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Cryoconite holes are known as foci of microbial diversity and activity on polar glacier surfaces, but are virtually unexplored microbial habitats in alpine regions. In addition, whether cryoconite community structure reflects ecosystem functionality is poorly understood. Terminal restriction fragment length polymorphism and Fourier transform infrared metabolite fingerprinting of cryoconite from glaciers in Austria, Greenland and Svalbard demonstrated cryoconite bacterial communities are closely correlated with cognate metabolite fingerprints. The influence of bacterial-associated fatty acids and polysaccharides was inferred, underlining the importance of bacterial community structure in the properties of cryoconite. Thus, combined application of T-RFLP and FT- IR metabolite fingerprinting promises high throughput, and hence, rapid assessment of community structure-function relationships. Pyrosequencing revealed Proteobacteria were particularly abundant, with Cyanobacteria likely acting as ecosystem engineers in both alpine and Arctic cryoconite communities. However, despite these generalities, significant differences in bacterial community structures, compositions and metabolomes are found between alpine and Arctic cryoconite habitats, reflecting the impact of local and regional conditions on the challenges of thriving in glacial ecosystems. [ABSTRACT FROM AUTHOR]

Subglacial environments are largely anoxic, contain organic carbon ( OC) overridden by glacier ice during periods of advance, and harbour active microbial communities. This creates favourable conditions for OC degradation via methanogenesis. It has been hypothesized that OC beneath ice sheets is converted to methane ( CH4) and may be released to the atmosphere during retreat. However, there are limited data available to support this contention. Here, we present new data on the abundance, diversity and activity of methanogenic archaea and the amount and character of OC in subglacial sediments from Arctic and Antarctic glacial systems based on different substrate types. We employed long-term laboratory incubations to quantify the CH4 production potential in different subglacial settings. Significant numbers of methanogens (up to 7 × 104 cells g−1) were detected in the samples and clones of Methanomicrobiales and Methanosarcinales were identified in clone libraries. Long lag periods (up to >200 days) were observed before significant CH4 concentrations were measured. We report order of magnitude differences in rates of CH4 production (101-105 fmol g−1 d−1) in different subglacial sediments, reflecting contrasts in the origin of the sediment and the OC character. Hence, we predict that contrasting rates of CH4 production are likely to occur beneath glaciers and ice sheets that overran different types of substrate. We subsequently estimated the potential for CH4 production beneath the Laurentide/Inuitian/Cordilleran and Fennoscandian Ice Sheets during a typical 85 ka Quaternary glacial/interglacial cycle. CH4 production from lacustrine-derived OC is likely to be an order of magnitude higher (~6.3-27 Pg C) than that from overridden soils (~0.55-0.68 Pg C), possibly due to a difference in lability between lacustrine and soil OC. While representing a fraction of the entire OC pool (~418-610 Pg C), this finding highlights the importance of considering the character of different OC pools when calculating subglacial CH4 production. [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.

In recent years, genomic analyses have arisen as an exciting way of investigating the functional capacity and environmental adaptations of numerous micro-organisms of global relevance, including cyanobacteria. In the extreme cold of Arctic, Antarctic and alpine environments, cyanobacteria are of fundamental ecological importance as primary producers and ecosystem engineers. While their role in biogeochemical cycles is well appreciated, little is known about the genomic makeup of polar and alpine cyanobacteria. In this article, we present ways that genomic techniques might be used to further our understanding of cyanobacteria in cold environments in terms of their evolution and ecology. Existing examples from other environments (e.g. marine/hot springs) are used to discuss how methods developed there might be used to investigate specific questions in the cryosphere. Phylogenomics, comparative genomics and population genomics are identified as methods for understanding the evolution and biogeography of polar and alpine cyanobacteria. Transcriptomics will allow us to investigate gene expression under extreme environmental conditions, and metagenomics can be used to complement tradition amplicon-based methods of community profiling. Finally, new techniques such as single cell genomics and metagenome assembled genomes will also help to expand our understanding of polar and alpine cyanobacteria that cannot readily be cultured.

