Fate of glacier surface snow-originating bacteria in the glacier-fed hydrologic continuums.

Glaciers represent important biomes of Earth and are recognized as key species pools for downstream aquatic environments. Worldwide, rapidly receding glaciers are driving shifts in hydrology, species distributions and threatening microbial diversity in glacier-fed aquatic ecosystems. However, the impact of glacier surface snow-originating taxa on the microbial diversity in downstream aquatic environments has been little explored. To elucidate the contribution of glacier surface snow-originating taxa to bacterial diversity in downstream aquatic environments, we collected samples from glacier surface snows, downstream streams and lakes along three glacier-fed hydrologic continuums on the Tibetan Plateau. Our results showed that glacier stream acts as recipients and vectors of bacteria originating from the glacier environments. The contributions of glacier surface snow-originating taxa to downstream bacterial communities decrease from the streams to lakes, which was consistently observed in three geographically separated glacier-fed ecosystems. Our results also revealed that some rare snow-originating bacteria can thrive along the hydrologic continuums and become dominant in downstream habitats. Finally, our results indicated that the dispersal patterns of bacterial communities are largely determined by mass effects and increasingly subjected to local sorting of species along the glacier-fed hydrologic continuums. Collectively, this study provides insights into the fate of bacterial assemblages in glacier surface snow following snow melt and how bacterial communities in aquatic environments are affected by the influx of glacier snow-originating bacteria.

Sulfuriferula nivalis sp. nov., a sulfur oxidizer isolated from snow and emended description of Sulfuriferula plumbiphila.

A chemolithoautotrophic sulfur-oxidizing bacterium, strain SGTMT was isolated from snow collected in Japan. As electron donors for growth, SGTMT oxidized thiosulfate, tetrathionate and elemental sulfur. Heterotrophic growth was not observed. Growth of the novel isolate was observed at a temperature range of 5-28 °C, with optimum growth at 18 °C. SGTMT grew at a pH range of 4.3-7.4, with optimum growth at pH 6.1-7.1. Major components in the cellular fatty acid profile were summed feature 3 (C16 : 1ω7c and/or C16 : 1ω6c) and C16 : 0. The complete genome of SGTMT consisted of a circular chromosome of approximately 3.4 Mbp and two plasmids. Phylogenetic analysis based on the 16S rRNA gene indicated that SGTMT represented a member of the genus Sulfuriferula, and its closest relative is Sulfuriferula thiophila mst6T with a sequence identity of 98 %. A comparative genome analysis showed dissimilarity between the genomes of SGTMT and S. thiophila mst6T, as low values of average nucleotide identity (74.9 %) and digital DNA-DNA hybridization (20.4%). On the basis of its genomic and phenotypic properties, SGTMT (=DSM 109609T=BCRC 81185T) is proposed as the type strain of a novel species, Sulfuriferula nivalis sp. nov. Some characteristics of another species in the same genus, Sulfuriferula plumbiphila, were also investigated to revise and supplement its description. The type strain of S. plumbiphila can grow on thiosulfate, tetrathionate and elemental sulfur. The strain showed optimum growth at pH 6.3-7.0 and shared major cellular fatty acids with the other species of the genus Sulfuriferula.

Microbial Ecology of Snow Reveals Taxa-Specific Biogeographical Structure.

Snows that persist late into the growing season become colonized with numerous metabolically active microorganisms, yet underlying mechanisms of community assembly and dispersal remain poorly known. We investigated (Illumina MiSeq) snow-borne bacterial, fungal, and algal communities across a latitudinal gradient in Fennoscandia and inter-continental distribution between northern Europe and North America. Our data indicate that bacterial communities are ubiquitous regionally (across Fennoscandia), whereas fungal communities are regionally heterogeneous. Both fungi and bacteria are biogeographically heterogeneous inter-continentally. Snow algae, generally thought to occur in colorful algae blooms (red, green, or yellow) on the snow surface, are molecularly described here as an important component of snows even in absence of visible algal growth. This suggests that snow algae are a previously underestimated major biological component of visually uncolonized snows. In contrast to fungi and bacteria, algae exhibit no discernible inter-continental or regional community structure and exhibit little endemism. These results indicate that global and regional snow microbial communities and their distributions may be dictated by a combination of size-limited propagule dispersal potential and restrictions (bacteria and fungi) and homogenization of ecologically specialized taxa (snow algae) across the globe. These results are among the first to compare inter-continental snow microbial communities and highlight how poorly understood microbial communities in these threatened ephemeral ecosystems are.

