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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.

Bacterial communities form the basis of biogeochemical processes and determine plant growth and health. Mosses harbour diverse bacterial communities that are involved in nitrogen fixation and carbon cycling. Global climate change is causing changes in aboveground plant biomass and shifting species composition in the Arctic, but little is known about the response of moss microbiomes in these environments. Here, we studied the total and potentially active bacterial communities associated with Racomitrium lanuginosum in response to a 20-yr in situ warming in an Icelandic heathland. We evaluated the effect of warming and warming-induced shrub expansion on the moss bacterial community composition and diversity, and nifH gene abundance. Warming changed both the total and the potentially active bacterial community structure, while litter abundance only affected the total bacterial community structure. The abundance of nifH genes was negatively affected by litter abundance. We also found shifts in the potentially nitrogen-fixing community, with Nostoc decreasing and noncyanobacterial diazotrophs increasing in relative abundance. Our data suggest that the moss microbial community and potentially nitrogen fixing taxa will be sensitive to future warming, partly via changes in litter and shrub abundance.

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.

In Arctic regions, thawing permafrost soils are projected to release 50 to 250 Gt of carbon by 2100. This data is mostly derived from carbon-rich wetlands, although 71% of this carbon pool is stored in faster-thawing mineral soils, where ecosystems close to the outer boundaries of permafrost regions are especially vulnerable. Although extensive data exists from currently thawing sites and short-term thawing experiments, investigations of the long-term changes following final thaw and co-occurring drainage are scarce. Here we show ecosystem changes at two comparable tussock tundra sites with distinct permafrost thaw histories, representing 15 and 25 years of natural drainage, that resulted in a 10-fold decrease in CH4 emissions (3.2 ± 2.2 vs. 0.3 ± 0.4 mg C-CH4 m−2 day−1), while CO2 emissions were comparable. These data extend the time perspective from earlier studies based on short-term experimental drainage. The overall microbial community structures did not differ significantly between sites, although the drier top soils at the most advanced site led to a loss of methanogens and their syntrophic partners in surface layers while the abundance of methanotrophs remained unchanged. The resulting deeper aeration zones likely increased CH4 oxidation due to the longer residence time of CH4 in the oxidation zone, while the observed loss of aerenchyma plants reduced CH4 diffusion from deeper soil layers directly to the atmosphere. Our findings highlight the importance of including hydrological, vegetation and microbial specific responses when studying long-term effects of climate change on CH4 emissions and underscores the need for data from different soil types and thaw histories.

Mosses are among the first colonizing organisms after glacier retreat and can develop into thick moss mats during later successional stages. They are key players in N2 fixation through their microbiome, which is an important process for nutrient build-up during primary succession. How these moss-microbe interactions develop during succession is not well-studied and is relevant in the light of climate change and increased glacier retreat.We examined how the bacterial communities associated with two moss species of the genus Racomitrium and the underlying substrate, as well as moss traits and nitrogen fixation, develop along a successional gradient in the glacier forefield of Fláajökull in southeast Iceland. In addition, tested whether moss functional traits, such as total carbon (TC) and nitrogen contents (TN) are drivers of moss and underlying soil bacterial communities.Although time since deglaciation did not affect TN and moisture content, TC and shoot length increased with time since deglaciation. Moss and underlying soil bacterial communities were distinct. While the soil bacterial community structure was driven by the time since deglaciation and moss C/N ratios, the moss bacterial community structure was linked to time since deglaciation and moss moisture content. Moss N2-fixation rates were linked to bacterial community composition and nifH gene abundance rather than moss TN or time since deglaciation. This was accompanied by a shift from autotrophic to heterotrophic diazotrophs.Overall, our results suggest that there is little lateral transfer between moss and soil bacterial communities and that moss traits and time since deglaciation affect moss and soil bacterial community structure. In addition, moss N2-fixation rates are determined by bacterial community structure, rather than moss traits or time since deglaciation.

