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Algal blooms occur during the summer melt on the Greenland Ice Sheet and on other melting supraglacial environments globally. Snow habitats are mostly inhabited by chlorophytes (Chlorophyceae and Trebouxiophyceae), while bare ice is dominated by streptophytes (Zygnematophyceae). These eukaryotes thrive at low temperatures and under high light and low nutrients, and they have specialised cellular mechanisms allowing for life under extreme conditions. However, little empirical data exists about the cellular adaptations of snow and glacial ice algae under these conditions, despite our growing knowledge of species diversity and associated abiotic conditions. We address this knowledge gap by identifying protein groups essential for the maintenance of cellular homeostasis under relevant environmental conditions from samples rich in snow and glacial ice algae from the Greenland Ice Sheet. Samples collected during summer algal blooms were subjected to a metaproteomics workflow using an established protein extraction protocol and LC-MS/MS analysis. Proteins were identified using a predicted protein database built from RNA sequencing data comprising of sequences from algae, but also other microorganisms present in the community such as bacteria and fungi. We assigned 35% of the recorded MS2 spectra to predominantly algal proteins, as well as bacterial and fungal proteins, corresponding to more than 5800 protein groups. This large dataset provides a starting point for dissecting cellular functions of cryogenic algae in these environments and allows us, for example, to evaluate the relative abundance of proteins linked to specific cellular pathways such as photosynthesis or lipid metabolism.

Phototrophic organisms blooming during the summer melt season on snow and ice surfaces are dominated by eukaryotic green algae (Chlorophytes and Streptophytes, respectively), with Cyanobacteria restricted to cryoconite habitats. However, the role and interactions between the algae and other light-harvesting organisms are largely understudied in these ecosystems.We searched metagenomes of snow and ice samples collected from the Greenland Ice Sheet during the summer melting season for signatures indicating anoxygenic photosystems, which are used by certain groups of bacteria to gain energy from light without releasing oxygen. Two metagenome assembled genomes (MAGs) carrying all genes necessary to perform anoxygenic photosynthesis and carbon-fixation were found. Whole-genome phylogenetic comparison placed the MAGs within the Alpha- and Gamma-proteobacteria, which was confirmed by alignment of the respective functional marker genes pufL and pufM to known sequences from cultured anoxygenic phototrophic bacteria. The identified functions and phylogeny suggest that the MAGs belong within the group of purple non-sulfur bacteria, a pigmented and metabolically versatile group of bacteria often found in shallow aqueous environments, but very little is documented in cryogenic environments. Our data show that these procaryotic organisms are preferably linked to glacial ice algae habitats and, to a lesser extent, to algae in snow habitats. Our results pose intriguing ecological questions for supraglacial habitats, such as the contribution of these procaryotes to the ongoing biological darkening of ice surfaces or the potential mutualistic light-harvesting strategies on ice, as the used wavelength of purple non-sulfur bacteria are complementary to those used by indigenous high abundant glacial ice algae.

Pigment-producing microorganisms are prevalent on glacier surfaces, decreasing the snow and ice albedo. This impacts the absorption of solar radiation and accelerates rates of glacier surface melting. Studying the glacier surface ecosystem is important to understand the effects of anthropogenic climate change, and to further our knowledge of how glacier-dwelling organisms and their evolution impact downstream ecosystems. Recent studies have revealed links between habitat type, seasonality, physicochemical characteristics and the microbial composition of the supraglacial environment. However, these studies are limited in number, time points and locations. Thus, global drivers of supraglacial microbial community composition remain unknown. To fill this gap, we used data produced by our team over the last five years, as well as gathered from public repositories, to investigate the prokaryotic and eukaryotic composition of supraglacial environments worldwide.We used 18S and 16S rRNA gene amplicon sequencing to study the microbial composition and diversity of more than eight hundred surface snow, ice and cryoconite samples from glaciers and snowfields all over the world, including Arctic, Antarctic and temperate glaciers. Results reveal a worldwide core microbiome specific to this environment, composed of generalist, freshwater and cold-adapted taxa. Distance-decay and latitudinal patterns can be identified, but key factors in determining microbial community diversity and composition rest on local to regional biogeographical scales. Habitat biology shows different responses to biogeographical drivers, likely influenced by their structure: cryoconite communities present a higher distance-decay and location specialization than snow and ice communities. In addition, communities of prokaryotes are less location-specific than those of eukaryotes. This study highlights the need for further investigation into the drivers of microbial dissemination onto glaciers and their response to local biogeography.