Bioavailability of Mineral-Bound Iron to a Snow Algal-Bacterial Coculture and Implications for Albedo-Altering Snow Algal Blooms.

Snow algae can form large-scale blooms across the snowpack surface and near-surface environments. These pigmented blooms can decrease snow albedo and increase local melt rates, and they may impact the global heat budget and water cycle. Yet, the underlying causes for the geospatial occurrence of these blooms remain unconstrained. One possible factor contributing to snow algal blooms is the presence of mineral dust as a micronutrient source. We investigated the bioavailability of iron (Fe)-bearing minerals, including forsterite (Fo90, Mg1.8Fe0.2SiO4), goethite, smectite, and pyrite as Fe sources for a Chloromonas brevispina-bacterial coculture through laboratory-based experimentation. Fo90 was capable of stimulating snow algal growth and increased the algal growth rate in otherwise Fe-depleted cocultures. Fo90-bearing systems also exhibited a decrease in the ratio of bacteria to algae compared to those of Fe-depleted conditions, suggesting a shift in microbial community structure. The C. brevispina coculture also increased the rate of Fo90 dissolution relative to that of an abiotic control. Analysis of 16S rRNA genes in the coculture identified Gammaproteobacteria, Betaproteobacteria, and Sphingobacteria, all of which are commonly found in snow and ice environments. Archaea were not detected. Collimonas and Pseudomonas, which are known to enhance mineral weathering rates, comprised two of the top eight (>1%) operational taxonomic units (OTUs). These data provide unequivocal evidence that mineral dust can support elevated snow algal growth under otherwise Fe-depleted growth conditions and that snow algal microbial communities can enhance mineral dissolution under these conditions.IMPORTANCE Fe, a key micronutrient for photosynthetic growth, is necessary to support the formation of high-density snow algal blooms. The laboratory experiments described herein allow for a systematic investigation of the interactions of snow algae, bacteria, and minerals and their ability to mobilize and uptake mineral-bound Fe. Results provide unequivocal and comprehensive evidence that mineral-bound Fe in Fe-bearing Fo90 was bioavailable to Chloromonas brevispina snow algae within an algal-bacterial coculture. This evidence includes (i) an observed increase in snow algal density and growth rate, (ii) decreased ratios of bacteria to algae in Fo90-containing cultures relative to those of cultures grown under similarly Fe-depleted conditions with no mineral-bound Fe present, and (iii) increased Fo90 dissolution rates in the presence of algal-bacterial cocultures relative to those of abiotic mineral controls. These results have important implications for the role of mineral dust in supplying micronutrients to the snow microbiome, which may help support dense snow algal blooms capable of lowering snow albedo and increasing snow melt rates on regional, and possibly global, scales.

Snow angels - the microbiology of freshly fallen snow: implications for immunocompromised patients.

The frequency of seasonal snowfall results in the transient covering of gardens/amenity sites/open public spaces, which encourages recreational interaction mainly with children. No data is available demonstrating the microbiological composition of such fallen snow and therefore a study was undertaken to examine the microbiology of snow from 37 sites, estimating (i) total viable count (TVC), (ii) identification of bacteria, and (iii) the presence of Pseudomonas aeruginosa. Mean TVC count of 8.3 colony-forming units (cfu)/ml snow melt water, 51.7 cfu/ml, 865 cfu/ml and 2,197 cfu/ml, was obtained for public amenity sites, domestic gardens, public open spaces and melting snow from public footpaths, respectively. No bacterial organisms (<10 cfu/ml) were detected in 5/14 (35.7%) open public spaces, 2/5 (40%) amenity sites and in 1/10 (10%) domestic gardens. Pseudomonas aeruginosa was not detected from any snow sample examined. Bacterial diversity consisted of 15 bacterial species (11 Gram-positive/four Gram-negative). The six Gram-positive genera identified from snow were Actinomyces, Bacillus, Brevibacillus, Micrococcus, Staphylococcus and Streptococcus. The four Gram-negative genera identified were Enterobacter, Pantoea, Pseudomonas and Xanthomonas. Bacillus licheniformis was the most commonly isolated organism from snow; it was isolated from every snow type. Snow may contain a diverse range of bacteria, many of which are capable of causing human infections.

Linking microbial diversity and functionality of arctic glacial surface habitats.

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 for the first time we have 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 12 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. It has been shown 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.

Muricauda marina sp. nov., isolated from marine snow of Yellow Sea.

