Photoinduced Evolutions of Permafrost-Derived Carbon in Subarctic Thermokarst Pond Surface Waters.

In subarctic regions, rising temperature and permafrost thaw lead to the formation of thermokarst ponds, where organics from eroding permafrost accumulate. Despite its environmental significance, limited knowledge exists regarding the photosensitivity of permafrost-derived carbon in these ponds. In this study, laboratory experiments were conducted to explore the photochemical transformations of organic matter in surface water samples from thermokarst ponds from different environments in northern Quebec, Canada. One pond near Kuujjuarapik is characterized by the presence of a collapsing palsa and is therefore organically rich, while the other pond near Umiujaq is adjacent to a collapsing lithalsa and thus contains fewer organic matters. Photobleaching occurred in the Umiujaq sample upon irradiation, whereas the Kuujjuarapik sample exhibited an increase in light absorbance at wavelength related to aromatic functionalities, indicating different photochemical aging processes. Ultrahigh-resolution mass spectrometry analysis reveals that the Kuujjuarapik sample preferentially photoproduced highly unsaturated CHO compounds with great aromaticity, while the irradiated Umiujaq sample produced a higher proportion of CHON aromatics with reduced nitrogen functionalities. Overall, this study illustrates that the photochemical reactivity of thermokarst pond water varies with the source of organic matter. The observed differences in reactivity contribute to an improved understanding of the photochemical emission of volatile organic compounds discovered earlier. Further insights into the photoinduced evolutions in thermokarst ponds may require the classification of permafrost-derived carbon therein.

Extensive loss of past permafrost carbon but a net accumulation into present-day soils.

Atmospheric concentrations of carbon dioxide increased between the Last Glacial Maximum (LGM, around 21,000 years ago) and the preindustrial era1. It is thought that the evolution of this atmospheric carbon dioxide (and that of atmospheric methane) during the glacial-to-interglacial transition was influenced by organic carbon that was stored in permafrost during the LGM and then underwent decomposition and release following thaw2,3. It has also been suggested that the rather erratic atmospheric δ13C and ∆14C signals seen during deglaciation1,4 could partly be explained by the presence of a large terrestrial inert LGM carbon stock, despite the biosphere being less productive (and therefore storing less carbon)5,6. Here we present an empirically derived estimate of the carbon stored in permafrost during the LGM by reconstructing the extent and carbon content of LGM biomes, peatland regions and deep sedimentary deposits. We find that the total estimated soil carbon stock for the LGM northern permafrost region is smaller than the estimated present-day storage (in both permafrost and non-permafrost soils) for the same region. A substantial decrease in the permafrost area from the LGM to the present day has been accompanied by a roughly 400-petagram increase in the total soil carbon stock. This increase in soil carbon suggests that permafrost carbon has made no net contribution to the atmospheric carbon pool since the LGM. However, our results also indicate potential postglacial reductions in the portion of the carbon stock that is trapped in permafrost, of around 1,000 petagrams, supporting earlier studies7. We further find that carbon has shifted from being primarily stored in permafrost mineral soils and loess deposits during the LGM, to being roughly equally divided between peatlands, mineral soils and permafrost loess deposits today.

Genome-centric view of carbon processing in thawing permafrost.

As global temperatures rise, large amounts of carbon sequestered in permafrost are becoming available for microbial degradation. Accurate prediction of carbon gas emissions from thawing permafrost is limited by our understanding of these microbial communities. Here we use metagenomic sequencing of 214 samples from a permafrost thaw gradient to recover 1,529 metagenome-assembled genomes, including many from phyla with poor genomic representation. These genomes reflect the diversity of this complex ecosystem, with genus-level representatives for more than sixty per cent of the community. Meta-omic analysis revealed key populations involved in the degradation of organic matter, including bacteria whose genomes encode a previously undescribed fungal pathway for xylose degradation. Microbial and geochemical data highlight lineages that correlate with the production of greenhouse gases and indicate novel syntrophic relationships. Our findings link changing biogeochemistry to specific microbial lineages involved in carbon processing, and provide key information for predicting the effects of climate change on permafrost systems.

Methane release from carbonate rock formations in the Siberian permafrost area during and after the 2020 heat wave.

