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A multi-omics approach, integrating metagenomics, metatranscriptomics and metaproteomics, determines the phylogenetic composition of the microbial community and assesses its functional potential and activity along a thaw transition from intact permafrost to thermokast bog. Multi-omics survey of frozen-soil microbiomes The application of the various individual 'omics' tools to the study of microbial ecosystems has dramatically altered our view of their constituents and ecology over the past decade. Here Janet Jansson and colleagues develop an multi-omics approach, integrating metagenomics, metatranscriptomics and metaproteomics to analyse microbial gene expression in frozen soils that form part of the Alaska Peatland Experiment. The results show that the community shifts along a natural thaw gradient from permafrost to seasonally thawed active layer to thermokarst bog and the authors find that there is a transition in the potential for several biogeochemical cycles with thaw, including those for denitrification, nitrate reduction, iron reduction and methane oxidation. Over 20% of Earth’s terrestrial surface is underlain by permafrost with vast stores of carbon that, once thawed, may represent the largest future transfer of carbon from the biosphere to the atmosphere 1 . This process is largely dependent on microbial responses, but we know little about microbial activity in intact, let alone in thawing, permafrost. Molecular approaches have recently revealed the identities and functional gene composition of microorganisms in some permafrost soils 2 , 3 , 4 and a rapid shift in functional gene composition during short-term thaw experiments 3 . However, the fate of permafrost carbon depends on climatic, hydrological and microbial responses to thaw at decadal scales 5 , 6 . Here we use the combination of several molecular ‘omics’ approaches to determine the phylogenetic composition of the microbial communities, including several draft genomes of novel species, their functional potential and activity in soils representing different states of thaw: intact permafrost, seasonally thawed active layer and thermokarst bog. The multi-omics strategy reveals a good correlation of process rates to omics data for dominant processes, such as methanogenesis in the bog, as well as novel survival strategies for potentially active microbes in permafrost.

Permafrost microbes await the thaw Permafrost soil in the Arctic contains a huge reservoir of carbon. If the climate warms and the permafrost thaws, this carbon would become accessible to microbial degradation, releasing greenhouse gases in the process. The microbes responsible for this process are largely unknown. Metagenomic analysis of DNA isolated from two permafrost soils collected in Alaska reveals a rapid microbial response to thawing, with many functional gene abundances increasing. A draft genome of a novel methanogen was constructed from the sequence data. This study highlights the importance of rapid cycling of methane and nitrogen in thawing permafrost. Permafrost contains an estimated 1672 Pg carbon (C), an amount roughly equivalent to the total currently contained within land plants and the atmosphere 1 , 2 , 3 . This reservoir of C is vulnerable to decomposition as rising global temperatures cause the permafrost to thaw 2 . During thaw, trapped organic matter may become more accessible for microbial degradation and result in greenhouse gas emissions 4 , 5 . Despite recent advances in the use of molecular tools to study permafrost microbial communities 6 , 7 , 8 , 9 , their response to thaw remains unclear. Here we use deep metagenomic sequencing to determine the impact of thaw on microbial phylogenetic and functional genes, and relate these data to measurements of methane emissions. Metagenomics, the direct sequencing of DNA from the environment, allows the examination of whole biochemical pathways and associated processes, as opposed to individual pieces of the metabolic puzzle. Our metagenome analyses reveal that during transition from a frozen to a thawed state there are rapid shifts in many microbial, phylogenetic and functional gene abundances and pathways. After one week of incubation at 5 °C, permafrost metagenomes converge to be more similar to each other than while they are frozen. We find that multiple genes involved in cycling of C and nitrogen shift rapidly during thaw. We also construct the first draft genome from a complex soil metagenome, which corresponds to a novel methanogen. Methane previously accumulated in permafrost is released during thaw and subsequently consumed by methanotrophic bacteria. Together these data point towards the importance of rapid cycling of methane and nitrogen in thawing permafrost.

In permafrost (perennially frozen ground) microbes survive oligotrophic conditions, sub-zero temperatures, low water availability and high salinity over millennia. Viable life exists in permafrost tens of thousands of years old but we know little about the metabolic and physiological adaptations to the challenges presented by life in frozen ground over geologic time. In this study we asked whether increasing age and the associated stressors drive adaptive changes in community composition and function. We conducted deep metagenomic and 16 S rRNA gene sequencing across a Pleistocene permafrost chronosequence from 19 000 to 33 000 years before present (kyr). We found that age markedly affected community composition and reduced diversity. Reconstruction of paleovegetation from metagenomic sequence suggests vegetation differences in the paleo record are not responsible for shifts in community composition and function. Rather, we observed shifts consistent with long-term survival strategies in extreme cryogenic environments. These include increased reliance on scavenging detrital biomass, horizontal gene transfer, chemotaxis, dormancy, environmental sensing and stress response. Our results identify traits that may enable survival in ancient cryoenvironments with no influx of energy or new materials.

