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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]

Background: Cyanobacteria are major primary producers in extreme cold ecosystems. Many lineages of cyanobacteria thrive in these harsh environments, but it is not fully understood how they survive in these conditions and whether they have evolved specific mechanisms of cold adaptation. Phormidesmis priestleyi is a cyanobacterium found throughout the cold biosphere (Arctic, Antarctic and alpine habitats). Genome sequencing of P. priestleyi BC1401, an isolate from a cryoconite hole on the Greenland Ice Sheet, has allowed for the examination of genes involved in cold shock response and production of extracellular polymeric substances (EPS). EPSs likely enable cyanobacteria to buffer the effects of extreme cold and by identifying mechanisms for EPS production in P. priestleyi BC1401 this study lays the way for investigating transcription and regulation of EPS production in an ecologically important cold tolerant cyanobacterium. Results: We sequenced the draft genome of P. priestleyi BC1401 and implemented a new de Bruijn graph visualisation approach combined with BLAST analysis to separate cyanobacterial contigs from a simple metagenome generated from non-axenic cultures. Comparison of known cold adaptation genes in P. priestleyi BC1401 with three relatives from other environments revealed no clear differences between lineages. Genes involved in EPS biosynthesis were identified from the Wzy- and ABC-dependent pathways. The numbers of genes involved in cell wall and membrane biogenesis in P. priestleyi BC1401 were typical relative to the genome size. A gene cluster implicated in biofilm formation was found homologous to the Wps system, although the intracellular signalling pathways by which this could be regulated remain unclear. Conclusions: Results show that the genomic characteristics and complement of known cold shock genes in P. priestleyi BC1401 are comparable to related lineages from a wide variety of habitats, although as yet uncharacterised cold shock genes in this organism may still exist. EPS production by P. priestleyi BC1401 likely contributes to its ability to survive efficiently in cold environments, yet this mechanism is widely distributed throughout the cyanobacterial phylum. Discovering how these EPS related mechanisms are regulated may help explain why P. priestleyi BC1401 is so successful in cold environments where related lineages are not. [ABSTRACT FROM AUTHOR]

Microbial communities in glacial ecosystems are diverse, active, and subjected to strong viral pressures and infection rates. In this study we analyse putative virus genomes assembled from three dsDNA viromes from cryoconite hole ecosystems of Svalbard and the Greenland Ice Sheet to assess the potential hosts and functional role viruses play in these habitats. We assembled 208 million reads from the virus-size fraction and developed a procedure to select genuine virus scaffolds from cellular contamination. Our curated virus library contained 546 scaffolds up to 230 Kb in length, 54 of which were circular virus consensus genomes. Analysis of virus marker genes revealed a wide range of viruses had been assembled, including bacteriophages, cyanophages, nucleocytoplasmic large DNA viruses and a virophage, with putative hosts identified as Cyanobacteria, Alphaproteobacteria, Gammaproteobacteria, Actinobacteria, Firmicutes, eukaryotic algae and amoebae. Whole genome comparisons revealed the majority of circular genome scaffolds (CGS) formed 12 novel groups, two of which contained multiple phage members with plasmid-like properties, including a group of phage-plasmids possessing plasmid-like partition genes and toxin-antitoxin addiction modules to ensure their replication and a satellite phage-plasmid group. Surprisingly we also assembled a phage that not only encoded plasmid partition genes, but a clustered regularly interspaced short palindromic repeat (CRISPR)/Cas adaptive bacterial immune system. One of the spacers was an exact match for another phage in our virome, indicating that in a novel use of the system, the lysogen was potentially capable of conferring immunity on its bacterial host against other phage. Together these results suggest that highly novel and diverse groups of viruses are present in glacial environments, some of which utilize very unusual life strategies and genes to control their replication and maintain a long-term relationship with their hosts. [ABSTRACT FROM AUTHOR]

The seasonal melting of ice entombed cryoconite holes on McMurdo Dry Valley glaciers provides oases for life in the harsh environmental conditions of the polar desert where surface air temperatures only occasionally exceed 0°C during the Austral summer. Here we follow temporal changes in cryoconite hole biogeochemistry on Canada Glacier from fully frozen conditions through the initial stages of spring thaw toward fully melted holes. The cryoconite holes had a mean isolation age from the glacial drainage system of 3.4 years, with an increasing mass of aqueous nutrients (dissolved organic carbon, total nitrogen, total phosphorus) with longer isolation age. During the initial melt there was a mean nine times enrichment in dissolved chloride relative to mean concentrations of the initial frozen holes indicative of an ionic pulse, with similar mean nine times enrichments in nitrite, ammonium, and dissolved organic matter. Nitrate was enriched twelve times and dissolved organic nitrogen six times, suggesting net nitrification, while lower enrichments for dissolved organic phosphorus and phosphate were consistent with net microbial phosphorus uptake. Rates of bacterial production were significantly elevated during the ionic pulse, likely due to the increased nutrient availability. There was no concomitant increase in photosynthesis rates, with a net depletion of dissolved inorganic carbon suggesting inorganic carbon limitation. Potential nitrogen fixation was detected in fully melted holes where it could be an important source of nitrogen to support microbial growth, but not during the ionic pulse where nitrogen availability was higher. This study demonstrates that ionic pulses significantly alter the timing and magnitude of microbial activity within entombed cryoconite holes, and adds credence to hypotheses that ionic enrichments during freeze-thaw can elevate rates of microbial growth and activity in other icy habitats, such as ice veins and subglacial regelation zones. [ABSTRACT FROM AUTHOR]