Glaciers represent a unique inventory of microbial genetic diversity and a record of evolution. The Tibetan Plateau contains the largest area of low-latitude glaciers and is particularly vulnerable to global warming. By sequencing 85 metagenomes and 883 cultured isolates from 21 Tibetan glaciers covering snow, ice and cryoconite habitats, we present a specialized glacier microbial genome and gene catalog to archive glacial genomic and functional diversity. This comprehensive Tibetan Glacier Genome and Gene (TG2G) catalog includes 883 genomes and 2,358 metagenome-assembled genomes, which represent 968 candidate species spanning 30 phyla. The catalog also contains over 25 million non-redundant protein-encoding genes, the utility of which is demonstrated by the exploration of secondary metabolite biosynthetic potentials, virulence factor identification and global glacier metagenome comparison. The TG2G catalog is a valuable resource that enables enhanced understanding of the structure and functions of Tibetan glacial microbiomes.

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.

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.

Snow algae are predicted to expand in polar regions due to climate warming, which can accelerate snowmelt by reducing albedo. Green snow frequently occurs near penguin colonies, and red snow distributes widely along ocean shores. However, the mechanisms underpinning the assemblage of algae and heterotrophs in colored snow remain poorly characterized. We investigated algal, bacterial, and fungal communities and their interactions in red and green snows in the Antarctic Peninsula using a high-throughput sequencing method. We found distinct algal community structure in red and green snows, and the relative abundance of dominant taxa varied, potentially due to nutrient status differences. Contrastingly, red and green snows exhibited similar heterotrophic communities (bacteria and fungi), whereas the relative abundance of fungal pathogens was substantially higher in red snow by 3.8-fold. Red snow exhibited a higher network complexity, indicated by a higher number of nodes and edges. Red snow exhibited a higher proportion of negative correlations among heterotrophs (62.2% vs 3.4%) and stronger network stability, suggesting the red-snow network is more resistant to external disturbance. Our study revealed that the red snow microbiome exhibits a more stable microbial network than the green snow microbiome.

Glacier and ice sheet surfaces host diverse communities of microorganisms whose activity (or inactivity) influences biogeochemical cycles and ice melting. Supraglacial microbes endure various environmental extremes including resource scarcity, frequent temperature fluctuations above and below the freezing point of water, and high UV irradiance during summer followed by months of total darkness during winter. One strategy that enables microbial life to persist through environmental extremes is dormancy, which despite being prevalent among microbial communities in natural settings, has not been directly measured and quantified in glacier surface ecosystems. Here, we use a combination of metabarcoding and metatranscriptomic analyses, as well as cell-specific activity (BONCAT) incubations to assess the diversity and activity of microbial communities from glacial surfaces in Iceland and Greenland. We also present a new ecological model for glacier microorganisms and simulate physiological state-changes in the glacial microbial community under idealized (i) freezing, (ii) thawing, and (iii) freeze-thaw conditions. We show that a high proportion (>50%) of bacterial cells are translationally active in-situ on snow and ice surfaces, with Actinomycetota, Pseudomonadota, and Planctomycetota dominating the total and active community compositions, and that glacier microorganisms, even when frozen, could resume translational activity within 24 h after thawing. Our data suggest that glacial microorganisms respond rapidly to dynamic and changing conditions typical of their natural environment. We deduce that the biology and biogeochemistry of glacier surfaces are shaped by processes occurring over short (i.e., daily) timescales, and thus are susceptible to change following the expected alterations to the melt-regime of glaciers driven by climate change. A better understanding of the activity of microorganisms on glacier surfaces is critical in addressing the growing concern of climate change in Polar regions, as well as for their use as analogues to life in potentially habitable icy worlds.

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.

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.

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.