Anthropogenic global warming may be accelerated by a positive feedback from the mobilization of methane from thawing Arctic permafrost. There are large uncertainties about the size of carbon stocks and the magnitude of possible methane emissions. Methane cannot only be produced from the microbial decay of organic matter within the thawing permafrost soils (microbial methane) but can also come from natural gas (thermogenic methane) trapped under or within the permafrost layer and released when it thaws. In the Taymyr Peninsula and surroundings in North Siberia, the area of the worldwide largest positive surface temperature anomaly for 2020, atmospheric methane concentrations have increased considerably during and after the 2020 heat wave. Two elongated areas of increased atmospheric methane concentration that appeared during summer coincide with two stripes of Paleozoic carbonates exposed at the southern and northern borders of the Yenisey-Khatanga Basin, a hydrocarbon-bearing sedimentary basin between the Siberian Craton to the south and the Taymyr Fold Belt to the north. Over the carbonates, soils are thin to nonexistent and wetlands are scarce. The maxima are thus unlikely to be caused by microbial methane from soils or wetlands. We suggest that gas hydrates in fractures and pockets of the carbonate rocks in the permafrost zone became unstable due to warming from the surface. This process may add unknown quantities of methane to the atmosphere in the near future.

Reduced methane emissions in former permafrost soils driven by vegetation and microbial changes following drainage.

In Arctic regions, thawing permafrost soils are projected to release 50 to 250 Gt of carbon by 2100. This data is mostly derived from carbon-rich wetlands, although 71% of this carbon pool is stored in faster-thawing mineral soils, where ecosystems close to the outer boundaries of permafrost regions are especially vulnerable. Although extensive data exists from currently thawing sites and short-term thawing experiments, investigations of the long-term changes following final thaw and co-occurring drainage are scarce. Here we show ecosystem changes at two comparable tussock tundra sites with distinct permafrost thaw histories, representing 15 and 25 years of natural drainage, that resulted in a 10-fold decrease in CH4 emissions (3.2 ± 2.2 vs. 0.3 ± 0.4 mg C-CH4  m-2  day-1 ), while CO2 emissions were comparable. These data extend the time perspective from earlier studies based on short-term experimental drainage. The overall microbial community structures did not differ significantly between sites, although the drier top soils at the most advanced site led to a loss of methanogens and their syntrophic partners in surface layers while the abundance of methanotrophs remained unchanged. The resulting deeper aeration zones likely increased CH4 oxidation due to the longer residence time of CH4 in the oxidation zone, while the observed loss of aerenchyma plants reduced CH4 diffusion from deeper soil layers directly to the atmosphere. Our findings highlight the importance of including hydrological, vegetation and microbial specific responses when studying long-term effects of climate change on CH4 emissions and underscores the need for data from different soil types and thaw histories.

Microbiome assembly in thawing permafrost and its feedbacks to climate.

The physical and chemical changes that accompany permafrost thaw directly influence the microbial communities that mediate the decomposition of formerly frozen organic matter, leading to uncertainty in permafrost-climate feedbacks. Although changes to microbial metabolism and community structure are documented following thaw, the generality of post-thaw assembly patterns across permafrost soils of the world remains uncertain, limiting our ability to predict biogeochemistry and microbial community responses to climate change. Based on our review of the Arctic microbiome, permafrost microbiology, and community ecology, we propose that Assembly Theory provides a framework to better understand thaw-mediated microbiome changes and the implications for community function and climate feedbacks. This framework posits that the prevalence of deterministic or stochastic processes indicates whether the community is well-suited to thrive in changing environmental conditions. We predict that on a short timescale and following high-disturbance thaw (e.g., thermokarst), stochasticity dominates post-thaw microbiome assembly, suggesting that functional predictions will be aided by detailed information about the microbiome. At a longer timescale and lower-intensity disturbance (e.g., active layer deepening), deterministic processes likely dominate, making environmental parameters sufficient for predicting function. We propose that the contribution of stochastic and deterministic processes to post-thaw microbiome assembly depends on the characteristics of the thaw disturbance, as well as characteristics of the microbial community, such as the ecological and phylogenetic breadth of functional guilds, their functional redundancy, and biotic interactions. These propagate across space and time, potentially providing a means for predicting the microbial forcing of greenhouse gas feedbacks to global climate change.

Ancient RNA from Late Pleistocene permafrost and historical canids shows tissue-specific transcriptome survival.