Tailed bacteriophages are the most abundant viruses in the biosphere. Here we examined the T4-type bacteriophage community inhabiting the surface of two glaciers in Svalbard. We used a molecular approach to target g23, the major capsid protein gene, to demonstrate that in the extreme cryoconite hole habitats the T4-type phages are surprisingly diverse. Phylogenetic analysis revealed that cryoconite hole sediments harbour a mixed phage community spanning multiple T4-type phage subgroups. The majority (71 %) of phage sequences clustered into three novel phylogenetically distinct groups, whilst the remainder clustered with known marine and soil derived phage sequences. The meltwater in cryoconite holes also contained a further distinct phage community which was related to previously detected marine phage variants. The ability of phages to move between marine and glacial habitats was tested in a transplantation experiment. Phages from the nearby marine fjord were found to be capable of initiating infection of supraglacial bacteria, suggesting suitable hosts could be found by non-native phages. Together this evidence suggests that the surface of glaciers contain both novel and cosmopolitan phages, some of which may have arrived in the cryosphere from other biomes.

The tundra is the coldest biome described in typical geography and biology textbooks. Within the cryosphere, there are large expanses of ice in the Antarctic, Arctic and alpine regions that are not regarded as being part of any biome. During the summer, there is significant melt on the surface of glaciers, ice caps and ice shelves, at which point microbial communities become active and play an important role in the cycling of carbon and other elements within the cryosphere. In this review, we suggest that it is time to recognise the cryosphere as one of the biomes of Earth. The cryospheric biome encompasses extreme environments and is typified by truncated food webs dominated by viruses, bacteria, protozoa and algae with distinct biogeographical structures., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Bacteriophages are found wherever microbial life is present and play a significant role in aquatic ecosystems. They mediate microbial abundance, production, respiration, diversity, genetic transfer, nutrient cycling and particle size distribution. Most studies of bacteriophage ecology have been undertaken at temperate latitudes. Data on bacteriophages in polar inland waters are scant but the indications are that they play an active and dynamic role in these microbially dominated polar ecosystems. This review summarises what is presently known about polar inland bacteriophages, ranging from subglacial Antarctic lakes to glacial ecosystems in the Arctic. The review examines interactions between bacteriophages and their hosts and the abiotic and biotic variables that influence these interactions in polar inland waters. In addition, we consider the proportion of the bacteria in Arctic and Antarctic lake and glacial waters that are lysogenic and visibly infected with viruses. We assess the relevance of bacteriophages in the microbial loop in the extreme environments of Antarctic and Arctic inland waters with an emphasis on carbon cycling.

Bacterial production in glacial runoff and aquatic habitats along a c. 500 m transect from the ablation area of a Svalbard glacier (Midre Lovénbreen, 79 degrees N, 12 degrees E) down to a series of proglacial lakes in its forefield were assessed. In addition, a series of in situ experiments were conducted to test how different nutrient sources (glacial flour and dissolved organic matter derived from goose faeces) and temperature affect bacterial abundance and production in these ecosystems. Bacterial abundance and production increased significantly along this transect and reached a maximum in the proglacial lakes. Bacterial diversity profiles as assessed by denaturing gradient gel electrophoresis indicated that communities in glacial runoff were different from those in proglacial lakes. Heterotrophic bacterial production was mainly controlled by temperature and phosphorus limitation. Addition of both glacial flour and dissolved organic matter derived from goose faeces stimulated bacterial production in those lakes. The results suggest that glacial runoff sustains an active bacterial community which is further stimulated in proglacial lakes by higher temperatures and nutrient inputs from bird faeces. Thus, as in maritime temperate and Antarctic settings, bacterial communities developing in the recently deglaciated terrain of Svalbard receive important inputs of nutrients via faunal transfers from adjacent ecosystems.

The frequency of visibly phage-infected bacterial cells (FVIB) and the average number of phages per cell [i.e. burst size (BS)] were determined in Antarctic and Arctic ultra-oligotrophic freshwater environments. Water samples were collected from two Antarctic freshwater lakes and cryoconite holes from a glacier in the Arctic. Data from this bipolar study show the highest FVIB (average 26.1%, range 5.1% to 66.7%) and the lowest BS (average 4, range 2-15) ever reported in the literature. The bacterial density is low in these ultra-oligotrophic freshwater environments but a large proportion of the bacteria are visibly infected. Our results suggest that a constant virioplankton population can be maintained in these extreme environments even though host density is low and often slow growing.