Microbial analysis at the micro scale of soil is essential to the overall understanding of microbial organization and interactions, and necessary for a better understanding of soil ecosystem functioning. While bacterial communities have been extensively described, little is known about the organization of fungal communities as well as functional potentials at scales relevant to microbial interactions. Fungal and bacterial communities and changes in nitrogen cycling potentials in the pristine Rothamsted Park Grass soil (bulk soil) as well as in its particle size sub-fractions (PSFs; > 250 μm, 250-63 μm, 63-20 μm, 20-2 μm, < 2 μm and supernatant) were studied. The potential for nitrogen reduction was found elevated in bigger aggregates. The relative abundance of Basidiomycota deceased with decreasing particle size, Ascomycota showed an increase and Mucoromycota became more prominent in particles less than 20 μm.Bacterial community structures changed below 20 μm at the scale where microbes operate. Strikingly, only members of two bacterial and one fungal phyla (Proteobacteria, Bacteroidota and Ascomycota, respectively) were washed-off the soil during fractionation and accumulated in the supernatant fraction where most of the detected bacterial genera (e.g., Pseudomonas, Massilia, Mucilaginibacter, Edaphobaculum, Duganella, Janthinobacterium and Variovorax) were previously associated with exopolysaccharide production and biofilm formation.Overall, the applied method shows potential to study soil microbial communities at micro scales which might be useful in studies focusing on the role of specific fungal taxa in soil structure formation as well as research on how and by whom biofilm-like structures are distributed and organized in soil.ImportanceIntensive exploitation of soils has led to increasing environmental concerns such as pollution, erosion, emission of greenhouse gases and, in general, the weakening of its ecosystem services that are mainly regulated by microbial activity. Microbial activity and metabolism drive the formation of soil aggregates, ranging in size from a few micrometres to several millimetres. Understanding biological mechanisms related to aggregate size classes can provide insight into large-scale processes, but most research has focused on macroaggregates. Here, we investigated the microbial community and its functional changes at these smaller scales that are clearly more relevant for assessing microbial activity. We demonstrated that fungal communities are more sensitive to bigger size classes than bacteria, suggesting their dominant role in soil structure formation and turnover. We also identified preferential niches for reductive processes within the nitrogen cycle and a selection of specific taxa by analysing the water used for the wet-fractionation approach.

Primary biological aerosol particles and microorganisms are ubiquitous in the atmosphere. Investigations of airborne chemical markers and microbial communities are critical for identifying sources, transport and transformation processes of aerosols. One potential major source of airborne chemical compounds and microbial communities (e.g. L- and D-amino acids, Flavobacteria) could be related to phytoplankton blooms that occur during the spring season in Arctic fjord systems. Here, we conducted a field study in a polar environment to investigate the occurrence in coarse and fine particles of water-soluble compounds (major ions, carboxylic acids and free L- and D-amino acids) and airborne bacterial communities in aerosol samples. The sampling was conducted with a 6 day sampling frequency at the Gruvebadet observatory, close to Ny-Alesund (Svalbard Islands). Glycine, D-amino acids and C4- organic acids increased during the exponential phase of a marine bloom that occurred in Kongsfjorden and started to drop at the beginning of the main-bloom phase. On the other hand, Polaribacter together with free L-amino acids overlapped with the Chlorophyll a peak and the subsequent decline, and thus might constitute a useful marker for the main-bloom phase.

The atmosphere is an important route for transporting and disseminating microorganisms over short and long distances. Understanding how microorganisms are distributed in the atmosphere is critical due to their role in public health, meteorology and atmospheric chemistry. In order to determine the dominant processes that structure airborne microbial communities, we investigated the diversity and abundance of both bacteria and fungi from the PM10 particle size (particulate matter of 10 micrometers or less in diameter) as well as particulate matter chemistry and local meteorological characteristics over time at nine different meteorological stations around the world. The bacterial genera Bacillus and Sphingomonas as well as the fungal species Pseudotaeniolina globaosa and Cladophialophora proteae were the most abundant taxa of the dataset, although their relative abundances varied greatly based on sampling site. Bacterial and fungal concentration was the highest at the high-altitude and semi-arid plateau of Namco (China; 3.56 × 10

International audience; Fungal denitrification is claimed to produce non‐negligible amounts of N2O in soils, but few tested species have shown significant activity. We hypothesized that denitrifying fungi would be found among those with assimilatory nitrate reductase, and tested 20 such batch cultures for their respiratory metabolism, including two positive controls, Fusarium oxysporum and Fusarium lichenicola, throughout the transition from oxic to anoxic conditions in media supplemented with urn:x-wiley:14622912:media:emi14980:emi14980-math-0001. Enzymatic reduction of urn:x-wiley:14622912:media:emi14980:emi14980-math-0002 (NIR) and NO (NOR) was assessed by correcting measured NO‐ and N2O‐kinetics for abiotic NO‐ and N2O‐production (sterile controls). Significant anaerobic respiration was only confirmed for the positive controls and for two of three Fusarium solani cultures. The NO kinetics in six cultures showed NIR but not NOR activity, observed through the accumulation of NO. Others had NOR but not NIR activity, thus reducing abiotically produced NO to N2O. The presence of candidate genes (nirK and p450nor) was confirmed in the positive controls, but not in some of the NO or N2O accumulating cultures. Based on our results, we conclude that only the Fusarium cultures were able to sustain anaerobic respiration and produced low amounts of N2O as a response to an abiotic NO production from the medium.

