Characteristics of methane emissions from alpine thermokarst lakes on the Tibetan Plateau.

Understanding methane (CH4) emission from thermokarst lakes is crucial for predicting the impacts of abrupt thaw on the permafrost carbon-climate feedback. However, observational evidence, especially from high-altitude permafrost regions, is still scarce. Here, by combining field surveys, radio- and stable-carbon isotopic analyses, and metagenomic sequencing, we present multiple characteristics of CH4 emissions from 120 thermokarst lakes in 30 clusters along a 1100 km transect on the Tibetan Plateau. We find that thermokarst lakes have high CH4 emissions during the ice-free period (13.4 ± 1.5 mmol m-2 d-1; mean ± standard error) across this alpine permafrost region. Ebullition constitutes 84% of CH4 emissions, which are fueled primarily by young carbon decomposition through the hydrogenotrophic pathway. The relative abundances of methanogenic genes correspond to the observed CH4 fluxes. Overall, multiple parameters obtained in this study provide benchmarks for better predicting the strength of permafrost carbon-climate feedback in high-altitude permafrost regions.

Microbial methane cycling in sediments of Arctic thermokarst lagoons.

Thermokarst lagoons represent the transition state from a freshwater lacustrine to a marine environment, and receive little attention regarding their role for greenhouse gas production and release in Arctic permafrost landscapes. We studied the fate of methane (CH4 ) in sediments of a thermokarst lagoon in comparison to two thermokarst lakes on the Bykovsky Peninsula in northeastern Siberia through the analysis of sediment CH4 concentrations and isotopic signature, methane-cycling microbial taxa, sediment geochemistry, lipid biomarkers, and network analysis. We assessed how differences in geochemistry between thermokarst lakes and thermokarst lagoons, caused by the infiltration of sulfate-rich marine water, altered the microbial methane-cycling community. Anaerobic sulfate-reducing ANME-2a/2b methanotrophs dominated the sulfate-rich sediments of the lagoon despite its known seasonal alternation between brackish and freshwater inflow and low sulfate concentrations compared to the usual marine ANME habitat. Non-competitive methylotrophic methanogens dominated the methanogenic community of the lakes and the lagoon, independent of differences in porewater chemistry and depth. This potentially contributed to the high CH4 concentrations observed in all sulfate-poor sediments. CH4 concentrations in the freshwater-influenced sediments averaged 1.34 ± 0.98 µmol g-1 , with highly depleted δ13 C-CH4 values ranging from -89‰ to -70‰. In contrast, the sulfate-affected upper 300 cm of the lagoon exhibited low average CH4 concentrations of 0.011 ± 0.005 µmol g-1 with comparatively enriched δ13 C-CH4 values of -54‰ to -37‰ pointing to substantial methane oxidation. Our study shows that lagoon formation specifically supports methane oxidizers and methane oxidation through changes in pore water chemistry, especially sulfate, while methanogens are similar to lake conditions.

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.

Development of permafrost-affected peatlands in the southern limit of the European Russian cryolithozone and their vulnerability to future warming.

Permafrost degradation due to climate warming is currently observed in the northeastern part of European Russia. Peat plateaus underlain by permafrost cover only about 20% of the Russian European cryolithozone but contain almost 50% of soil organic carbon stocks (SOC), which are considered to be vulnerable to microbial mineralization after permafrost thaw. The current study was performed at three key sites of peat plateaus located along the southern permafrost limit. SOC decomposition was studied by aerobic and anaerobic incubation experiments, conducted at 4 °C over a period of 1301 days. The CO2 production was measured in peat samples at three key sites from the active layer (AL), transitional layer (TL), permafrost layer (PL), and at one site from the deep permafrost layer (DPL), which is in contact with mineral soil at 3.7 m depth. During the experiment, the initial СО2 respiration rates significantly differed in the samples AL, TL and PL in all key sites. However, at each site in the majority of samples the CO2 respiration rates were 2-5 times aerobically higher than anaerobically. In anaerobic conditions, in all sites, the СО2 respiration rate in PL was the lowest, higher in TL and the highest in AL in all 3 sites. Projections of CO2 aerobically production for 80 years represent 1.44 ± 0.11, 6.31 ± 0.47, 30.64 ± 17.98% of initial permafrost carbon from the samples of Inta 1, Inta 11 and Kolva respectively. But under anaerobical conditions estimates are close and indicate insignificant amounts 0.30…1.90% of carbon release over a period of 80 years. We suggest that even under ideal conditions of the incubation experiment, without considering ecological inertia under natural conditions, while also permafrost temperature is close to zero, greenhouse gas release from initial SOC is significantly less than estimated.

