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Cryopeg brines are isolated volumes of hypersaline water in subzero permafrost. The cryopeg system at Utqiaġvik, Alaska, is estimated to date back to 40 ka BP or earlier, a remnant of a late Pleistocene Ocean. Surprisingly, the cryopeg brines contain high concentrations of organic carbon, including extracellular polysaccharides, and high densities of bacteria. How can these physiologically extreme, old, and geologically isolated systems support such an ecosystem? This study addresses this question by examining the energetics of the Utqiaġvik cryopeg brine ecosystem. Using literature-derived assumptions and new measurements on archived borehole materials, we first estimated the quantity of organic carbon when the system formed. We then considered two bacterial growth trajectories to calculate the lower and upper bounds of the cellspecific metabolic rate of these communities. These bounds represent the first community estimates of metabolic rate in a subzero hypersaline environment. To assess the plausibility of the different growth trajectories, we developed a model of the organic carbon cycle and applied it to three borehole scenarios. We also used dissolved inorganic carbon and nitrogen measurements to independently estimate the metabolic rate. The model reconstructs the growth trajectory of the microbial community and predicts the present-day cell density and organic carbon content. Model input included measured rates of the in-situ enzymatic conversion of particulate to dissolved organic carbon under subzero brine conditions. A sensitivity analysis of model parameters was performed, revealing an interplay between growth rate, cell-specific metabolic rate, and extracellular enzyme activity. This approach allowed us to identify plausible growth trajectories consistent with the observed bacterial densities in the cryopeg brines. We found that the cell-specific metabolic rate in this system is relatively high compared to marine sediments. We attribute this finding to the need to invest energy in the production of extracellular enzymes, for generating bioavailable carbon from particulate organic carbon, and the production of extracellular polysaccharides for cryoprotection and osmoprotection. These results may be relevant to other isolated systems in the polar regions of Earth and to possible ice-bound brines on worlds such as Europa, Enceladus, and Mars. [ABSTRACT FROM AUTHOR]

Cryopeg brines are isolated volumes of hypersaline water in subzero permafrost. The cryopeg system at Utqiaġvik, Alaska, is estimated to date back to 40  ka BP or earlier, a remnant of a late Pleistocene Ocean. Surprisingly, the cryopeg brines contain high concentrations of organic carbon, including extracellular polysaccharides, and high densities of bacteria. How can these physiologically extreme, old, and geologically isolated systems support such an ecosystem? This study addresses this question by examining the energetics of the Utqiaġvik cryopeg brine ecosystem. Using literature-derived assumptions and new measurements on archived borehole materials, we first estimated the quantity of organic carbon when the system formed. We then considered two bacterial growth trajectories to calculate the lower and upper bounds of the cellspecific metabolic rate of these communities. These bounds represent the first community estimates of metabolic rate in a subzero hypersaline environment. To assess the plausibility of the different growth trajectories, we developed a model of the organic carbon cycle and applied it to three borehole scenarios. We also used dissolved inorganic carbon and nitrogen measurements to independently estimate the metabolic rate. The model reconstructs the growth trajectory of the microbial community and predicts the present-day cell density and organic carbon content. Model input included measured rates of the in-situ enzymatic conversion of particulate to dissolved organic carbon under subzero brine conditions. A sensitivity analysis of model parameters was performed, revealing an interplay between growth rate, cell-specific metabolic rate, and extracellular enzyme activity. This approach allowed us to identify plausible growth trajectories consistent with the observed bacterial densities in the cryopeg brines. We found that the cell-specific metabolic rate in this system is relatively high compared to marine sediments. We attribute this finding to the need to invest energy in the production of extracellular enzymes, for generating bioavailable carbon from particulate organic carbon, and the production of extracellular polysaccharides for cryoprotection and osmoprotection. These results may be relevant to other isolated systems in the polar regions of Earth and to possible ice-bound brines on worlds such as Europa, Enceladus, and Mars. [ABSTRACT FROM AUTHOR]

In Arctic soils, warming accelerates decomposition of organic matter and increases emission of greenhouse gases (GHGs), contributing to a positive feedback to climate change. Although microorganisms play a key role in the processes between decomposition of organic matter and GHGs emission, the effects of warming on temporal responses of microbial activity are still elusive. In this study, treatments of warming and precipitation were conducted from 2012 to 2018 in Cambridge Bay, Canada. Soils of organic and mineral layers were collected monthly from June to September in 2018 and analyzed for extracellular enzyme activities and bacterial community structures. The activity of hydrolases was the highest in June and decreased thereafter over summer in both organic and mineral layers. Bacterial community structures changed gradually over summer, and the responses were distinct depending on soil layers and environmental factors; water content and soil temperature affected the shift of bacterial community structures in both layers, whereas bacterial abundance, dissolved organic carbon, and inorganic nitrogen did so in the organic layer only. The activity of hydrolases and bacterial community structures did not differ significantly among treatments but among months. Our results demonstrate that temporal variations may control extracellular enzyme activities and microbial community structure rather than the small effect of warming over a long period in high Arctic soil. Although the effects of the treatments on microbial activity were minor, our study provides insight that microbial activity may increase due to an increase in carbon availability, if the growing season is prolonged in the Arctic. [ABSTRACT FROM AUTHOR]