Lichens are traditionally defined as a symbiosis between a fungus and a green alga and or a cyanobacterium. This idea has been challenged by the discovery of bacterial communities inhabiting the lichen thalli. These bacteria are thought to contribute to the survival of lichens under extreme and changing environmental conditions. How these changing environmental conditions affect the lichen-associated bacterial community composition remains unclear.We describe the total (rDNA-based) and potentially metabolically active (rRNA-based) bacterial community of the lichen Cetaria islandica and its response to long-term warming using a 20-year warming experiment in an Icelandic sub-Arctic tundra. 16S rRNA and rDNA amplicon sequencing showed that the orders Acetobacterales (of the class Alphaproteobacteria) and Acidobacteriales (of the phylum Acidobacteria) dominated the bacterial community. Numerous ASVs (amplicon sequence variants) taxa could only be detected in the potentially active community but not in the total community. Long-term warming led to increases in relative abundance on class, order and ASV level. Warming altered the relative abundance of ASVs of the most common bacterial genera, such as Granulicella and Endobacter. The potentially metabolically active bacterial community was also more responsive to warming than the total community.Our results suggest that the bacterial community of the lichen C. islandica is dominated by acidophilic taxa and harbors disproportionally active rare taxa. We also show for the first time that climate warming can lead to shifts in lichen-associated bacterial community composition.

Primary biological aerosols are transported over large distances, are traveling in various media such as dry air masses, clouds or fog, and eventually deposited with dry deposition, especially for larger particles, or precipitation like rain, hail or snow. To investigate relative abundance and diversity of airborne bacterial and fungal communities, samples have been collected with a liquid impinger (Coriolis μ) from the top of Mount Sonnblick (3106 m asl, Austrian Alps) from the respective sources under a temporal aspect over four seasons over the year to include all climatic conditions. Bacterial and fungal samples (16S rRNA and ITS) were sequenced using Illumina MiSeq paired-end sequencing, investigated for relative abundance by qPCR(16S rRNA and 18S rRNA) and ice nucleation activity. Results show that there is no stable free tropospheric air microbial community and air mass origin was different for the four sampling periods which exerted influence on the microbial composition of the atmosphere although a core microbiome could be identified consisting of 61 bacterial OTUs and eight fungal genera. Differentiation between seasons was stronger pronounced in air than in precipitation, with rain being most different and variable of precipitation types, indicating distinct forces driving microbial fate in the air. Microorganisms precipitated with snow, hail or rain or being transported by clouds differ in their species composition from free tropospheric air masses and do not mirror the air community structure. They were more diverse, distinct in composition, 16S:18S ratio and abundance from free-floating PBA. Hence, snow or cloud samples are not suitable proxies for free tropospheric air microbiome composition, since separation processes in aerosolization, transport and scavenging occur. The microbial composition of arriving precipitation or clouds represents only a part of the microbial air composition communities of the cumulative sources of origin. Relative abundance and composition of ice nucleation-active bacteria showed a higher share of relative % reads of known ice nucleation-active bacteria present in all wet phases compared to air. Results propose a separation of IN-active reads with higher shares occurring in precipitation. This study presents the first comparison of free tropospheric bacterial and fungal abundance and diversity in time

Bacterial communities form the basis of biogeochemical processes and determine plant growth and health. Mosses, an abundant plant group in Arctic ecosystems, harbour diverse bacterial communities that are involved in nitrogen fixation and carbon cycling. Global climate change is causing changes in aboveground plant biomass and shifting species composition in the Arctic, but little is known about the response of moss microbiomes.Here, we studied the total and potentially active bacterial community associated with Racomitrium lanuginosum, in response to 20-year in situ warming in an Icelandic heathland. We evaluated the effect of warming and warming-induced shrub expansion on the moss bacterial community composition and diversity, nifH gene abundance and nitrogen-fixation rates.Warming changed both the total and the potentially active bacterial community structure, while litter abundance only affected the total bacterial community structure. The relative abundance of Proteobacteria increased, while the relative abundance of Cyanobacteria and Acidobacteria decreased. NifH gene abundance and nitrogen-fixation rates were negatively affected by litter and Betula nana abundance, respectively. We also found shifts in the potentially nitrogen-fixing community, with Nostoc decreasing and non-cyanobacterial diazotrophs increasing in relative abundance. Our data suggests that the moss microbial community including the potentially nitrogen-fixing taxa is sensitive to future warming.Synthesis. Long-term warming led to a shift in moss-associated bacterial community composition, while the abundance of nitrogen-fixing bacteria and nitrogen-fixation rates were negatively affected by increased litter and Betula nana abundance respectively. Warming and increased shrub abundance as a result of warming can affect moss-associated bacterial communities and nitrogen fixation rates in tundra ecosystems.

Fungal denitrification is claimed to produce non-negligible amounts of N