While sequencing ancient DNA (aDNA) from archaeological material is now commonplace, very few attempts to sequence ancient transcriptomes have been made, even from typically stable deposition environments such as permafrost. This is presumably due to assumptions that RNA completely degrades relatively quickly, particularly when dealing with autolytic, nuclease-rich mammalian tissues. However, given the recent successes in sequencing ancient RNA (aRNA) from various sources including plants and animals, we suspect that these assumptions may be incorrect or exaggerated. To challenge the underlying dogma, we generated shotgun RNA data from sources that might normally be dismissed for such study. Here, we present aRNA data generated from two historical wolf skins, and permafrost-preserved liver tissue of a 14,300-year-old Pleistocene canid. Not only is the latter the oldest RNA ever to be sequenced, but it also shows evidence of biologically relevant tissue specificity and close similarity to equivalent data derived from modern-day control tissue. Other hallmarks of RNA sequencing (RNA-seq) data such as exon-exon junction presence and high endogenous ribosomal RNA (rRNA) content confirms our data's authenticity. By performing independent technical library replicates using two high-throughput sequencing platforms, we show not only that aRNA can survive for extended periods in mammalian tissues but also that it has potential for tissue identification. aRNA also has possible further potential, such as identifying in vivo genome activity and adaptation, when sequenced using this technology.

Approaches for the complex assessment of polychemical pollution of permafrost-affected soils and the upper layer of permafrost.

The analysis of about 200 samples taken from 42 permafrost-affected soil profiles was carried out on four key sites in different regions of cryolithozone (West Siberia, Central, North, and North-East Yakutia) characterized by different active layer depths and soil lithology. The aim of the study was to determine the influence of different processes of cryogenic mass-exchange on the redistribution and accumulation of major pollutants such as petroleum products, acid-soluble forms of trace elements, polycyclic hydrocarbons, and technogenic radionuclides transferred via atmospheric transport or after the local anthropogenic impact in different soil horizons of Cryosols and in the upper layers of permafrost. Samples were analyzed using modern precise techniques (direct γ-spectrometric measurements with Ge(Li) and NaI(Tl) detectors; fluorometric method; reversed-phase high-performance liquid chromatography; spectrofluorimetric detection; atomic absorption spectrometry with flame atomization). The study has shown that processes (cryoturbations, frost heaving, gelifluction along with fluvial processes) that strongly affect Cryosols' profile structure can also lead to the active migration and accumulation of local and global pollutants in the middle and lowermost suprapermafrost soil horizons. The accumulation of some pollutants in suprapermafrost horizons of cryogenic soils and in the upper layers of permafrost (in particular, petroleum products and mobile forms of trace elements) can be associated with a combination of factors, such as the relatively light particle size distribution, relatively weak manifestation of cryoturbation processes, and low thickness of the active layer (about 40-60 cm). The integral calculation of the geoaccumulation index values has shown that all of the groups of human-affected soil horizons are moderately to extremely polluted by petroleum hydrocarbons (and at a relatively lower level by trace elements) and the maximum pollution stands for the suprapermafrost horizons as well as in cryoturbated or buried fragments of organogenic matter in some cases. The maxima of the heavy PAH content in permafrost-affected soils can be confined to horizons enriched with anthropogenic inclusions and artifacts (for example, construction slag, coal) and to individual horizons of soils buried as a result of both cryogenic and alluvial processes. The specific activity of the technogenic radionuclide cesium in cryogenic soils revealed its association mainly with the surface organogenic and organomineral horizons of the studied profiles and rarely observed in the cryoturbated fragments of these horizons in the middle and suprapermafrost layers of soil profiles. The necessity of the complex analytical assessment of the permafrost-affected soils has been revealed especially in case of studying of the ecological state of the anthropogenically affected Cryosols.

Quantification of endospores in ancient permafrost using time-resolved terbium luminescence.

We describe herein a simple procedure for quantifying endospore abundances in ancient and organic-rich permafrost. We repeatedly (10x) extracted and fractionated permafrost using a tandem filter assembly composed of 3 and 0.2 µm filters. Then, the 0.2 µm filter was washed (7x), autoclaved, and the contents eluted, including dipicolinic acid (DPA). Time-resolved luminescence using Tb(EDTA) yielded a LOD of 1.46 nM DPA (6.55 × 103 endospores/mL). In review, DPA/endospore abundances were ~2.2-fold greater in older 33 ky permafrost (258 ± 36 pmol DPA gdw-1; 1.15 × 106 ± 0.16 × 106 spores gdw-1) versus younger 19 ky permafrost (p = 0.007297). This suggests that dormancy increases with permafrost age.

