The role of heterotrophs in glacier surface ecosystem productivity

Cryoconite holes are miniature freshwater ecosystems on glacier surfaces worldwide that harbour a diverse microbial community. Microbial isolates from cryoconite holes show exceptional environmental tolerance: they are able to grow in a wide range of pH, high salinities and scavenge a wide range of organic carbon sources. All the isolates tested were facultative anaerobes The community resilience exceeds the ‘normal’ conditions encountered, and enables their functionality in microenvironments, and in transition environments caused by freeze-thaw. This study demonstrates for the first time that environmental conditions, in particular concentrations of oxygen, are spatially heterogenous at the microscale. Anoxic zones develop rapidly following disturbance and can be found inside cryoconite granules and in thick layers of cryoconite sediment. These microhabitats, with limited oxygen, create microniches for anaerobic metabolism. Facultatively anaerobic microorganisms isolated from cryoconite holes are capable of anaerobic biodegradation via fermentation, and anaerobic metabolism in the holes is dominated by fermentation. An excess of fermentation products, especially acetate, can be detected in defrosted cryoconite sediments and in long-term incubations, indicating incomplete metabolic pathways. Analysis of the microbial community structure revealed that terminal oxidisers are present in low abundance in some of the samples, but the terminal metabolic steps such as sulphate reduction or methanogenesis are not coupled with fermentation. This means that fermentation products could be released to downstream environments, but the extent and impact of this process requires further quantification. Anaerobic metabolism and anaerobic microorganisms are therefore important components of cryoconite ecosystem, which degrade organic matter and impact the glacier carbon cycle

Rapid development of anoxic niches in supraglacial ecosystems

Microorganisms play a significant role in changing the physical properties of the surface of the Greenland Ice Sheet. Cryoconite holes are a hotspot for this microbial activity, yet little is known about the REDOX conditions that develop within them. In this study, we used oxygen microelectrodes and microoptodes to measure for anoxic conditions at the microscale, for the first time revealing a potential niche for anaerobic microorganisms and anaerobic processes. The development of an anoxic zone 2 mm deep within a 6 mm-thick layer of cryoconite sediment was observed within an hour of disturbance, showing rapid acclimation to changing physical conditions. Long-term (half year) incubations of cryoconite material showed a peak of oxygen production and consumption after forty days and reached a low-activity, steady state by day 116, with a persisting anoxic zone beginning between 2 mm and 4 mm deep. Anaerobic microorganisms, which have received little attention to date, should therefore be considered an important component of the cryoconite ecosystem. We discuss the possible dynamics of oxygen concentrations in the supraglacial system and infer that anoxic zones are an important factor in the development of cryoconite sediment communities

Physiological capabilities of cryoconite hole microorganisms

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

Polar cryoconite associated microbiota is dominated by hemispheric specialist genera

Cryoconite holes, supraglacial depressions containing water and microbe-mineral aggregates, are known to be hotspots of microbial diversity on glacial surfaces. Cryoconite holes form in a variety of locations and conditions, which impacts both their structure and the community that inhabits them. Using high-throughput 16S and 18S rRNA gene sequencing, we have investigated the communities of a wide range of cryoconite holes from 15 locations across the Arctic and Antarctic. Around 24 bacterial and 11 eukaryotic first-rank phyla were observed in total. The various biotic niches (grazer, predator, photoautotroph, and chemotroph), are filled in every location. Significantly, there is a clear divide between the bacterial and microalgal communities of the Arctic and that of the Antarctic. We were able to determine the groups contributing to this difference and the family and genus level. Both polar regions contain a “core group” of bacteria that are present in the majority of cryoconite holes and each contribute >1% of total amplicon sequence variant (ASV) abundance. Whilst both groups contain Microbacteriaceae, the remaining members are specific to the core group of each polar region. Additionally, the microalgal communities of Arctic cryoconite holes are dominated by Chlamydomonas whereas the Antarctic cryoconite holes are dominated by Pleurastrum. Therefore cryoconite holes may be a global feature of glacier landscapes, but they are inhabited by regionally distinct microbial communities. Our results are consistent with the notion that cryoconite microbiomes are adapted to differing conditions within the cryosphere

Algal photophysiology drives darkening and melt of the Greenland Ice Sheet

Blooms of Zygnematophycean “glacier algae” lower the bare ice albedo of the Greenland Ice Sheet (GrIS), amplifying summer energy absorption at the ice surface and enhancing meltwater runoff from the largest cryospheric contributor to contemporary sea-level rise. Here, we provide a step change in current understanding of algal-driven ice sheet darkening through quantification of the photophysiological mechanisms that allow glacier algae to thrive on and darken the bare ice surface. Significant secondary phenolic pigmentation (11 times the cellular content of chlorophyll a) enables glacier algae to tolerate extreme irradiance (up to ∼4,000 µmol photons⋅m−2⋅s−1) while simultaneously repurposing captured ultraviolet and short-wave radiation for melt generation. Total cellular energy absorption is increased 50-fold by phenolic pigmentation, while glacier algal chloroplasts positioned beneath shading pigments remain low-light–adapted (Ek ∼46 µmol photons⋅m−2⋅s−1) and dependent upon typical nonphotochemical quenching mechanisms for photoregulation. On the GrIS, glacier algae direct only ∼1 to 2.4% of incident energy to photochemistry versus 48 to 65% to ice surface melting, contributing an additional ∼1.86 cm water equivalent surface melt per day in patches of high algal abundance (∼104 cells⋅mL−1). At the regional scale, surface darkening is driven by the direct and indirect impacts of glacier algae on ice albedo, with a significant negative relationship between broadband albedo (Moderate Resolution Imaging Spectroradiometer [MODIS]) and glacier algal biomass (R2 = 0.75, n = 149), indicating that up to 75% of the variability in albedo across the southwestern GrIS may be attributable to the presence of glacier algae