Greenhouse gas production and lipid biomarker distribution in Yedoma and Alas thermokarst lake sediments in Eastern Siberia.

Permafrost thaw leads to thermokarst lake formation and talik growth tens of meters deep, enabling microbial decomposition of formerly frozen organic matter (OM). We analyzed two 17-m-long thermokarst lake sediment cores taken in Central Yakutia, Russia. One core was from an Alas lake in a Holocene thermokarst basin that underwent multiple lake generations, and the second core from a young Yedoma upland lake (formed ~70 years ago) whose sediments have thawed for the first time since deposition. This comparison provides a glance into OM fate in thawing Yedoma deposits. We analyzed total organic carbon (TOC) and dissolved organic carbon (DOC) content, n-alkane concentrations, and bacterial and archaeal membrane markers. Furthermore, we conducted 1-year-long incubations (4°C, dark) and measured anaerobic carbon dioxide (CO2 ) and methane (CH4 ) production. The sediments from both cores contained little TOC (0.7 ± 0.4 wt%), but DOC values were relatively high, with the highest values in the frozen Yedoma lake sediments (1620 mg L-1 ). Cumulative greenhouse gas (GHG) production after 1 year was highest in the Yedoma lake sediments (226 ± 212 µg CO2 -C g-1  dw, 28 ± 36 µg CH4 -C g-1  dw) and 3 and 1.5 times lower in the Alas lake sediments, respectively (75 ± 76 µg CO2 -C g-1  dw, 19 ± 29 µg CH4 -C g-1  dw). The highest CO2 production in the frozen Yedoma lake sediments likely results from decomposition of readily bioavailable OM, while highest CH4 production in the non-frozen top sediments of this core suggests that methanogenic communities established upon thaw. The lower GHG production in the non-frozen Alas lake sediments resulted from advanced OM decomposition during Holocene talik development. Furthermore, we found that drivers of CO2 and CH4 production differ following thaw. Our results suggest that GHG production from TOC-poor mineral deposits, which are widespread throughout the Arctic, can be substantial. Therefore, our novel data are relevant for vast ice-rich permafrost deposits vulnerable to thermokarst formation.

Methanogenic response to long-term permafrost thaw is determined by paleoenvironment.

Methane production in thawing permafrost can be substantial, yet often evolves after long lag phases or is even lacking. A central question is to which extent the production of methane after permafrost thaw is determined by the initial methanogenic community. We quantified the production of methane relative to carbon dioxide (CO2) and enumerated methanogenic (mcrA) gene copies in long-term (2-7 years) anoxic incubations at 4 °C using interglacial and glacial permafrost samples of Holocene and Pleistocene, including Eemian, origin. Changes in archaeal community composition were determined by sequencing of the archaeal 16S rRNA gene. Long-term thaw stimulated methanogenesis where methanogens initially dominated the archaeal community. Deposits of interstadial and interglacial (Eemian) origin, formed under higher temperatures and precipitation, displayed the greatest response to thaw. At the end of the incubations, a substantial shift in methanogenic community composition and a relative increase in hydrogenotrophic methanogens had occurred except for Eemian deposits in which a high abundance of potential acetoclastic methanogens were present. This study shows that only anaerobic CO2 production but not methane production correlates significantly with carbon and nitrogen content and that the methanogenic response to permafrost thaw is mainly constrained by the paleoenvironmental conditions during soil formation.