Climate change and the permafrost carbon feedback.

Large quantities of organic carbon are stored in frozen soils (permafrost) within Arctic and sub-Arctic regions. A warming climate can induce environmental changes that accelerate the microbial breakdown of organic carbon and the release of the greenhouse gases carbon dioxide and methane. This feedback can accelerate climate change, but the magnitude and timing of greenhouse gas emission from these regions and their impact on climate change remain uncertain. Here we find that current evidence suggests a gradual and prolonged release of greenhouse gas emissions in a warming climate and present a research strategy with which to target poorly understood aspects of permafrost carbon dynamics.

Intra- and inter-cores fungal diversity suggests interconnection of different habitats in an Antarctic frozen lake (Boulder Clay, Northern Victoria Land).

A perennially frozen lake at Boulder Clay site (Victoria Land, Antarctica), characterized by the presence of frost mounds, have been selected as an in situ model for ecological studies. Different samples of permafrost, glacier ice and brines have been studied as a unique habitat system. An additional sample of brines (collected in another frozen lake close to the previous one) was also considered. Alpha- and beta-diversity of fungal communities showed both intra- and inter-cores significant (p < 0.05) differences, which suggest the presence of interconnection among the habitats. Therefore, the layers of frost mound and the deep glacier could be interconnected while the brines could probably be considered as an open habitat system not interconnected with each other. Moreover, the absence of similarity between the lake ice and the underlying permafrost suggested that the lake is perennially frozen based. The predominance of positive significant (p < 0.05) co-occurrences among some fungal taxa allowed to postulate the existence of an ecological equilibrium in the habitats systems. The positive significant (p < 0.05) correlation between salt concentration, total organic carbon and pH, and some fungal taxa suggests that a few abiotic parameters could drive fungal diversity inside these ecological niches.

Effect of Cysteine Residue Substitution in the GCSAG Motif of the PMGL2 Esterase Active Site on the Enzyme Properties.

The gene coding for PMGL2 esterase, which belongs to the family of mammalian hormone-sensitive lipases (HSLs), was discovered by screening a metagenomic DNA library from a permafrost soil. The active site of PMGL2 contains conserved GXSXG motif which includes Cys173 residue next to the catalytic Ser174. In order to clarify the functional role of the cysteine residue in the GCSAG motif, we constructed a number of PMGL2 mutants with Cys173 substitutions and studied their properties. The specific activity of the C173D mutant exceeded the specific activity of the wild-type enzyme (wtPMGL2) by 60%, while the C173T/C202S mutant displayed reduced catalytic activity. The activity of the C173D mutant with p-nitrophenyl octanoate was 15% higher, while the activity of the C173T/C202S mutant was 17% lower compared to wtPMGL2. The C173D mutant was also characterized by a high activity at low temperatures (20-35°C) and significant loss of thermal stability. The kcat value for this protein was 56% higher than for the wild-type enzyme. The catalytic constants of the C173S mutant were close to those of wtPMGL2; this enzyme also demonstrated the highest thermal stability among the studied mutants. The obtained results demonstrate that substitutions of amino acid residues adjacent to the catalytic serine residue in the GXSXG motif can have a significant effect on the properties of PMGL2 esterase.

Diversity and Distribution of Methanogenic Community Between Two Typical Alpine Ecosystems on the Qinghai-Tibetan Plateau.