A Gram-staining-negative, long rod-shaped, non-motile, strictly aerobic bacterial strain, designated H19-56T, was isolated from a surface marine snow sample collected from the Yellow Sea near China. Phylogenetic analysis based on 16S rRNA gene sequences indicated that H19-56T represented a member of the genus Muricauda and showed the highest sequence similarity to Muricauda ruestringensis B1T (96.9 %). H19-56T grew optimally at pH 8.0, 32 °C and in the presence of 3 % (w/v) NaCl. The DNA G+C content was 43.6 mol%. H19-56T contained MK-6 as the predominant respiratory quinone and had iso-C15 : 0 and iso-C15 : 1G as the major cellular fatty acids. The major polar lipids were phosphatidylethanolamine, two unidentified aminolipids and three unidentified lipids. On the basis of the results of the polyphasic analyses, this isolate was considered to represent a novel species of the genus Muricauda, for which the name Muricauda marina sp. nov. is proposed. The type strain is H19-56T (CGMCC 1.15774T=JCM 31456T=MCCC 1K03196T=KCTC 52374T).

Description of Conyzicola nivalis sp. nov., isolated from glacial snow, and emended description of the genus Conyzicola and Conyzicola lurida.

A cold-tolerant, translucent, yellow-pigmented, Gram-stain-positive, non-motile, rod-shaped bacteria was isolated from snow of the Zadang Glacier on the Tibetan Plateau, PR China. 16S rRNA gene sequence similarity analysis indicated that the isolate was closely related to Conyzicola lurida KCTC 29231T and Leifsonia psychrotolerans DSM 22824T at a level of 97.72 and 97.49 %, respectively. Other close relatives had a 16S rRNA gene sequence similarity of less than 97 %. The major cell-wall amino acid was 2,4-diaminobutyric acid. The cell-wall sugars were rhamnose, ribose, glucose, galactose and mannose. The major menaquinones were MK-9, MK-10 and MK-11. The major polar lipids were diphosphatidylglycerol and phosphatidylglycerol. The predominant fatty acids were anteiso-C15 : 0 and iso-C16 : 0. In DNA-DNA hybridization tests, strain ZD5-4T shared 43.6 and 34.2 % relatedness with C. lurida KCTC 29231T and L. psychrotolerans DSM 22824T, respectively. Based on phenotypic and chemotaxonomic characteristics, strain ZD5-4T was considered to represent a novel species of the genus Conyzicola, for which the name Conyzicola nivalis sp. nov. is proposed. The type strain is ZD5-4T (=JCM 30076T=CGMCC 1.12813T). Emended descriptions of the genus Conyzicola and Conyzicola lurida are also provided.

Hydrophobic ice-binding sites confer hyperactivity of an antifreeze protein from a snow mold fungus.

Snow mold fungus, Typhula ishikariensis, secretes seven antifreeze protein isoforms (denoted TisAFPs) that assist in the survival of the mold under snow cover. Here, the X-ray crystal structure of a hyperactive isoform, TisAFP8, at 1.0 Å resolution is presented. TisAFP8 folds into a right-handed ß-helix accompanied with a long α-helix insertion. TisAFP8 exhibited significantly high antifreeze activity that is comparable with other hyperactive AFPs, despite its close structural and sequence similarity with the moderately active isoform TisAFP6. A series of mutations introduced into the putative ice-binding sites (IBSs) in the ß-sheet and adjacent loop region reduced antifreeze activity. A double-mutant A20T/A212S, which comprises a hydrophobic patch between the ß-sheet and loop region, caused the greatest depression of antifreeze activity of 75%, when compared with that of the wild-type protein. This shows that the loop region is involved in ice binding and hydrophobic residues play crucial functional roles. Additionally, bound waters around the ß-sheet and loop region IBSs were organized into an ice-like network and can be divided into two groups that appear to mediate separately TisAFP and ice. The docking model of TisAFP8 with the basal plane via its loop region IBS reveals a better shape complementarity than that of TisAFP6. In conclusion, we present new insights into the ice-binding mechanism of TisAFP8 by showing that a higher hydrophobicity and better shape complementarity of its IBSs, especially the loop region, may render TisAFP8 hyperactive to ice binding.

Survival and Antibiotic Resistance of Bacteria in Artificial Snow Produced from Contaminated Water.