Is oxygenation related to the decomposition of organic matter in cryoconite holes?

Cryoconite is a sediment occurring on glacier surfaces worldwide which reduces ice albedo and concentrates glacier surface meltwater into small reservoirs called cryoconite holes. It consists of mineral and biogenic matter, including active microorganisms. This study presents an experimental insight into the influence of sediment oxygenation on the cryoconite ability to produce and decomposition of organic matter. Samples were collected from five glaciers in the Arctic and the European mainland. Cryoconite from three glaciers was incubated in stagnant and mechanically mixed conditions to imitate inter-hole water–sediment mixing by meltwater occurring on glaciers in Northern Hemisphere, and its effect on oxygen profiles and organic matter content. Moreover, we investigated short-term changes of oxygen conditions in cryoconite from four glaciers in illuminated and dark conditions. An anaerobic zone was present or approaching zero oxygen in all illuminated cryoconite samples, varying in depth depending on the origin of cryoconite: from 1500 µm from Steindalsbreen (Scandinavian Peninsula) and Forni Glacier (The Alps) to 3100 µm from Russell Glacier and Longyearbreen (Arctic) after incubation. Organic matter content varied between glaciers from 6.11% on Longyearbreen to 16.36% on Russell Glacier. The mixed sediment from the Forni Glacier had less organic matter than stagnant, the sediment from Longyearbreen followed this trend, but the difference was not statistically significant, while the sediment from Ebenferner did not differ between groups. Our results have implications for the understanding of biogeochemical processes on glacier surfaces, the adaptation of organisms to changing physical conditions due to abrupt sediment mixing, but also on the estimation of productivity of supraglacial systems

Cryoconite – from minerals and organic matter to bioengineeredsediments on glacier's surfaces

Cryoconite is a mixture of mineral and organic material covering glacial ice, playing important roles in biogeochemical cycles and lowering the albedo of a glacier surface. Understanding the differences in structure of cryoconite across the globe can be important in recognizing past and future changes in supraglacial environments and ice-organisms-minerals interactions. Despite the worldwide distribution and over a century of studies, the basic characteristics of cryoconite, including its forms and geochemistry, remain poorly studied. The major purpose of our study is the presentation and description of morphological diversity, chemical and photoautotrophs composition, and organic matter content of cryoconite sampled from 33 polar and mountain glaciers around the globe. Observations revealed that cryoconite is represented by various morphologies including loose and granular forms. Granular cryoconite includes smooth, rounded, or irregularly shaped forms; with some having their surfaces covered by cyanobacteria filaments. The occurrence of granules increased with the organic matter content in cryoconite.Moreover, amajor driver of cryoconite colouringwas the concentration of organicmatter and its interplay with minerals. The structure of cyanobacteria and algae communities in cryoconite differs between glaciers, but representatives of cyanobacteria families Pseudanabaenaceae and Phormidiaceae, and algae families Mesotaeniaceae and Ulotrichaceaewere themost common. Themost of detected cyanobacterial taxa are known to produce polymeric substances (EPS) that may cement granules. Organic matter content in cryoconite varied between glaciers, ranging from 1% to 38%. The geochemistry of all the investigated samples reflected local sediment sources, except of highly concentrated Pb andHg in cryoconite collected fromEuropean glaciers near industrialized regions, corroborating cryoconite as element-specific collector and potential environmental indicator of anthropogenic activity. Our work supports a notion that cryoconite may bemore than just simple sediment and instead exhibits complex structure with relevance for biodiversity and the functioning of glacial ecosystem

Anthropogenic contaminants in glacial environments II: Release and downstream consequences

Anthropogenic contamination has been detected in glacial and proglacial environments around the globe. Through mechanisms of secondary release, these contaminants are finding their way into glacial hydrological systems and downstream environments, with potential to impact hundreds of millions of people who rely on glacial meltwater for water, food and energy security worldwide. The first part of our progress report outlined the sources and accumulation mechanisms of contaminants in glacial environments (Part I: Inputs and accumulation). Here we assess processes of contaminant release, pathways to downstream environments, and socio-environmental consequences. We reflect on the potential impacts these contaminants could have for human, ecosystem, and environmental health, as well as framing glacial contaminants within the context of the water-food-energy nexus. Improved understanding of these processes and impacts, while crucially embedding local knowledge, will help to develop key policy and mitigation strategies to address future risk of contaminant release from glaciers