Arwyn Edwards

Possible interactions between bacterial diversity, microbial activity and supraglacial hydrology of cryoconite holes in Svalbard

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.

Coupled cryoconite ecosystem structure–function relationships are revealed by comparing bacterial communities in alpine and Arctic glaciers

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.

The dynamic bacterial communities of a melting High Arctic glacier snowpack

Snow environments can occupy over a third of land surface area, but little is known about the dynamics of snowpack bacteria. The effect of snow melt on bacterial community structure and diversity of surface environments of a Svalbard glacier was examined using analyses of 16S rRNA genes via T-RFLP, qPCR and 454 pyrosequencing. Distinct community structures were found in different habitat types, with changes over 1 week apparent, in particular for the dominant bacterial class present, Betaproteobacteria. The differences observed were consistent with influences from depositional mode (snowfall vs aeolian dusts), contrasting snow with dust-rich snow layers and near-surface ice. Contrary to that, slush as the decompositional product of snow harboured distinct lineages of bacteria, further implying post-depositional changes in community structure. Taxa affiliated to the betaproteobacterial genus Polaromonas were particularly dynamic, and evidence for the presence of betaproteobacterial ammonia-oxidizing bacteria was uncovered, inviting the prospect that the dynamic bacterial communities associated with snowpacks may be active in supraglacial nitrogen cycling and capable of rapid responses to changes induced by snowmelt. Furthermore the potential of supraglacial snowpack ecosystems to respond to transient yet spatially extensive melting episodes such as that observed across most of Greenland’s ice sheet in 2012 merits further investigation.

A comparison of the microbiome and the metabolome of different regions of the equine hindgut

The microbiome and associated metabolome of faecal samples were compared to those from the caecum and right dorsal colon of horses and ponies euthanised for nonresearch purposes by investigating the microbial population community structure as well as their functional metabolic products. Through the use of 16S rRNA gene dendrograms, the caecum microbiome was shown to cluster separately from the other gut regions. 16S rRNA gene-based quantitative PCR (q-PCR) also demonstrated differences between the caecum and the other gut regions. Metabolites as identified by Fourier transform infrared clustered in a similar way and specific metabolic products (volatile fatty acids and ammonia) also varied by region. Protozoal 18S rDNA concentration and archaeal mcrA gene concentration quantified by q-PCR were found in higher numbers in the colon than the other gut regions. Diversity calculations using Simpson and Shannon-Wiener indices demonstrated higher diversity in the right dorsal colon and faeces than in the caecum. All findings of this study suggest that faecal samples are likely to represent the microbial population of the right dorsal colon to some extent but not that of the caecum, indicating careful consideration is required when planning microbial investigations of the hindgut of the horse.

A metagenomic snapshot of taxonomic and functional diversity in an alpine glacier cryoconite ecosystem

Cryoconite is a microbe‐mineral aggregate which darkens the ice surface of glaciers. Microbial process and marker gene PCR-dependent measurements reveal active and diverse cryoconite microbial communities on polar glaciers. Here, we provide the first report of a cryoconite metagenome and culture-independent study of alpine cryoconite microbial diversity. We assembled 1.2 Gbp of metagenomic DNA sequenced using an Illumina HiScanSQ from cryoconite holes across the ablation zone of Rotmoosferner in the Austrian Alps. The metagenome revealed a bacterially-dominated community, with Proteobacteria (62% of bacterialassigned contigs) and Bacteroidetes (14%) considerably more abundant than Cyanobacteria (2.5%). Streptophyte DNA dominated the eukaryotic metagenome. Functional genes linked to N, Fe, S and P cycling illustrated an acquisitive trend and a nitrogen cycle based upon efficient ammonia recycling. A comparison of 32 metagenome datasets revealed a similarity in functional profiles between the cryoconite and metagenomes characterized from other cold microbe‐mineral aggregates. Overall, the metagenomic snapshot reveals the cryoconite ecosystem of this alpine glacier as dependent on scavenging carbon and nutrients from allochthonous sources, in particular mosses transported by wind from ice-marginal habitats, consistent with net heterotrophy indicated by productivity measurements. A transition from singular snapshots of cryoconite metagenomes to comparative analyses is advocated.