This study shows microbiological contamination of water in two main Podhale rivers, whose resources are used for the production of artificial snow, and the resulting snow contamination. Thirty-one E. coli strains were isolated from snow at two ski stations in the studied region, their antimicrobial resistance was determined, and the presence of extended spectrum ß-lactamase (ESBL) genes was searched for. The results indicate that the waters of both rivers are severely contaminated, resulting in the contamination of artificial snow with, among others, thermotolerant E. coli. E. coli isolated from snow were most frequently resistant to ampicillin (74.19%) and amoxicillin/clavulanate (51.61% isolates). Aminoglycosides and third generation cephalosporins were most efficient among the tested antimicrobials. Some bacterial strains were multidrug resistant and three strains exhibited the ESBL mechanism. Molecular analyses showed the presence of ESBL genes in the same three strains. Genetic variation among E. coli indicates that only some genotypes are able to survive the artificial snow production process.

Schwertmannite formation at cell junctions by a new filament-forming Fe(II)-oxidizing isolate affiliated with the novel genus Acidithrix.

A new acidophilic iron-oxidizing strain (C25) belonging to the novel genus Acidithrix was isolated from pelagic iron-rich aggregates ('iron snow') collected below the redoxcline of an acidic lignite mine lake. Strain C25 catalysed the oxidation of ferrous iron [Fe(II)] under oxic conditions at 25 °C at a rate of 3.8 mM Fe(II) day(-1) in synthetic medium and 3.0 mM Fe(II) day(-1) in sterilized lake water in the presence of yeast extract, producing the rust-coloured, poorly crystalline mineral schwertmannite [Fe(III) oxyhydroxylsulfate]. During growth, rod-shaped cells of strain C25 formed long filaments, and then aggregated and degraded into shorter fragments, building large cell-mineral aggregates in the late stationary phase. Scanning electron microscopy analysis of cells during the early growth phase revealed that Fe(III)-minerals were formed as single needles on the cell surface, whereas the typical pincushion-like schwertmannite was observed during later growth phases at junctions between the cells, leaving major parts of the cell not encrusted. This directed mechanism of biomineralization at specific locations on the cell surface has not been reported from other acidophilic iron-oxidizing bacteria. Strain C25 was also capable of reducing Fe(III) under micro-oxic conditions which led to a dissolution of the Fe(III)-minerals. Thus, strain C25 appeared to have ecological relevance for both the formation and transformation of the pelagic iron-rich aggregates at oxic/anoxic transition zones in the acidic lignite mine lake.

Myrmecridium hiemale sp. nov. from snow-covered alpine soil is the first eurypsychrophile in this genus of anamorphic fungi.

Myrmecridium hiemale sp. nov. was isolated from snow-covered alpine bare soil and is described as the first eurypsychrophilic species of this genus of filamentous fungi. Colony growth temperature experiments were carried out in the range 4-37 °C. Morphological characteristics and colony appearance were in accordance with characteristics typical for Myrmecridium, but M. hiemale does not grow at temperatures of 25 °C and above. Sequence analyses of the internal transcribed spacer and LSU rRNA D1/D2 regions indicated that the strain in question represents a distinct taxon within the genus Myrmecridium (Myrmecridiaceae, Sordariomycetes, Ascomycota). The type strain of M. hiemale is CBS 141017T(=JMRC 12083T). A morphological description is provided, and a key is presented for the currently known taxa of Myrmecridium, a group of interesting fungi that are either saprobes or plant endophytes.

Hymenobacter nivis sp. nov., isolated from red snow in Antarctica.

A novel aerobic bacterial strain, P3T, was isolated from a red snow obtained from Antarctica. Cells of strain P3T were rod-shaped, non-motile, catalase-negative, oxidase-positive and Gram-stain-negative. Growth was observed at temperatures ranging from 0 to 25 °C, with optimum growth at 15 °C. The pH range for growth was pH 5.3-7.8. The G+C content of the genomic DNA was 55.0 mol%. The major components in the fatty acid profile were iso-C15 : 0, summed feature 4 (iso-C17 : 1 I and/or anteiso-C17 : 1 B), anteiso-C15 : 0 and summed feature 3 (iso-C16 : 1ω7c and/or iso-C16 : 1ω6c). The predominant isoprenoid quinone was MK-7. Phylogenetic analysis based on the 16S rRNA gene sequence indicated that the novel isolate was a member of the genus Hymenobacter, and the strain showed highest sequence similarity (94 %) with H.ymenpbacter glaciei VUG-A130T, Hymenobacter. soli PB17T and Hymenobacter. antarcticus VUG-A42aaT. On the basis of phylogenetic and phenotypic properties strain P3T represents a novel species of the genus Hymenobacter (for which the name Hymenobacter nivis sp. nov. is proposed. The types strain is P3T (=DSM 101755T=NBRC 111535T).