Arctic terrestrial vegetation is responding to warming with higher biomass and productivity, but the effects for freshwater ecosystems are unclear. Biofilm extracellular enzymes are important mediators of organic matter processing in aquatic ecosystems, thus understanding how enzyme activities change with water temperature and organic matter availability is important to assess the effects of climate change. Here, we studied biofilm enzyme activities in six streams differing in their riparian vegetation cover and water thermal regime on Disko Island, Greenland. For all streams, biofilms enzymes showed a low activity to decompose plant material, as expected in the Arctic. However, in contrast to simple polysaccharides, there was a significant increase in the capacity for hemicellulose decomposition, in streams with high-vegetation cover. Moreover, biofilms in high-vegetation streams showed greater autotrophy (chlorophyll a to total organic matter content) and higher phosphatase activity than stream with low-vegetation cover ones. Enzyme temperature sensitivity (measured as the rate of change of enzyme activity by biofilms incubated at 3 and 22 °C) did not change across the streams, but differed among enzymes types. Phosphatase and phenol oxidase enzymes presented the highest sensitivity to water temperature, which suggests that the degradation of organic phosphorous compounds and lignin could be disproportionately enhanced under warming climate scenarios. [ABSTRACT FROM AUTHOR]

Permafrost thaw in the Arctic driven by climate change is mobilizing ancient terrigenous organic carbon ( OC) into fluvial networks. Understanding the controls on metabolism of this OC is imperative for assessing its role with respect to climate feedbacks. In this study, we examined the effect of inorganic nutrient supply and dissolved organic matter ( DOM) composition on aquatic extracellular enzyme activities ( EEAs) in waters draining the Kolyma River Basin (Siberia), including permafrost-derived OC. Reducing the phenolic content of the DOM pool resulted in dramatic increases in hydrolase EEAs (e.g., phosphatase activity increased >28-fold) supporting the idea that high concentrations of polyphenolic compounds in DOM (e.g., plant structural tissues) inhibit enzyme synthesis or activity, limiting OC degradation. EEAs were significantly more responsive to inorganic nutrient additions only after phenolic inhibition was experimentally removed. In controlled mixtures of modern OC and thawed permafrost endmember OC sources, respiration rates per unit dissolved OC were 1.3-1.6 times higher in waters containing ancient carbon, suggesting that permafrost-derived OC was more available for microbial mineralization. In addition, waters containing ancient permafrost-derived OC supported elevated phosphatase and glucosidase activities. Based on these combined results, we propose that both composition and nutrient availability regulate DOM metabolism in Arctic aquatic ecosystems. Our empirical findings are incorporated into a mechanistic conceptual model highlighting two key enzymatic processes in the mineralization of riverine OM: (i) the role of phenol oxidase activity in reducing inhibitory phenolic compounds and (ii) the role of phosphatase in mobilizing organic P. Permafrost-derived DOM degradation was less constrained by this initial 'phenolic- OM' inhibition; thus, informing reports of high biological availability of ancient, permafrost-derived DOM with clear ramifications for its metabolism in fluvial networks and feedbacks to climate. [ABSTRACT FROM AUTHOR]

Abstract: Arctic systems, which store ∼50% of global soil carbon, are undergoing rapid climatic warming that may drive significant carbon release to the atmosphere. To better understand how warming impacts arctic decomposition, we characterized the effects of a twenty-two year long tundra greenhouse warming experiment on decomposer-produced extracellular enzymes, nutrients, and microbial biomass across a year. This experiment, which is the longest running tundra ecosystem warming study in existence, was previously shown to have altered the plant and soil communities. The greenhouse treatment has also changed the seasonal soil temperature regime by indirectly increasing winter soil temperature, an effect that was likely facilitated through an increase in snow-trapping shrub biomass. Irrespective of the warming treatment, we observed that peak nutrient pools, microbial biomass, and hydrolytic enzyme activities all occurred from the late winter through thaw. This pattern was decoupled from peak oxidative enzyme activities, which occurred during the summer. The greenhouse treatment amplified the natural seasonal cycle of extracellular enzyme activities, suggesting that tundra decomposer communities maintain a temporal niche space which is critical to understanding how arctic biogeochemical cycling will respond to warming. A spatial separation was also observed; extracellular enzyme activities in the deeper soil horizons were more sensitive to warming than at the surface. Direct greenhouse warming did not strongly stimulate decomposition: only oxidative enzyme activities in the surface horizon increased during the summer. Unexpectedly, the strongest treatment effect observed was a stimulation of hydrolytic enzyme activities at depth in the mineral horizon from the late winter through thaw (which also affected extracellular enzyme stoichiometry, increasing C:N and C:P acquisition activities), before the greenhouse treatment was directly active. This effect declined during senescence and was reversed in early winter, suggesting that negative biotic-abiotic feedbacks may curtail increased decomposer activity in warming arctic systems. [Copyright &y& Elsevier]