In the present study, cyanobacterium isolate CHS1 isolated from Hopar glacier, Pakistan, was analyzed for the first time for cell membrane fatty acids and production of pigments. Sequencing of the 16-23S intergenetic region confirmed identification of the isolate CHS1 as Nodularia spumigena. All chlorophyll and carotenoid pigments were quantified using high-performance liquid chromatography and experiments to test tolerance against a range of physico-chemical conditions were conducted. Likewise, the fatty acid profile of the cell membrane CHS1 was analyzed using gas chromatography and mass spectroscopy. The cyanobacterium isolate CHS1 demonstrated tolerance to 8 g/L% NaCl, 35°C and pH 5–9. The characteristic polyunsaturated fatty acid (PUFA) of isolate CHS1, C18:4, was observed in fatty acid methyl esters (FAMEs) extracted from the cell membrane. CHS1 was capable of producing saturated fatty acids (SFA) (e.g., C16:0), monounsaturated fatty acids (MUFA) (e.g., C18:1) and polyunsaturated fatty acids (e.g., C20:5) in the cell membrane. In this study, we hypothesize that one mechanism of cold adaptation displayed by isolate CHS1 is the accumulation of high amounts of PUFA in the cell membrane.

Algal blooms are natural phenomena that may cause human health problems, millions of dollars in losses and ecological disasters worldwide. Anthropogenic pressures like eutrophication may increase the frequency and intensity of these phenomena. The Black Sea is characterized by rapid changes in salinity and temperature in surface waters. In addition, it has suffered increasing environmental pressure from human activities. This work presents the first cultures of Gymnodinium aureolum to be isolated from the Black Sea. Morphological and phylogenetic analyses confirmed our strain as G. aureolum. The effects of temperature and salinity on growth were tested in experiments combining two temperatures and five salinities in 10 experimental treatments. This provides baseline data on the physiological adaption and acclimatization potential of the species to bloom under present and future climatic scenarios in the Black Sea. Gymnodinium aureolum grew exponentially in all experimental treatments, except for cultures at salinity 5. Growth rate increased significantly with increasing temperature reaching the maximum at 20 °C and salinity 15 (0.38 ± 0.02 d−1). This suggests an adaptation to the salinity and temperature of Black Sea waters and, together with previous records of G. aureolum in both water and sediments, supports the idea that this may be a bloom-forming population of G. aureolum.

The global cryosphere, Earth's frozen water, is in precipitous decline. The ongoing and predicted impacts of cryosphere loss are diverse, ranging from disappearance of entire biomes to crises of water availability. Covering approximately one-fifth of the planet, mass loss from the terrestrial cryosphere is driven primarily by a warming atmosphere but reductions in albedo (the proportion of reflected light) also contribute by increasing absorption of solar radiation. In addition to dust and other abiotic impurities, biological communities substantially reduce albedo worldwide. In this review, we provide a global synthesis of biological albedo reduction (BAR) in terrestrial snow and ice ecosystems. We first focus on known drivers—algal blooms and cryoconite (granular sediment on the ice that includes both mineral and biological material)—as they account for much of the biological albedo variability in snow and ice habitats. We then consider an array of potential drivers of BAR whose impacts may be overlooked, such as arthropod deposition, resident organisms (e.g., dark-bodied glacier ice worms), and larger vertebrates, including humans, that transiently visit the cryosphere. We consider both primary (e.g., BAR due to the presence of pigmented algal cells) and indirect (e.g., nutrient addition from arthropod deposition) effects, as well as interactions among biological groups (e.g., birds feeding on ice worms). Collectively, we highlight that in many cases, overlooked drivers and interactions among factors have considerable potential to alter BAR, perhaps rivaling the direct effects of algal blooms and cryoconite. We conclude by highlighting knowledge gaps for the field with an emphasis on the underrepresentation of genomic tools, understudied areas (particularly high-elevation glaciers at tropical latitudes), and a dearth of temporal sampling in current efforts. We detail a global framework for long-term BAR monitoring that, if implemented, would yield a tremendous amount of insight for BAR and would be particularly valuable in light of the rapid ecological and physical changes occurring in the contemporary cryosphere.