Gas production from dredged sediment.

The anaerobic degradation of sediment organic matter leads to considerable gas production in constructions made from sediments and in landfills where contaminated sediments are disposed of, inducing problems with the mechanical stability of constructions or necessitating extraction and treatment of gas. However, little is known about the magnitude of gas generation from dredged sediment, hence validated input parameters for gas production modelling are missing. On the occasion of drillings performed for the installation of inclinometers on a mono-landfill for contaminated dredged sediment, eleven waste layers of known were sampled. Samples were analysed for gas generation in a long-term laboratory incubation experiment carried out for 757 days. It was found that the residual gas potential of the deposited dredged material ranged between 2 and 12 m3 MgDW-1, relating to 3-11% of the organic matter being degraded. Correlation analyses with material properties suggest a strong role of nitrogen, either directly or as indicative parameter, with the gas potential increasing with total nitrogen content and the share of degradable carbon decreasing with increasing TOC/TN ratio. The by far greatest share of organic matter was bound in the heavy density fraction >1.4 g cm-3, suggesting that the readily available light organic matter pool had already been depleted during pre-treatment of the dredged sediment in dewatering fields and the subsequent years of landfilling. Consequently, the correlation of the remaining gas potential with heavy fraction nitrogen was even stronger than for bulk nitrogen. The gas potential as revealed from the long-term test correlated well with short-term values, but outreached the commonly applied potential measured for 21 days by the factor of four. The data improve the state of knowledge on gas production from the large mineral waste stream of dredged material and serve to improve gas production modelling for these types of wastes. The strong correlation of gas potential to TN suggests that TN may serve as a proxy to estimate total gas potential.

Influence of Anthropogenic Activities on Metals in Arctic Permafrost: A Characterization of Benchmark Soils on the Yamal and Gydan Peninsulas in Russia.

Permafrost-affected region in Russian Arctic is an important study area for investigating fate of trace metals in soils by geological processes and human-induced trace metals through atmospheric deposition. Two plots of soils in Yamal region were selected: Northern Trans-Urals area (PU1, PU2, PU3) adjacent to urban areas and Gydan Peninsula representing reference groups as natural landscapes (Yavai, Gyda, Enysei). The levels of most metals in Urals area were more than those in Gydan Peninsula. In soil profile, Histic horizon revealed the accumulation of most metals. Cd and Pb were classified as metals, which were transported by atmosphere from urban areas and accumulated in surficial organic layers. Gleying processes and cryogenic mass exchanges transported metals from bottom to top layers in mineral horizons. Moreover, gleying horizon functioned as a geochemical barrier for metal transporting below permafrost table. The levels of As, Mn, and Fe were obliviously higher than threshold limit values of Russian Siberia. However, these values cannot represent the natural hydromorphic soils in Arctic tundra. The Geoaccumulation Index (Igeo) were determined for assessing surface soil samples regarding to metals' pollution. The results suggested local geology pollution for Gydan Peninsula and atmospheric transport pollution for Urals area. More investigations with respect to trace metals behavior in permafrost-affected soil profile needed to be studied for understanding levels of trace metals with changes of active layer depth due to climate changing.

Anaerobic methanotrophic communities thrive in deep submarine permafrost.

Thawing submarine permafrost is a source of methane to the subsurface biosphere. Methane oxidation in submarine permafrost sediments has been proposed, but the responsible microorganisms remain uncharacterized. We analyzed archaeal communities and identified distinct anaerobic methanotrophic assemblages of marine and terrestrial origin (ANME-2a/b, ANME-2d) both in frozen and completely thawed submarine permafrost sediments. Besides archaea potentially involved in anaerobic oxidation of methane (AOM) we found a large diversity of archaea mainly belonging to Bathyarchaeota, Thaumarchaeota, and Euryarchaeota. Methane concentrations and δ13C-methane signatures distinguish horizons of potential AOM coupled either to sulfate reduction in a sulfate-methane transition zone (SMTZ) or to the reduction of other electron acceptors, such as iron, manganese or nitrate. Analysis of functional marker genes (mcrA) and fluorescence in situ hybridization (FISH) corroborate potential activity of AOM communities in submarine permafrost sediments at low temperatures. Modeled potential AOM consumes 72-100% of submarine permafrost methane and up to 1.2 Tg of carbon per year for the total expected area of submarine permafrost. This is comparable with AOM habitats such as cold seeps. We thus propose that AOM is active where submarine permafrost thaws, which should be included in global methane budgets.

