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Polar regions harbor a diversity of cold-adapted (cryophilic) algae, which can be categorized into psychrophilic (obligate cryophilic) and cryotrophic (nonobligate cryophilic) snow algae. Both can accumulate significant biomasses on glacier and snow habitats and play major roles in global climate dynamics. Despite their significance, genomic studies on these organisms remain scarce, hindering our understanding of their evolutionary history and adaptive mechanisms in the face of climate change. Here, we present the draft genome assembly and annotation of the psychrophilic snow algal strain CCCryo 101-99 (cf. Sphaerocystis sp.). The draft haploid genome assembly is 122.5 Mb in length and is represented by 664 contigs with an N50 of 0.86 Mb, a Benchmarking Universal Single-Copy Orthologs (BUSCO) completeness of 92.9% (n = 1,519), a maximum contig length of 5.3 Mb, and a guanine-cystosine (GC) content of 53.1%. In total, 28.98% of the genome (35.5 Mb) contains repetitive elements. We identified 417 noncoding RNAs and annotated the chloroplast genome. The predicted proteome comprises 14,805 genes with a BUSCO completeness of 97.8%. Our preliminary analyses reveal a genome with a higher repeat content compared with mesophilic chlorophyte relatives, alongside enrichment in gene families associated with photosynthesis and flagella functions. Our current data will facilitate future comparative studies, improving our understanding of the likely response of polar algae to a warming climate as well as their evolutionary trajectories in permanently cold environments. [ABSTRACT FROM AUTHOR]

Microbial nitrogen fixation is crucial for building labile nitrogen stocks and facilitating higher plant colonisation in oligotrophic glacier forefield soils. Here, the diazotrophic bacterial community structure across four Arctic glacier forefields was investigated using metagenomic analysis. In total, 70 soil metagenomes were used for taxonomic interpretation based on 185 nitrogenase (nif) sequences, extracted from assembled contigs. The low number of recovered genes highlights the need for deeper sequencing in some diverse samples, to uncover the complete microbial populations. A key group of forefield diazotrophs, found throughout the forefields, was identified using a nifH phylogeny, associated with nifH Cluster I and III. Sequences related most closely to groups including Alphaproteobacteria, Betaproteobacteria, Cyanobacteria and Firmicutes. Using multiple nif genes in a Last Common Ancestor analysis revealed a diverse range of diazotrophs across the forefields. Key organisms identified across the forefields included Nostoc, Geobacter, Polaromonas and Frankia. Nitrogen fixers that are symbiotic with plants were also identified, through the presence of root associated diazotrophs, which fix nitrogen in return for reduced carbon. Additional nitrogen fixers identified in forefield soils were metabolically diverse, including fermentative and sulphur cycling bacteria, halophiles and anaerobes.

During the ablation season, active microbial communities colonise large areas of the Greenland Ice Sheet surface and produce dissolved organic matter (DOM) that may be exported downstream by surface melt. Meltwater flow through the bare ice interfluvial area, characterized by a porous weathering crust, is slow (~ 10-2 m d-1), meaning that it presents a potential site for photochemical and/or microbial alteration of supraglacial DOM. Transformations of supraglacial DOM during transport through the supraglacial drainage system remain unexplored, limiting our understanding of supraglacial DOM inputs to downstream subglacial and coastal ecosystems. Here, we employ negative-ion electrospray ionization 21 tesla Fourier transform ion cyclotron resonance mass spectrometry to catalogue the molecular composition of DOM in supraglacial dark ice, weathering crust meltwater, and supraglacial stream water sampled in a hydrologically connected supraglacial micro- catchment to address this knowledge gap. Dark ice DOM contained significantly more aromatic (25 ± 3 %) and less biolabile (13 ± 4 %) DOM than weathering crust meltwater (3 ± 0 and 50 ± 0 %, respectively), pointing to retention of DOM on the ice surface and microbial, as well as photochemical alteration of DOM during transit through the supraglacial drainage system. These findings have implications for our understanding of supraglacial biogeochemical cycling, highlighting the importance of including the weathering crust photic zone when assessing supraglacial inputs to subglacial and downstream ecosystems. [ABSTRACT FROM AUTHOR]