Cryoconite The dark biological secret of the cryosphere

Cryoconite is granular sediment found on glacier surfaces comprising both mineral and biological material. Despite long having been recognised as an important glaciological and biological phenomenon cryoconite remains relatively poorly understood. Here, we appraise the literature on cryoconite for the first time, with the aim of synthesising and evaluating current knowledge to direct future investigations. We review the properties of cryoconite, the environments in which it is found, the biology and biogeochemistry of cryoconite, and its interactions with climate and anthropogenic pollutants. We generally focus upon cryoconite in the Arctic in summer, with Antarctic and lower latitude settings examined individually. We then compare the current state-of-the-science with that at the turn of the twentieth century, and suggest directions for future research including specific recommendations for studies at a range of spatial scales and a framework for integrating these into a more holistic understanding of cryoconite and its role in the cryosphere.Cryoconite is granular sediment found on glacier surfaces comprising both mineral and biological material. Despite long having been recognised as an important glaciological and biological phenomenon cryoconite remains relatively poorly understood. Here, we appraise the literature on cryoconite for the first time, with the aim of synthesising and evaluating current knowledge to direct future investigations. We review the properties of cryoconite, the environments in which it is found, the biology and biogeochemistry of cryoconite, and its interactions with climate and anthropogenic pollutants. We generally focus upon cryoconite in the Arctic in summer, with Antarctic and lower latitude settings examined individually. We then compare the current state-of-the-science with that at the turn of the twentieth century, and suggest directions for future research including specific recommendations for studies at a range of spatial scales and a framework for integrating these into a more holistic understanding of cr...

Microbial cell budgets of an Arctic glacier surface quantified using flow cytometry

Uncertainty surrounds estimates of microbial cell and organic detritus fluxes from glacier surfaces. Here, we present the first enumeration of biological particles draining from a supraglacial catchment, on Midtre Lovénbreen (Svalbard) over 36 days. A stream cell flux of 1.08 × 10(7)  cells m(-2)  h(-1) was found, with strong inverse, non-linear associations between water discharge and biological particle concentrations. Over the study period, a significant decrease in cell-like particles exhibiting 530 nm autofluorescence was noted. The observed total fluvial export of ~7.5 × 10(14) cells equates to 15.1-72.7 g C, and a large proportion of these cells were small (< 0.5 μm in diameter). Differences between the observed fluvial export and inputs from ice-melt and aeolian deposition were marked: results indicate an apparent storage rate of 8.83 × 10(7)  cells m(-2)  h(-1). Analysis of surface ice cores revealed cell concentrations comparable to previous studies (6 × 10(4)  cells ml(-1)) but, critically, showed no variation with depth in the uppermost 1 m. The physical retention and growth of particulates at glacier surfaces has two implications: to contribute to ice mass thinning through feedbacks altering surface albedo, and to potentially seed recently deglaciated terrain with cells, genes and labile organic matter. This highlights the merit of further study into glacier surface hydraulics and biological processes.

Contrasts between the cryoconite and ice-marginal bacterial communities of Svalbard glaciers

Cryoconite holes are foci of unusually high microbial diversity and activity on glacier surfaces worldwide, comprising melt-holes formed by the darkening of ice by biogenic granular debris. Despite recent studies linking cryoconite microbial community structure to the functionality of cryoconite habitats, little is known of the processes shaping the cryoconite bacterial community. In particular, the assertions that the community is strongly influenced by aeolian transfer of biota from ice-marginal habitats and the potential for cryoconite microbes to inoculate proglacial habitats are poorly quantified despite their longevity in the literature. Therefore, the bacterial community structures of cryoconite holes on three High-Arctic glaciers were compared to bacterial communities in adjacent moraines and tundra using terminal-restriction fragment length polymorphism. Distinct community structures for cryoconite and ice-marginal communities were observed. Only a minority of phylotypes are present in both habitat types, implying that cryoconite habitats comprise distinctive niches for bacterial taxa when compared to ice-marginal habitats. Curiously, phylotype abundance distributions for both cryoconite and ice-marginal sites best fit models relating to succession. Our analyses demonstrate clearly that cryoconites have their own, distinct functional microbial communities despite significant inputs of cells from other habitats.

Microbial diversity on Icelandic glaciers and ice caps.