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., SCOPUS: ar.j, info:eu-repo/semantics/published

Bacteriophage genomes rapidly evolve via mutation and horizontal gene transfer to counter evolving bacterial host defenses; such arms race dynamics should lead to divergence between phages from similar, geographically isolated ecosystems. However, near-identical phage genomes can reoccur over large geographical distances and several years apart, conversely suggesting many are stably maintained. Here, we show that phages with near-identical core genomes in distant, discrete aquatic ecosystems maintain diversity by possession of numerous flexible gene modules, where homologous genes present in the pan-genome interchange to create new phage variants. By repeatedly reconstructing the core and flexible regions of phage genomes from different metagenomes, we show a pool of homologous gene variants co-exist for each module in each location, however, the dominant variant shuffles independently in each module. These results suggest that in a natural community, recombination is the largest contributor to phage diversity, allowing a variety of host recognition receptors and genes to counter bacterial defenses to co-exist for each phage., 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.

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.

The surface (supraglacial) environment of the Greenland Ice Sheet (GrIS) is an active site for the storage, transformation and transport of carbon, which is driven by extremely high levels of solar radiation throughout the ablation season. Within the south west of the GrIS, blooms of Streptophyte micro-algae (hereafter glacier algae) at abundances of ~ 105 cell mL−1 dominate primary production in the surface ice and provide dissolved organic matter (DOM) to the heterotrophic bacterial community. Glacier algae contain photoprotective secondary phenolic pigment that comprises a large proportion of the cell (~ 4 % of the dry weight) and could represent a substantial, additional carbon source for the heterotrophic community. The transformation and degradation of DOM by solar radiation (photodegradation) and heterotrophic communities (biodegradation) represent two crucial controls on DOM composition and quantity; however, the influence of these processes within the surface ice is yet to be constrained. This study therefore assessed responses in the composition and quantity of two carbon sources (glacier algae secondary pigment and surface ice DOM) following exposure to UV, PAR, UV+PAR (photodegradation) and subsequent incubation with bacterial communities isolated from the ambient environment (biodegradation). Our results indicate that exposure to predominantly UV radiation altered the composition of glacier algal pigment and surface ice DOM; however, the quantity of DOM remained constant. Biodegradation caused the greatest changes to both DOM composition and quantity, particularly in surface ice DOM. Secondary pigment extracted from glacier algae was not a highly bioavailable source of carbon and did not support significant growth of surface ice heterotrophic bacterial communities. Conversely, low molecular weight compounds in surface ice DOM were rapidly utilised by heterotrophic bacteria supporting between a 3 and 9-fold increase in bacterial abundance over a 30-day incubation. We found that photodegradation of glacier algal pigment and surface ice DOM did not influence heterotrophic consumption. Photodegradation and biodegradation of DOM in the surface ice habitat are likely intimately linked and act as fundamental controls on the composition and quantity of DOM exported to downstream environments.

International audience; 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.

Ice sheets overlay active and putatively widespread microbial ecosystems. An active subglacial biota has the potential to impact strongly on the (bio)geochemistry of local as well as downstream environments. Such impacts partly depend on the distribution of microbial populations, the types of habitats present beneath the ice, and their connectivity. In the ablation zone of the Greenland Ice Sheet (GrIS), supraglacial meltwaters are routed to the ice-sheet bed during the melt season, flushing out subglacial waters, sediments, and cells to proglacial environments via runoff. Here, we report on the diversity, composition, and niche differentiation of microbial assemblages exported in bulk runoff from a large (~600 km2) GrIS catchment. Proglacial river samples were collected over a period of subglacial drainage evolution in order to capture potential shifts in exported microbial community alongside hydrochemical transitions. We use high-resolution hydrochemical and hydrological information from the proglacial river to guide microbial (16S rRNA gene) interpretations. Core populations closely matched sequences previously isolated from other (pro)glacial environments, and phylogenetic characterisation of main OTUs alluded to a central role for subglacial iron, sulphur, and methane cycling. Whilst results indicate that bulk populations exported are likely true members of sub ice-sheet communities, we also find evidence of a supraglacial signature influencing composition of exported assemblages. Changes in assemblage structure accompanied those of major hydrological periods, with enhanced subglacial flushing coinciding with distinct shifts in microbial composition. Timing of sampling therefore matters when attempting to infer more nuanced changes in exported communities, or reveal the biogeochemical processes likely occurring in regions of the bed less influenced by surface melt. This is likely especially true when studying larger glacial systems, which experience complex hydrological changes throughout the melt-season, and that periods of extensive subglacial flushing offer opportunities to assess diversity from more isolated regions of the bed. Still, an apparent strong buffering signal from marginal zones appear to mask some of the diversity intrinsic to more remote, likely anoxic, subglacial niches, which may ultimately only be sampled via direct access to the subsurface.