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]

Abstract: Cryoconite holes on glacier surfaces are ice-cold hot spots of microbial diversity and activity but still little is known about their fungal inhabitants. We provide the first report of distinctive fungal communities in cryoconite debris from three valley glaciers at Kongsfjorden, Svalbard. Multivariate analysis of terminal-restriction fragment length polymorphism (T-RFLP) profiles of rRNA ITS amplicons revealed that quite distinct fungal communities were found in cryoconite holes compared with soils from adjacent moraine and tundra sites, and that communities on glaciers with contrasting ice-surface hydrology also differed. Most of the fungi cultured from cryoconite sediment were basidiomycetous yeasts or filamentous Ascomycota (Helotiales/Pleosporales). The latter included aeroaquatic fungi, such as Articulospora and Varicosporium, implying a role for these important freshwater decomposers in the carbon dynamics of cryoconite holes. Matching of the dominant peaks from T-RFLP analysis to predicted peaks of cultured isolates confirmed the abundance of these aeroaquatic fungi but also revealed that most of the dominant T-RFLP peaks did not match any cultured isolates. Considering the prevalence and endangerment of glacial environments worldwide, these findings would suggest that their potential as reservoirs of fungal diversity should not be overlooked. [Copyright &y& Elsevier]

The diversity of highly active bacterial communities in cryoconite holes on three Arctic glaciers in Svalbard was investigated using terminal restriction fragment length polymorphism (T-RFLP) of the 16S rRNA locus. Construction and sequencing of clone libraries allowed several members of these communities to be identified, with Proteobacteria being the dominant one, followed by Cyanobacteria and Bacteroidetes. T-RFLP data revealed significantly different communities in holes on the (cold) valley glacier Austre Brøggerbreen relative to two adjacent (polythermal) valley glaciers, Midtre Lovénbreen and Vestre Brøggerbreen. These population compositions correlate with differences in organic matter content, temperature and the metabolic activity of microbial communities concerned. No within-glacier spatial patterns were observed in the communities identified over the 2-year period and with the 1 km-spaced sampling. We infer that surface hydrology is an important factor in the development of cryoconite bacterial communities. [ABSTRACT FROM AUTHOR]

Cryoconite holes, which can cover 0.1–10% of the surface area of glaciers, are small, water-filled depressions (typically <1 m in diameter and usually <0.5 m deep) that form on the surface of glaciers when solar-heated inorganic and organic debris melts into the ice. Recent studies show that cryoconites are colonized by a diverse range of microorganisms, including viruses, bacteria and algae. Whether microbial communities on the surface of glaciers are actively influencing biogeochemical cycles or are just present in a dormant state has been a matter of debate for long time. Here, we report primary production and community respiration of cryoconite holes upon glaciers in Svalbard, Greenland and the European Alps. Microbial activity in cryoconite holes is high despite maximum temperatures seldom exceeding 0.1 °C. In situ primary production and respiration in cryoconites during the summer is often comparable with that found in soils in warmer and nutrient richer regions. Considering only glacier areas outside Antarctica and a conservative average cryoconite distribution on glacial surfaces, we found that on a global basis cryoconite holes have the potential to fix as much as 64 Gg of carbon per year (i.e. 98 Gg of photosynthesis minus 34 Gg of community respiration). Most lakes and rivers are generally considered as heterotrophic systems, but our results suggest that glaciers, which contain 75% of the freshwater of the planet, are largely autotrophic systems. [ABSTRACT FROM AUTHOR]

There is now compelling evidence that microbially mediated reactions impart a significant effect upon the dynamics, composition, and abundance of nutrients in glacial melt water. Consequently, we must now consider ice masses as ecosystem habitats in their own right and address their diversity, functional potential, and activity as part of alpine and polar environments. Although such research is already underway, its fragmentary nature provides little basis for developing modern concepts of glacier ecology. This paper therefore provides a much-needed framework for development by reviewing the physical, biogeochemical, and microbiological characteristics of microbial habitats that have been identified within glaciers and ice sheets. Two key glacial ecosystems emerge, one inhabiting the glacier surface (the supraglacial ecosystem) and one at the ice-bed interface (the subglacial ecosystem). The supraglacial ecosystem is characterized by a diverse consortium of microbes (usually bacteria, algae, phytoflagellates, fungi, viruses and occasional rotifers, tardigrades, and diatoms) within the snowpack, supraglacial streams, and melt pools (cryoconite holes). The subglacial system is dominated by aerobic/anaerobic bacteria and most probably viruses in basal ice/till mixtures and subglacial lakes. A third, so-called englacial ecosystem is also described, but it is demonstrated that conditions within glacier ice are sufficient to make metabolic activity and its impact upon nutrient dynamics negligible at the glacier scale. Consideration of the surface and internal heat balances of the glacier show that all glacial ecosystems are sensitive to climate change, although at different timescales. Thus, while rapid, melt-driven habitat changes lead to melt-out, resuscitation, and redistribution of microorganisms in many supraglacial ecosystems, much slower climatic and glacial mass-balance processes effect such changes in the subglacial ecosystem. Paradoxically, it is shown that these forces have brought about net refreezing and the onset of cryostasis in the subglacial ecosystems of many Arctic glaciers subject to thinning in recent decades. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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