Transformation of terrestrial organic matter along thermokarst-affected permafrost coasts in the Arctic.

The changing climate in the Arctic has a profound impact on permafrost coasts, which are subject to intensified thermokarst formation and erosion. Consequently, terrestrial organic matter (OM) is mobilized and transported into the nearshore zone. Yet, little is known about the fate of mobilized OM before and after entering the ocean. In this study we investigated a retrogressive thaw slump (RTS) on Qikiqtaruk - Herschel Island (Yukon coast, Canada). The RTS was classified into an undisturbed, a disturbed (thermokarst-affected) and a nearshore zone and sampled systematically along transects. Samples were analyzed for total and dissolved organic carbon and nitrogen (TOC, DOC, TN, DN), stable carbon isotopes (δ13C-TOC, δ13C-DOC), and dissolved inorganic nitrogen (DIN), which were compared between the zones. C/N-ratios, δ13C signatures, and ammonium (NH4-N) concentrations were used as indicators for OM degradation along with biomarkers (n-alkanes, n-fatty acids, n-alcohols). Our results show that OM significantly decreases after disturbance with a TOC and DOC loss of 77 and 55% and a TN and DN loss of 53 and 48%, respectively. C/N-ratios decrease significantly, whereas NH4-N concentrations slightly increase in freshly thawed material. In the nearshore zone, OM contents are comparable to the disturbed zone. We suggest that the strong decrease in OM is caused by initial dilution with melted massive ice and immediate offshore transport via the thaw stream. In the mudpool and thaw stream, OM is subject to degradation, whereas in the slump floor the nitrogen decrease is caused by recolonizing vegetation. Within the nearshore zone of the ocean, heavier portions of OM are directly buried in marine sediments close to shore. We conclude that RTS have profound impacts on coastal environments in the Arctic. They mobilize nutrients from permafrost, substantially decrease OM contents and provide fresh water and nutrients at a point source.

Circumpolar assessment of permafrost C quality and its vulnerability over time using long-term incubation data.

High-latitude ecosystems store approximately 1700 Pg of soil carbon (C), which is twice as much C as is currently contained in the atmosphere. Permafrost thaw and subsequent microbial decomposition of permafrost organic matter could add large amounts of C to the atmosphere, thereby influencing the global C cycle. The rates at which C is being released from the permafrost zone at different soil depths and across different physiographic regions are poorly understood but crucial in understanding future changes in permafrost C storage with climate change. We assessed the inherent decomposability of C from the permafrost zone by assembling a database of long-term (>1 year) aerobic soil incubations from 121 individual samples from 23 high-latitude ecosystems located across the northern circumpolar permafrost zone. Using a three-pool (i.e., fast, slow and passive) decomposition model, we estimated pool sizes for C fractions with different turnover times and their inherent decomposition rates using a reference temperature of 5 °C. Fast cycling C accounted for less than 5% of all C in both organic and mineral soils whereas the pool size of slow cycling C increased with C : N. Turnover time at 5 °C of fast cycling C typically was below 1 year, between 5 and 15 years for slow turning over C, and more than 500 years for passive C. We project that between 20 and 90% of the organic C could potentially be mineralized to CO2 within 50 incubation years at a constant temperature of 5 °C, with vulnerability to loss increasing in soils with higher C : N. These results demonstrate the variation in the vulnerability of C stored in permafrost soils based on inherent differences in organic matter decomposability, and point toward C : N as an index of decomposability that has the potential to be used to scale permafrost C loss across landscapes.