Alpine permafrost regions are important sources of biogenic CH4 and methanogens play an important role in the methane-producing process. The alpine permafrost on the Qinghai-Tibetan plateau comprises about one-sixth of China's land area, and there are various types of alpine ecosystems. However, the methanogenic communities in the typical alpine ecosystems are poorly understood. In this study, the active layers and permafrost layers of the natural ecosystem of alpine grassland (DZ2-1) and alpine swamp meadow (DZ2-5) were selected to investigate the diversity and abundance of methanogenic communities. Methanobacterium (63.65%) are overwhelmingly dominant in the active layer of the alpine grassland (DZ2-1A). ZC-I cluster (26.13%), RC-I cluster (19.56%), and Methanobacterium (15.02%) are the dominant groups in the permafrost layer of the alpine grassland (DZ2-1P). Methanosaeta (32.92%), Fen cluster (29.59%), Methanosarcina (16.33%), and Methanobacterium (13.95%) are the dominant groups in the active layer of the alpine swamp meadow (DZ2-5A), whereas the Fen cluster (50.85%), ZC-I cluster (27.63%), and RC-I cluster (14.15%) are relatively abundant in the permafrost layer of the alpine swamp meadow (DZ2-5P). qPCR data showed that the abundance of methanogens was higher in the natural ecosystem of alpine swamp meadow than in alpine grassland. We found that the community characteristics of methanogens were related to environmental factors. Pearson correlation analyses indicated that the relative abundance of Methanobacterium had a significantly positive correlation with hydrogen concentration (P < 0.01), while the relative abundances of Methanosaeta and Methanosarcina were positively correlated with acetate concentration (P < 0.05). This study will help us to understand the methanogenic communities and their surrounding environments in alpine ecosystems.

Geochemical pollution of trace metals in permafrost-affected soil in the Russian Arctic marginal environment.

The Arctic marginal environment has been considered as far from industrial areas and low population. During June-July of 2016 "Russian High Latitude" expedition, 93 samples of soil genetic horizon from 25 soil profiles dug till frozen ground were sampled from 8 islands and 2 capes of the Russian Arctic without direct anthropogenic influences. Nine trace metals (Pb, Cd, Cu, Ni, Co, Zn, Fe, Mn and Hg) were measured and quantified by energy-dispersive X-ray analysis for elemental concentrations. Through analysis of divided soil groups (Haplothels, Turbels, Historthels), the factors of organic matter and cryoturbation had a significant influence on metals' distribution except for Fe and Mn. From summarized soil master horizons (O, A, B, C), Fe and Mn are abundant in all horizons suggesting as geochemical background values. Cu, Pb, Co and Ni are distributed specifically in different horizons with leaching and accumulation process, whereas Hg is evenly disturbed in all horizons. The correlation analysis reveals that distribution of most metals in present soils is highly depended on soil properties (pH, TOC, clay and silt). Li was selected as normalizing element for metals' concentrations from mineral layers to establish geochemical baseline concentrations. The concentrations of trace metals have been assessed by geoaccumulation index (Igeo) and enrichment factor, showing only Co and Zn are moderately polluted and slightly polluted, and Co, Cu, Zn and Pb are enriched in topsoil. Other indices as modified degree of contamination (mCdegree) and pollution load index (PLI), mCdegree show moderate degree of pollution and PLI shows unpolluted to moderate pollution load. The ecological risk indices, e.g., ecological risk factor (Er) and potential ecological risk index, show low ecological risk potential.

Volatile emissions from thawing permafrost soils are influenced by meltwater drainage conditions.

Vast amounts of carbon are bound in both active layer and permafrost soils in the Arctic. As a consequence of climate warming, the depth of the active layer is increasing in size and permafrost soils are thawing. We hypothesize that pulses of biogenic volatile organic compounds are released from the near-surface active layer during spring, and during late summer season from thawing permafrost, while the subsequent biogeochemical processes occurring in thawed soils also lead to emissions. Biogenic volatile organic compounds are reactive gases that have both negative and positive climate forcing impacts when introduced to the Arctic atmosphere, and the knowledge of their emission magnitude and pattern is necessary to construct reliable climate models. However, it is unclear how different ecosystems and environmental factors such as drainage conditions upon permafrost thaw affect the emission and compound composition. Here we show that incubations of frozen B horizon of the active layer and permafrost soils collected from a High Arctic heath and fen release a range of biogenic volatile organic compounds upon thaw and during subsequent incubation experiments at temperatures of 10°C and 20°C. Meltwater drainage in the fen soils increased emission rates nine times, while having no effect in the drier heath soils. Emissions generally increased with temperature, and emission profiles for the fen soils were dominated by benzenoids and alkanes, while benzenoids, ketones, and alcohols dominated in heath soils. Our results emphasize that future changes affecting the drainage conditions of the Arctic tundra will have a large influence on volatile emissions from thawing permafrost soils - particularly in wetland/fen areas.