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.

Large blooms of purple-brownish pigmented glacier algae cover the ablation zones of the Greenland Ice Sheet (GrIS) and amplify its melt by lowering the ice surface albedo and increasing its solar radiation absorption. The darkening effect of these Zygnematophycean algae can be mainly attributed to their phenolic pigments, which absorb in the visible (VIS) and UV light ranges. Currently, a mechanistic understanding of the factors regulating the production of these pigments and their implications for the large-scale biologically-driven albedo reduction on the GrIS is missing. Here, we reveal how light (VIS vs. UV range) controls the phenolic pigment production, endo- and exometabolome, gene expression and photosynthetic performance of glacier algae. Two different algal communities (a mixed natural microbial community collected from snow-free ice and a laboratory-grown community of the ice algae Mesotaenium berggrenii without its original dark pigmentation) were used for a set of in situ incubations on Mittivakkat glacier in SE-Greenland. Pulse-amplitude-modulated (PAM) fluorometry revealed an overall higher photosynthetic performance (electron transport rate) at higher irradiances for the field population containing purpurogallin-like pigments compared to the lab community without dark pigmentation. The lab population showed a low maximum quantum efficiency of photosystem II under in situ light conditions, indicating a photo-damaging effect from high intensities of UV light in the absence of purpurogallin-derived phenolic pigments. Our study highlights the intracellular shading effect by purpurogallin-derived pigments, which are key for the survival of glacier algae on the ice and forms a cornerstone of understanding the large-scale variability in the biological darkening of the GrIS.

Significance Processes that darken the surface of the Greenland Ice Sheet (GrIS) enhance energy absorption and accelerate melt, with consequences for global sea-level rise. Here, we demonstrate how summer blooms of “glacier algae” darken the ice surface, significantly impacting the physical integrity of the environment. We identify and quantify the energy regulation mechanisms employed by glacier algae to balance their requirements for photosynthesis and growth with the extreme light and temperature regime of the GrIS, demonstrating how these mechanisms are optimized to darken and melt the ice surface. Our findings are critical for the incorporation of biological feedbacks into predictive models of GrIS surface runoff and provide unique insight into how photoautotrophic life excels within icy environments., Blooms of Zygnematophycean “glacier algae” lower the bare ice albedo of the Greenland Ice Sheet (GrIS), amplifying summer energy absorption at the ice surface and enhancing meltwater runoff from the largest cryospheric contributor to contemporary sea-level rise. Here, we provide a step change in current understanding of algal-driven ice sheet darkening through quantification of the photophysiological mechanisms that allow glacier algae to thrive on and darken the bare ice surface. Significant secondary phenolic pigmentation (11 times the cellular content of chlorophyll a) enables glacier algae to tolerate extreme irradiance (up to ∼4,000 µmol photons⋅m−2⋅s−1) while simultaneously repurposing captured ultraviolet and short-wave radiation for melt generation. Total cellular energy absorption is increased 50-fold by phenolic pigmentation, while glacier algal chloroplasts positioned beneath shading pigments remain low-light–adapted (Ek ∼46 µmol photons⋅m−2⋅s−1) and dependent upon typical nonphotochemical quenching mechanisms for photoregulation. On the GrIS, glacier algae direct only ∼1 to 2.4% of incident energy to photochemistry versus 48 to 65% to ice surface melting, contributing an additional ∼1.86 cm water equivalent surface melt per day in patches of high algal abundance (∼104 cells⋅mL−1). At the regional scale, surface darkening is driven by the direct and indirect impacts of glacier algae on ice albedo, with a significant negative relationship between broadband albedo (Moderate Resolution Imaging Spectroradiometer [MODIS]) and glacier algal biomass (R2 = 0.75, n = 149), indicating that up to 75% of the variability in albedo across the southwestern GrIS may be attributable to the presence of glacier algae.