Microbial blooms colonize the Greenland Ice Sheet bare ice surface during the ablation season and significantly reduce its albedo. On the ice surface, microbes are exposed to high levels of irradiance, freeze–thaw cycles, and low nutrient concentrations. It is well known that microorganisms secrete metabolites to maintain homeostasis, communicate with other microorganisms, and defend themselves. Yet, the exometabolome of supraglacial microbial blooms, dominated by the pigmented glacier ice algae Ancylonema alaskanum and Ancylonema nordenskiöldii, remains thus far unstudied. Here, we use a high‐resolution mass spectrometry‐based untargeted metabolomics workflow to identify metabolites in the exometabolome of microbial blooms on the surface of the southern tip of the Greenland Ice Sheet. Samples were collected every 6 h across two diurnal cycles at 5 replicate sampling sites with high similarity in community composition, in terms of orders and phyla present. Time of sampling explained 46% (permutational multivariate analysis of variance [PERMANOVA], pseudo‐F = 3.7771, p = 0.001) and 27% (PERMANOVA, pseudo‐F = 1.8705, p = 0.001) of variance in the exometabolome across the two diurnal cycles. Annotated metabolites included riboflavin, lumichrome, tryptophan, and azelaic acid, all of which have demonstrated roles in microbe–microbe interactions in other ecosystems and should be tested for potential roles in the development of microbial blooms on bare ice surfaces. [ABSTRACT FROM AUTHOR]

The microbiome of Greenland Ice Sheet supraglacial habitats is still underinvestigated, and as a result there is a lack of representative genomes from these environments. In this study, we investigated the supraglacial microbiome through a combination of culturing-dependent and -independent approaches. We explored ice, cryoconite, biofilm, and snow biodiversity to answer: (1) how microbial diversity differs between supraglacial habitats, (2) if obtained bacterial genomes reflect dominant community members, and (3) how culturing versus high throughput sequencing changes our observations of microbial diversity in supraglacial habitats. Genomes acquired through metagenomic sequencing (133 high-quality MAGs) and whole genome sequencing (73 bacterial isolates) were compared to the metagenome assemblies to investigate abundance within the total environmental DNA. Isolates obtained in this study were not dominant taxa in the habitat they were sampled from, in contrast to the obtained MAGs. We demonstrate here the advantages of using metagenome SSU rRNA genes to reflect whole-community diversity. Additionally, we demonstrate a proof-of-concept of the application of in situ culturing in a supraglacial setting. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

We examined the response of bacteria and proto- and metazooplankton to photomodified dissolved organic matter (DOM). Sterile filtered water from a eutrophic and a humic lake, that was either exposed to artificial ultraviolet (UV) radiation or kept dark, was added to semicontinuous laboratory microcosms that lasted for 7 weeks. Bacterial production responded positively to photochemical modification of DOM regardless of lake type. Final heterotrophic biomass (bacteria+proto+metazooplankton) was 47 [+ or -] 5 and 37 [+ or -] 5 [micro]g carbon (C) [L.sup.-1] in microcosms with UV-exposed and unaltered eutrophic water DOM and 15 [+ or -] 4 and 11 [+ or -] 2 [micro]g C [L.sup.-1] in microcosms with UV-exposed and unaltered humic water DOM, respectively. For the eutrophic water, there were no significant differences in proto- or metazooplankton biomasses between microcosms receiving UV-exposed or nonexposed DOM. Differences between eutrophic water microcosms were not significant when flagellates, ciliates, cladocerans, and copepods were examined separately. In microcosms with UV-exposed humic water, biomasses of heterotrophic flagellates, rotifers, nauplii, and cladocerans were higher than in those with nonexposed DOM. Higher final metazooplankton biomass following addition of UV-exposed humic water indicates that photochemically modified DOM can be effectively transferred through the microbial loop.

The bacterial and viral abundances were measured in 24 lakes with dissolved organic carbon (DOC) concentrations ranging from 3 to 19 mg of C liter(super -1). The results indicate that the way viruses control bacterial assemblages in humic lakes is different from the mechanism in clear water systems.

Summary: 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]