by temperature, with the level of br-FAs decreasing relative to n-FAs with increasing temperature. Notably, the production of polyunsaturated fatty acids (PUFAs) was only seen at lower temperatures. This study contributes to understanding, for the first time, the role of br-FAs in the maintenance of cell membrane fluidity of bacteria inhabiting non-polar habitats.

Contributions of iron sources to Southern Ocean CO2sequestration during the last glacial period remain uncertain. Here, based on the biogeochemical analysis of subglacial calcites, the authors propose Antarctic volcanism, via subglacial drainage of Fe-rich waters, as a key contributor.

Abstract Glaciers and ice sheets, like other biomes, occupy a significant area of the planet and harbour biological communities with distinct interactions and feedbacks with their physical and chemical environment. In the case of the glacial biome, the biological processes are dominated almost exclusively by microbial communities. Habitats on glaciers and ice sheets with enough liquid water to sustain microbial activity include snow, surface ice, cryoconite holes, englacial systems and the interface between ice and overridden rock/soil. There is a remarkable similarity between the different specific glacial habitats across glaciers and ice sheets worldwide, particularly regarding their main primary producers and ecosystem engineers. At the surface, cyanobacteria dominate the carbon production in aquatic/sediment systems such as cryoconite holes, while eukaryotic Zygnematales and Chlamydomonadales dominate ice surfaces and snow dynamics, respectively. Microbially driven chemolithotrophic processes associated with sulphur and iron cycle and C transformations in subglacial ecosystems provide the basis for chemical transformations at the rock interface under the ice that underpin an important mechanism for the delivery of nutrients to downstream ecosystems. In this review, we focus on the main ecosystem engineers of glaciers and ice sheets and how they interact with their chemical and physical environment. We then discuss the implications of this microbial activity on the icy microbiome to the biogeochemistry of downstream ecosystems.

The Arctic is melting at an unprecedented rate and key drivers are changes in snow and ice albedo. Here, the authors show that red pigmented snow algae play a crucial role in decreasing surface albedo and their patterns for diversity, pigmentation, and consequently albedo, are ubiquitous across the Arctic.

Microorganisms play a significant role in changing the physical properties of the surface of the Greenland Ice Sheet. Cryoconite holes are a hotspot for this microbial activity, yet little is known about the REDOX conditions that develop within them. In this study, we used oxygen microelectrodes and microoptodes to measure for anoxic conditions at the microscale, for the first time revealing a potential niche for anaerobic microorganisms and anaerobic processes. The development of an anoxic zone 2 mm deep within a 6 mm-thick layer of cryoconite sediment was observed within an hour of disturbance, showing rapid acclimation to changing physical conditions. Long-term (half year) incubations of cryoconite material showed a peak of oxygen production and consumption after forty days and reached a low-activity, steady state by day 116, with a persisting anoxic zone beginning between 2 mm and 4 mm deep. Anaerobic microorganisms, which have received little attention to date, should therefore be considered an important component of the cryoconite ecosystem. We discuss the possible dynamics of oxygen concentrations in the supraglacial system and infer that anoxic zones are an important factor in the development of cryoconite sediment communities.

Organic matter (OM) has an essential biogeochemical influence along the hydrological continuum and within aquatic ecosystems. Organic matter derived via microbial processes was investigated within a range of model freshwater samples over a 10-day period. For this, excitation-emission matrix (EEM) fluorescence spectroscopy in combination with parallel factor (PARAFAC) analysis was employed. This research shows the origin and processing of both protein-like and humic-like fluorescence within environmental and synthetic samples over the sampling period. The microbial origin of Peak T fluorescence is demonstrated within both synthetic samples and in environmental samples. Using a range of incubation temperatures provides evidence for the microbial metabolic origin of Peak T fluorescence. From temporally resolved experiments, evidence is provided that Peak T fluorescence is an indication of metabolic activity at the microbial community level and not a proxy for bacterial enumeration. This data also reveals that humic-like fluorescence can be microbially derived in situ and is not solely of terrestrial origin, likely to result from the upregulation of cellular processes prior to cell multiplication. This work provides evidence that freshwater microbes can engineer fluorescent OM, demonstrating that microbial communities not only process, but also transform, fluorescent organic matter.

Cryoconite holes are foci of unusually high microbial diversity and activity on glacier surfaces worldwide, comprising melt-holes formed by the darkening of ice by biogenic granular debris. Despite recent studies linking cryoconite microbial community structure to the functionality of cryoconite habitats, little is known of the processes shaping the cryoconite bacterial community. In particular, the assertions that the community is strongly influenced by aeolian transfer of biota from ice-marginal habitats and the potential for cryoconite microbes to inoculate proglacial habitats are poorly quantified despite their longevity in the literature. Therefore, the bacterial community structures of cryoconite holes on three High-Arctic glaciers were compared to bacterial communities in adjacent moraines and tundra using terminal-restriction fragment length polymorphism. Distinct community structures for cryoconite and ice-marginal communities were observed. Only a minority of phylotypes are present in both habitat types, implying that cryoconite habitats comprise distinctive niches for bacterial taxa when compared to ice-marginal habitats. Curiously, phylotype abundance distributions for both cryoconite and ice-marginal sites best fit models relating to succession. Our analyses demonstrate clearly that cryoconites have their own, distinct functional microbial communities despite significant inputs of cells from other habitats.

Snow algae are poly-extremophilic microalgae and important primary colonisers and producers on glaciers and snow fields. Depending on their pigmentation they cause green or red mass blooms during the melt season. This decreases surface albedo and thus further enhances snow and ice melting. Although the phenomenon of snow algal blooms has been known for a long time, large aspects of their physiology and ecology sill remain cryptic. This study provides the first in-depth and multi-omics investigation of two very striking adjacent green and red snow fields on a glacier in Svalbard. We have assessed the algal community composition of green and red snow including their associated microbiota, i.e., bacteria and archaea, their metabolic profiles (targeted and non-targeted metabolites) on the bulk and single-cell level, and assessed the feedbacks between the algae and their physico-chemical environment including liquid water content, pH, albedo and nutrient availability. We demonstrate that green and red snow clearly vary in their physico-chemical environment, their microbial community composition and their metabolic profiles. For the algae this likely reflects both different stages of their life cycles and their adaptation strategies. Green snow represents a wet, carbon and nutrient rich environment and is dominated by the algae Microglena sp. with a metabolic profile that is characterized by key metabolites involved in growth and proliferation. In contrast, the dry and nutrient poor red snow habitat is colonised by various Chloromonas species with a high abundance of storage and reserve metabolites likely to face upcoming severe conditions. Combining a multitude of techniques we demonstrate the power of such complementary approaches in elucidating the function and ecology of extremophiles such as green and red snow algal blooms, which play crucial roles in glacial ecosystems.