Predicting long-term carbon mineralization and trace gas production from thawing permafrost of Northeast Siberia.

The currently observed Arctic warming will increase permafrost degradation followed by mineralization of formerly frozen organic matter to carbon dioxide (CO2 ) and methane (CH4 ). Despite increasing awareness of permafrost carbon vulnerability, the potential long-term formation of trace gases from thawing permafrost remains unclear. The objective of the current study is to quantify the potential long-term release of trace gases from permafrost organic matter. Therefore, Holocene and Pleistocene permafrost deposits were sampled in the Lena River Delta, Northeast Siberia. The sampled permafrost contained between 0.6% and 12.4% organic carbon. CO2 and CH4 production was measured for 1200 days in aerobic and anaerobic incubations at 4 °C. The derived fluxes were used to estimate parameters of a two pool carbon degradation model. Total CO2 production was similar in Holocene permafrost (1.3 ± 0.8 mg CO2 -C gdw(-1) aerobically, 0.25 ± 0.13 mg CO2 -C gdw(-1) anaerobically) as in 34 000-42 000-year-old Pleistocene permafrost (1.6 ± 1.2 mg CO2 -C gdw(-1) aerobically, 0.26 ± 0.10 mg CO2 -C gdw(-1) anaerobically). The main predictor for carbon mineralization was the content of organic matter. Anaerobic conditions strongly reduced carbon mineralization since only 25% of aerobically mineralized carbon was released as CO2 and CH4 in the absence of oxygen. CH4 production was low or absent in most of the Pleistocene permafrost and always started after a significant delay. After 1200 days on average 3.1% of initial carbon was mineralized to CO2 under aerobic conditions while without oxygen 0.55% were released as CO2 and 0.28% as CH4 . The calibrated carbon degradation model predicted cumulative CO2 production over a period of 100 years accounting for 15.1% (aerobic) and 1.8% (anaerobic) of initial organic carbon, which is significantly less than recent estimates. The multiyear time series from the incubation experiments helps to more reliably constrain projections of future trace gas fluxes from thawing permafrost landscapes.

Regional, seasonal and interspecific variation in 15N and 13C in sympatric mouse lemurs.

Madagascar provides some of the rare examples where two or more primate species of the same genus and with seemingly identical niche requirements occur in sympatry. If congeneric primate species co-occur in other parts of the world, they differ in size in a way that is consistent with Hutchinson's rule for coexisting species, or they occupy different ecological niches. In some areas of Madagascar, mouse lemurs do not follow these "rules" and thus seem to violate one of the principles of community ecology. In order to understand the mechanisms that allow coexistence of sympatric congeneric species without obvious niche differentiation, we studied food composition of two identical sized omnivorous mouse lemur species, Microcebus griseorufus and M. murinus with the help of stable isotope analyses (δ(15)N and δ(13)C). The two species are closely related sister species. During the rich season, when food seems abundant, the two species do not differ in their nitrogen isotope composition, indicating that the two species occupy the same trophic level. But they differ in their δ(13)C values, indicating that M. griseorufus feeds more on C(4) and CAM (Crassulacean-acid-metabolism) plants than M. murinus. During the lean season, M. murinus has lower δ(15)N values, indicating that the two species feed at different trophic levels during times of food shortage. Hybrids between the two species showed intermediate food composition. The results reflect subtle differences in foraging or metabolic adaptations that are difficult to quantify by traditional observations but that represent possibilities to allow coexistence of species.

Methanogenic community composition and anaerobic carbon turnover in submarine permafrost sediments of the Siberian Laptev Sea.