Diversity and potential sources of microbiota associated with snow on western portions of the Greenland Ice Sheet.

Snow overlays the majority of the Greenland Ice Sheet (GrIS). However, there is very little information available on the microbiological assemblages that are associated with this vast and climate-sensitive landscape. In this study, the structure and diversity of snow microbial assemblages from two regions of the western GrIS ice margin were investigated through the sequencing of small subunit ribosomal RNA genes. The origins of the microbiota were investigated by examining correlations to molecular data obtained from marine, soil, freshwater and atmospheric environments and geochemical analytes measured in the snow. Snow was found to contain a diverse assemblage of bacteria (Alphaproteobacteria, Betaproteobacteria and Gammaproteobacteria) and eukarya (Alveolata, Fungi, Stramenopiles and Chloroplastida). Phylotypes related to archaeal Thaumarchaeota and Euryarchaeota phyla were also identified. The snow microbial assemblages were more similar to communities characterized in soil than to those documented in marine ecosystems. Despite this, the chemical composition of snow samples was consistent with a marine contribution, and strong correlations existed between bacterial beta diversity and the concentration of Na(+) and Cl(-) . These results suggest that surface snow from western regions of Greenland contains exogenous microbiota that were likely aerosolized from more distant soil sources, transported in the atmosphere and co-precipitated with the snow.

Colonization in the photic zone and subsequent changes during sinking determine bacterial community composition in marine snow.

Due to sampling difficulties, little is known about microbial communities associated with sinking marine snow in the twilight zone. A drifting sediment trap was equipped with a viscous cryogel and deployed to collect intact marine snow from depths of 100 and 400 m off Cape Blanc (Mauritania). Marine snow aggregates were fixed and washed in situ to prevent changes in microbial community composition and to enable subsequent analysis using catalyzed reporter deposition fluorescence in situ hybridization (CARD-FISH). The attached microbial communities collected at 100 m were similar to the free-living community at the depth of the fluorescence maximum (20 m) but different from those at other depths (150, 400, 550, and 700 m). Therefore, the attached microbial community seemed to be "inherited" from that at the fluorescence maximum. The attached microbial community structure at 400 m differed from that of the attached community at 100 m and from that of any free-living community at the tested depths, except that collected near the sediment at 700 m. The differences between the particle-associated communities at 400 m and 100 m appeared to be due to internal changes in the attached microbial community rather than de novo colonization, detachment, or grazing during the sinking of marine snow. The new sampling method presented here will facilitate future investigations into the mechanisms that shape the bacterial community within sinking marine snow, leading to better understanding of the mechanisms which regulate biogeochemical cycling of settling organic matter.

Microbial abundance and community structure in a melting alpine snowpack.

Snowmelt is a crucial period for alpine soil ecosystems, as it is related to inputs of nutrients, particulate matter and microorganisms to the underlying soil. Although snow-inhabiting microbial communities represent an important inoculum for soils, they have thus far received little attention. The distribution and structure of these microorganisms in the snowpack may be linked to the physical properties of the snowpack at snowmelt. Snow samples were taken from snow profiles at four sites (1930-2519 m a.s.l.) in the catchment of the Tiefengletscher, Canton Uri, Switzerland. Microbial (Archaea, Bacteria and Fungi) communities were investigated through T-RFLP profiling of the 16S and 18S rRNA genes, respectively. In parallel, we assessed physical and chemical parameters relevant to the understanding of melting processes. Along the snow profiles, density increased with depth due to compaction, while other physico-chemical parameters, such as temperature and concentrations of DOC and soluble ions, remained in the same range (e.g. <2 mg DOC L(-1), 5-30 µg NH4 (+)-N L(-1)) in all samples at all sites. Along the snow profiles, no major change was observed either in cell abundance or in bacterial and fungal diversity. No Archaea could be detected in the snow. Microbial communities, however, differed significantly between sites. Our results show that meltwater rearranges soluble ions and microbial communities in the snowpack.

Soil water flow is a source of the plant pathogen Pseudomonas syringae in subalpine headwaters.