Algae are important primary colonizers of snow and glacial ice, but hitherto little is known about their ecology on Icelands glaciers and ice caps. Due do the close proximity of active volcanoes delivering large amounts of ash and dust, they are special ecosystems. This study provides the first investigation of the presence and diversity of microbial communities on all major Icelandic glaciers and ice caps over a 3 year period. Using high-throughput sequencing of the small subunit ribosomal RNA genes (16S and 18S), we assessed the snow community structure and complemented these analyses with a comprehensive suite of physical-, geo-, and biochemical characterizations of the aqueous and solid components contained in snow and ice samples. Our data reveal that a limited number of snow algal taxa (Chloromonas polyptera, Raphidonema sempervirens and two uncultured Chlamydomonadaceae) support a rich community comprising of other micro-eukaryotes, bacteria and archaea. Proteobacteria and Bacteroidetes were the dominant bacterial phyla. Archaea were also detected in sites where snow algae dominated and they mainly belong to the Nitrososphaerales, which are known as important ammonia oxidizers. Multivariate analyses indicated no relationships between nutrient data and microbial community structure. However, the aqueous geochemical simulations suggest that the microbial communities were not nutrient limited because of the equilibrium of snow with the nutrient-rich and fast dissolving volcanic ash. Increasing algal secondary carotenoid contents in the last stages of the melt seasons have previously been associated with a decrease in surface albedo, which in turn could potentially have an impact on the melt rates of Icelandic glaciers.

A frozen asset: The potential of flow cytometry in constraining the glacial biome

TO THE EDITOR: Today, around two thirds ( 51M km) of Earth’s freshwater is locked away in glaciers, ice caps and ice sheets (1). Yet, wholesale recognition that these frozen assets constitute Earth’s largest freshwater ecosystem is only recent (2). Considering the global scale of this glacial habitat, and its important role in Earth’s climate system, it should neither be neglected nor overlooked. Here, we evaluate a cytometric definition of this globally significant, icy biome. A sceptical reader more familiar with marine and terrestrial biomes may dismiss the notion of a glacial ecosystem on the grounds that, being “as pure as driven snow,” this icy world is not significantly contaminated by microbes. Yet, long-standing evidence points to microbial habitats in unlikely, cold environments including clouds, precipitation, and snow-cover (3); while cores from the Earth’s two ice sheets reveal the prevalence of immured microbes in glacier ice, with concentrations in the order of 10210 cells/mL in ice of 750 ka in age (4–6). Assuming these microbial abundances are broadly representative, a total of between 4 3 10 and 7 3 10 cells may lie entombed in glacial ice. However, this is a conservative assessment for the glacial biome because the elevated biomass associated with active habitats at the ice surface and glacier bed remains poorly constrained and eukaryote communities are excluded. In comparison, Whitman et al. (7), omitting glacierized environments, reported global estimates for Bacteria and Archaea abundance in all non-glacial freshwater (1.3 3 10 cells), and rainforest, tundra, and alpine soils (Table 1). The comparable magnitude and associated uncertainties of estimates of glacial ice biomass illustrate its potential importance, and motivates continuing investigation. Microbial ecologists have typically preferred to concentrate cells on membrane filters and enumerate cells stained for nucleic acids using epifluorescence microscopy. However, the low concentrations of cells typical of individual samples from snow, glacial ice, or meltwaters, coupled with their presence in an aqueous media, would seem to readily lend flow cytometry (FCM) as an ideal analytical tool for the quantification of cells associated with glacial samples. Nevertheless, despite a much longer history of application in other aquatic habitats (8), and its capability for multi-parameter interrogation of heterogeneous microbial populations at the single-cell level (9), FCM has yet to become an established tool within glacial ecology. Notwithstanding infrequent application, the potential utility of FCM for probing the glacial cryosphere has been demonstrated by the small, but growing, number of glacial and snowpack studies. Initially, FCM enumerations employed Hoechst 33342 to discern microbes in ice from depths of 3 km below the Antarctic ice sheet surface (6). Subsequently, studies applied SYTO stains to enumerate microbes and explore cell size distributions for samples drawn from the GISP2 ice core (5), while measurements of cell concentrations in Tibetan glacier snowpacks (10) have paralleled the perspective that SYBR Green is a more effective nucleic acid stain for freshwater samples. However, these snapshots of communities within glacial ice or snow are not a true reflection of the complexity of community and habitat dynamics that characterize the glacial biome. While both water and cellular mobility in deep ice may be limited (11), the dynamic and porous nature of a glacier’s surface potentially facilitates the translocation of cells and nutrients. The atmosphere–ice interface exhibits complex topography and its hydraulic conductivity evolves as