The Siberian Laptev Sea shelf contains submarine permafrost, which was formed by flooding of terrestrial permafrost with ocean water during the Holocene sea level rise. This flooding resulted in a warming of the permafrost to temperatures close below 0 degrees C. The impact of these environmental changes on methanogenic communities and carbon dynamics in the permafrost was studied in a submarine permafrost core of the Siberian Laptev Sea shelf. Total organic carbon (TOC) content varied between 0.03% and 8.7% with highest values between 53 and 62 m depth below sea floor. In the same depth, maximum methane concentrations (284 nmol CH(4) g(-1)) and lowest carbon isotope values of methane (-72.2 per thousand VPDB) were measured, latter indicating microbial formation of methane under in situ conditions. The archaeal community structure was assessed by a nested polymerase chain reaction (PCR) amplification for DGGE, followed by sequencing of reamplified bands. Submarine permafrost samples showed a different archaeal community than the nearby terrestrial permafrost. Samples with high methane concentrations were dominated by sequences affiliated rather to the methylotrophic genera Methanosarcina and Methanococcoides as well as to uncultured archaea. The presented results give the first insights into the archaeal community in submarine permafrost and the first evidence for their activity at in situ conditions.

Desulfovibrio frigidus sp. nov. and Desulfovibrio ferrireducens sp. nov., psychrotolerant bacteria isolated from Arctic fjord sediments (Svalbard) with the ability to reduce Fe(III).

Strains 18T, 61T and 77 were isolated from two permanently cold fjord sediments on the west coast of Svalbard. The three psychrotolerant strains, with temperature optima at 20-23 degrees C, were able to grow at the freezing point of sea water, -2 degrees C. The strains oxidized important fermentation products such as hydrogen, formate and lactate with sulfate as the electron acceptor. Sulfate could be replaced by sulfite, thiosulfate or elemental sulfur. Poorly crystalline and soluble Fe(III) compounds were reduced in sulfate-free medium, but no growth occurred under these conditions. In the absence of electron acceptors, fermentative growth was possible. The pH optimum for the strains was around 7.1. The DNA G+C contents were 43.3 and 42.0 mol% for strains 18T and 61T, respectively. Strains 18T, 61T and 77 were most closely related to Desulfovibrio hydrothermalis (95.0-95.7 % 16S rRNA gene sequence similarity). Strains 18T and 77, exhibiting 99.9 % sequence similarity, represent a novel species for which the name Desulfovibrio frigidus sp. nov. is proposed. The type strain is strain 18T (=DSM 17176T = JCM 12924T). Strain 61T was closely related to strains 18T and 77 (97.6 and 97.5 % 16S rRNA gene sequence similarity), but on the basis of DNA-DNA hybridization strain 61T represents a novel species for which the name Desulfovibrio ferrireducens sp. nov. is proposed. The type strain is strain 61T (=DSM 16995T = JCM 12925T).

Desulfotomaculum arcticum sp. nov., a novel spore-forming, moderately thermophilic, sulfate-reducing bacterium isolated from a permanently cold fjord sediment of Svalbard.

Strain 15T is a novel spore-forming, sulfate-reducing bacterium isolated from a permanently cold fjord sediment of Svalbard. Sulfate could be replaced by sulfite or thiosulfate. Hydrogen, formate, lactate, propionate, butyrate, hexanoate, methanol, ethanol, propanol, butanol, pyruvate, malate, succinate, fumarate, proline, alanine and glycine were used as electron donors in the presence of sulfate. Growth occurred with pyruvate as sole substrate. Optimal growth was observed at pH 7.1-7.5 and concentrations of 1-1.5 % NaCl and 0.4 % MgCl2. Strain 15T grew between 26 and 46.5 degrees C and optimal growth occurred at 44 degrees C. Therefore, strain 15T apparently cannot grow at in situ temperatures of Arctic sediments from where it was isolated, and it was proposed that it was present in the sediment in the form of spores. The DNA G+C content was 48.9 mol%. Strain 15T was most closely related to Desulfotomaculum thermosapovorans MLF(T) (93.5 % 16S rRNA gene sequence similarity). Strain 15T represents a novel species, for which the name Desulfotomaculum arcticum sp. nov. is proposed. The type strain is strain 15T (=DSM 17038T = JCM 12923T).