Rock glaciers (RG) are assumed to influence the biogeochemistry of downstream ecosystems because of the high ratio of rock:water in those systems, but no studies have considered the effects of a RG inflow on the microbial ecology of sediments in a downstream pond. An alpine RG-pond system, located in the NW Italian Alps has been chosen as a model, and Bacteria and Archaea 16S rRNA genes abundance, distribution and diversity have been assessed by qPCR and Illumina sequencing, coupled with geochemical analyses on sediments collected along a distance gradient from the RG inflow. RG surface material and neighbouring soil have been included in the analysis to better elucidate relationships among different habitats. Our results showed that different habitats harboured different, well-separated microbial assemblages. Across the pond, the main variations in community composition (e.g. Thaumarchaeota and Cyanobacteria relative abundance) and porewater geochemistry (pH, DOC, TDN and NH4+) were not directly linked to RG proximity, but to differences in water depth. Some microbial markers potentially linked to the presence of meltwater inputs from the RG have been recognised, although the RG seems to have a greater influence on the pond microbial communities due to its contribution in terms of sedimentary material.

Glaciers and ice sheets are significant sources of dissolved organic carbon and nutrients to downstream subglacial and marine ecosystems. Climatically driven increases in glacial runoff are expected to intensify the impact of exported nutrients on local and regional downstream environments. However, the origin and bioreactivity of dissolved organic carbon from glacier surfaces are not fully understood. Here, we present simultaneous measurements of gross primary production, community respiration, dissolved organic carbon composition and export from different surface habitats of the Greenland ice sheet, throughout the ablation season. We found that microbial production was significantly correlated with the concentration of labile dissolved organic species in glacier surface meltwater. Further, we determined that freely available organic compounds made up 62% of the dissolved organic carbon exported from the glacier surface through streams. We therefore conclude that microbial communities are the primary driver for labile dissolved organic carbon production and recycling on glacier surfaces, and that glacier dissolved organic carbon export is dependent on active microbial processes during the melt season. Glacial systems are important sources of dissolved organic carbon to downstream ecosystems. Observations of carbon dynamics on the Greenland ice sheet reveal substantial melt season production and export of microbial dissolved organic carbon.

Hybrid assembly of Illumina and Oxford Nanopore sequencing was used here to produce the complete circular genome and plasmid sequences of Salmonella enterica subsp. enterica serovar Enteritidis PT1 (phage type 1). The organism was obtained from the Salmonella Reference Laboratory at Public Health England, Colindale, UK.

Glaciers and ice sheets host abundant and dynamic communities of microorganisms on the ice surface (supraglacial environments). Recently, it has been shown that Streptophyte ice algae blooming on the surface ice of the south-west coast of the Greenland Ice Sheet are a significant contributor to the 15-year marked decrease in albedo. Currently little is known about the constraints, such as the nutrient cycling, on this large-scale algal bloom. In this study, we present a preliminary data set that investigates the conversion of dissolved inorganic nutrients to the dissolved organic phase occurring in these darkening surface ice environments. Our results show a clear dominance of the organic phase, with 93 % of the total dissolved nitrogen and 67 % of the total dissolved phosphorus in the organic phase. Correlations between algal abundance and dissolved organic carbon and nitrogen, indicate ice algae are driving the dissolved nutrient phase shift occurring in these supraglacial environments. Dissolved organic nutrient ratios in these supraglacial environments are notably higher than the Redfield Ratio (DON : DOP = 49, 78, 116) and DOC : DOP = 797, 1166, 2013), suggesting these environments may be phosphorus limited.

Distinct microbial habitats on glacial surfaces are dominated by snow and ice algae, which are the critical players and the dominant primary colonisers and net producers during the melt season. Here we have for the first time evaluated the role of these algae in association with the full microbial community composition (i.e., algae, bacteria, archaea) in distinct surface habitats and on twelve glaciers and permanent snow fields in Svalbard and Arctic Sweden. We cross-correlated these data with the analyses of specific metabolites such as fatty acids and pigments, and a full suite of potential critical physico-chemical parameters including major and minor nutrients, and trace metals. We show that correlations between single algal species, metabolites, and specific geochemical parameters can be used to unravel mixed metabolic signals in complex communities, further assign them to single species and infer their functionality. The data also clearly show that the production of metabolites in snow and ice algae is driven mainly by nitrogen and less so by phosphorus limitation. This is especially important for the synthesis of secondary carotenoids, which cause a darkening of glacial surfaces leading to a decrease in surface albedo and eventually higher melting rates. This article is protected by copyright. All rights reserved.