The airborne plant pathogenic bacterium Pseudomonas syringae is ubiquitous in headwaters, snowpack and precipitation where its populations are genetically and phenotypically diverse. Here, we assessed its population dynamics during snowmelt in headwaters of the French Alps. We revealed a continuous and significant transport of P.syringae by these waters in which the population density is correlated with water chemistry. Via in situ observations and laboratory experiments, we validated that P.syringae is effectively transported with the snow melt and rain water infiltrating through the soil of subalpine grasslands, leading to the same range of concentrations as measured in headwaters (10(2) -10(5) CFU l(-1) ). A population structure analysis confirmed the relatedness between populations in percolated water and those above the ground (i.e. rain, leaf litter and snowpack). However, the transport study in porous media suggested that water percolation could have different efficiencies for different strains of P.syringae. Finally, leaching of soil cores incubated for up to 4 months at 8°C showed that indigenous populations of P.syringae were able to survive in subalpine soil under cold temperature. This study brings to light the underestimated role of hydrological processes involved in the long distance dissemination of P.syringae.

The nitrogen cycle in cryoconites: naturally occurring nitrification-denitrification granules on a glacier.

Cryoconites are microbial aggregates commonly found on glacier surfaces where they tend to take spherical, granular forms. While it has been postulated that the microbes in cryoconite granules play an important role in glacier ecosystems, information on their community structure is still limited, and their functions remain unclear. Here, we present evidence for the occurrence of nitrogen cycling in cryoconite granules on a glacier in Central Asia. We detected marker genes for nitrogen fixation, nitrification and denitrification in cryoconite granules by digital polymerase chain reaction (PCR), while digital reverse transcription PCR analysis revealed that only marker genes for nitrification and denitrification were abundantly transcribed. Analysis of isotope ratios also indicated the occurrence of nitrification; nitrate in the meltwater on the glacier surface was of biological origin, while nitrate in the snow was of atmospheric origin. The predominant nitrifiers on this glacier belonged to the order Nitrosomonadales, as suggested by amoA sequences and 16S ribosomal RNA pyrosequencing analysis. Our results suggest that the intense carbon and nitrogen cycles by nitrifiers, denitrifiers and cyanobacteria support abundant and active microbes on the Asian glacier.

Bacterial diversity in snow on North Pole ice floes.

The microbial abundance and diversity in snow on ice floes at three sites near the North Pole was assessed using quantitative PCR and 454 pyrosequencing. Abundance of 16S rRNA genes in the samples ranged between 43 and 248 gene copies per millilitre of melted snow. A total of 291,331 sequences were obtained through 454 pyrosequencing of 16S rRNA genes, resulting in 984 OTUs at 97 % identity. Two sites were dominated by Cyanobacteria (72 and 61 %, respectively), including chloroplasts. The third site differed by consisting of 95 % Proteobacteria. Principal component analysis showed that the three sites clustered together when compared to the underlying environments of sea ice and ocean water. The Shannon indices ranged from 2.226 to 3.758, and the Chao1 indices showed species richness between 293 and 353 for the three samples. The relatively low abundances and diversity found in the samples indicate a lower rate of microbial input to this snow habitat compared to snow in the proximity of terrestrial and anthropogenic sources of microorganisms. The differences in species composition and diversity between the sites show that apparently similar snow habitats contain a large variation in biodiversity, although the differences were smaller than the differences to the underlying environment. The results support the idea that a globally distributed community exists in snow and that the global snow community can in part be attributed to microbial input from the atmosphere.

The ice-binding proteins of a snow alga, Chloromonas brevispina: probable acquisition by horizontal gene transfer.

All ice-and snow-related unicellular algae examined so far secrete ice-binding proteins (IBPs) to mitigate freezing damage. Two types of IBP have been identified in chlorophytes. Type 1 IBPs are members of a large family of proteins that share a large domain of unknown function (DUF3494). Previous studies have suggested that the type 1 algal IBP genes were acquired by horizontal gene transfer. To test this hypothesis I sequenced the IBP genes of a snow alga, Chloromonas brevispina. The IBPs were identified by ice affinity purification, de novo sequencing of a tryptic peptide and large-scale sequencing of the transcriptome and genome. C. brevispina has genes for over 20 IBP isoforms, which strongly indicates their importance. The IBPs are all of type 1 and match fungal and bacterial proteins more closely than they match known algal IBPs, providing further evidence that the genes were acquired by horizontal transfer. Modeling of the 3D structures of the IBPs based on the known structure of a homologous protein suggests that the ice-binding site has characteristics that are shared by all DUF3494 proteins.