Abstract: The active lifetime of extracellular enzymes is a critical determinant of the effectiveness of enzyme production as a means for heterotrophic marine microbes to obtain organic substrates. Here, we report lifetimes of three classes of extracellular enzyme in Arctic seawater. We also investigated the relative importance of photochemical processes and particle-associated processes in inactivating extracellular enzymes. Enzyme inactivation in filtered seawater was slow, with apparent half-lives of enzyme activities on the order of hundreds of hours. The presence of particles (including cells) did not significantly change inactivation rates, suggesting that the long half-lives observed in filtered seawater were realistic for enzymes in unfiltered seawater. Phosphatase and leucine aminopeptidase were susceptible to photoinactivation, but only under high intensity UV-B and UV-C illumination; there was no evidence for increased inactivation rates under natural illumination at our study site in Ny Ålesund, Svalbard. Comparison of inactivation rates of commercially-obtained enzymes from non-marine sources with the extracellular enzymes naturally present in Arctic seawater suggests that the natural enzymes contain structural features that confer longer lifetimes, consistent with observations reported by others from a range of field sites that cell-free enzymes can contribute a substantial fraction of total hydrolytic activity in the water column. [Copyright &y& Elsevier]

Arctic soils contain large amounts of organic matter due to very slow rates of detritus decomposition. The first step in decomposition results from the activity of extracellular enzymes produced by soil microbes. We hypothesized that potential enzyme activities are low relative to the large stocks of organic matter in Arctic tundra soils, and that enzyme activity is low at in situ temperatures. We measured the potential activity of six hydrolytic enzymes at 4 and 20 °C on four sampling dates in tussock, intertussock, shrub organic, and shrub mineral soils at Toolik Lake, Alaska. Potential activities of N-acetyl glucosaminidase, β-glucosidase, and peptidase tended to be greatest at the end of winter, suggesting that microbes produced enzymes while soils were frozen. In general, enzyme activities did not increase during the Arctic summer, suggesting that enzyme production is N-limited during the period when temperatures would otherwise drive higher enzyme activity in situ. We also detected seasonal variations in the temperature sensitivity ( Q10) of soil enzymes. In general, soil enzyme pools were more sensitive to temperature at the end of the winter than during the summer. We modeled potential in situβ-glucosidase activities for tussock and shrub organic soils based on measured enzyme activities, temperature sensitivities, and daily soil temperature data. Modeled in situ enzyme activity in tussock soils increased briefly during the spring, then declined through the summer. In shrub soils, modeled enzyme activities increased through the spring thaw into early August, and then declined through the late summer and into winter. Overall, temperature is the strongest factor driving low in situ enzyme activities in the Arctic. However, enzyme activity was low during the summer, possibly due to N-limitation of enzyme production, which would constrain enzyme activity during the brief period when temperatures would otherwise drive higher rates of decomposition. [ABSTRACT FROM AUTHOR]

In the Arctic, climate changes contribute to enhanced mobilization of organic matter in streams. Microbial extracellular enzymes are important mediators of stream organic matter processing, but limited information is available on enzyme processes in this remote area. Here, we studied the variability of microbial extracellular enzyme activity in high-Arctic fluvial biofilms. We evaluated 12 stream reaches in Northeast Greenland draining areas exhibiting different geomorphological features with contrasting contents of soil organic matter to cover a wide range of environmental conditions. We determined stream nitrogen, phosphorus, and dissolved organic carbon concentrations, quantified algal biomass and bacterial density, and characterized the extracellular enzyme activities involved in catalyzing the cleavage of a range of organic matter compounds (e.g., β-glucosidase, phosphatase, β-xylosidase, cellobiohydrolase, and phenol oxidase). We found significant differences in microbial organic matter utilization among the study streams draining contrasting geomorphological features, indicating a strong coupling between terrestrial and stream ecosystems. Phosphatase and phenol oxidase activities were higher in solifluction areas than in alluvial areas. Besides dissolved organic carbon, nitrogen availability was the main driver controlling enzyme activities in the high-Arctic, which suggests enhanced organic matter mineralization at increased nutrient availability. Overall, our study provides novel information on the controls of organic matter usage by high-Arctic stream biofilms, which is of high relevance due to the predicted increase of nutrient availability in high-Arctic streams in global climate change scenarios.