A major goal of microbial ecology is to identify links between microbial community structure and microbial processes. Although this objective seems straightforward, there are conceptual and methodological challenges to designing studies that explicitly evaluate this link. Here, we analyzed literature documenting structure and process responses to manipulations to determine the frequency of structure-process links and whether experimental approaches and techniques influence link detection. We examined nine journals (published 2009–13) and retained 148 experimental studies measuring microbial community structure and processes. Many qualifying papers (112 of 148) documented structure and process responses, but few (38 of 112 papers) reported statistically testing for a link. Of these tested links, 75% were significant and typically used Spearman or Pearson's correlation analysis (68%). No particular approach for characterizing structure or processes was more likely to produce significant links. Process responses were detected earlier on average than responses in structure or both structure and process. Together, our findings suggest that few publications report statistically testing structure-process links. However, when links are tested for they often occur but share few commonalities in the processes or structures that were linked and the techniques used for measuring them. Few publications reported statistically testing microbial community structure-process links; 75% of tested links were significant, though had few commonalities in which processes or structures were measured and the techniques used.

For any enzyme-catalyzed reaction to occur, the corresponding protein-encoding genes and transcripts are necessary prerequisites. Thus, a positive relationship between the abundance of gene or transcripts and corresponding process rates is often assumed. To test this assumption, we conducted a meta-analysis of the relationships between gene and/or transcript abundances and corresponding process rates. We identified 415 studies that quantified the abundance of genes or transcripts for enzymes involved in carbon or nitrogen cycling. However, in only 59 of these manuscripts did the authors report both gene or transcript abundance and rates of the appropriate process. We found that within studies there was a significant but weak positive relationship between gene abundance and the corresponding process. Correlations were not strengthened by accounting for habitat type, differences among genes or reaction products versus reactants, suggesting that other ecological and methodological factors may affect the strength of this relationship. Our findings highlight the need for fundamental research on the factors that control transcription, translation and enzyme function in natural systems to better link genomic and transcriptomic data to ecosystem processes.

Permafrost underlies approximately one quarter of Northern Hemisphere terrestrial surfaces and contains 25–50% of the global soil carbon (C) pool. Permafrost soils and the C stocks within are vulnerable to ongoing and future projected climate warming. The biogeography of microbial communities inhabiting permafrost has not been examined beyond a small number of sites focused on local-scale variation. Permafrost is different from other soils. Perennially frozen conditions in permafrost dictate that microbial communities do not turn over quickly, thus possibly providing strong linkages to past environments. Thus, the factors structuring the composition and function of microbial communities may differ from patterns observed in other terrestrial environments. Here, we analyzed 133 permafrost metagenomes from North America, Europe, and Asia. Permafrost biodiversity and taxonomic distribution varied in relation to pH, latitude and soil depth. The distribution of genes differed by latitude, soil depth, age, and pH. Genes that were the most highly variable across all sites were associated with energy metabolism and C-assimilation. Specifically, methanogenesis, fermentation, nitrate reduction, and replenishment of citric acid cycle intermediates. This suggests that adaptations to energy acquisition and substrate availability are among some of the strongest selective pressures shaping permafrost microbial communities. The spatial variation in metabolic potential has primed communities for specific biogeochemical processes as soils thaw due to climate change, which could cause regional- to global- scale variation in C and nitrogen processing and greenhouse gas emissions.

A meta-analysis of soil incubation studies from the permafrost zone suggests that thawing under aerobic conditions, which releases CO 2 , will strengthen the permafrost carbon feedback more than waterlogged systems, which releases CO 2 and CH 4 . Increasing temperatures in northern high latitudes are causing permafrost to thaw 1 , making large amounts of previously frozen organic matter vulnerable to microbial decomposition 2 . Permafrost thaw also creates a fragmented landscape of drier and wetter soil conditions 3 , 4 that determine the amount and form (carbon dioxide (CO 2 ), or methane (CH 4 )) of carbon (C) released to the atmosphere. The rate and form of C release control the magnitude of the permafrost C feedback, so their relative contribution with a warming climate remains unclear 5 , 6 . We quantified the effect of increasing temperature and changes from aerobic to anaerobic soil conditions using 25 soil incubation studies from the permafrost zone. Here we show, using two separate meta-analyses, that a 10 °C increase in incubation temperature increased C release by a factor of 2.0 (95% confidence interval (CI), 1.8 to 2.2). Under aerobic incubation conditions, soils released 3.4 (95% CI, 2.2 to 5.2) times more C than under anaerobic conditions. Even when accounting for the higher heat trapping capacity of CH 4 , soils released 2.3 (95% CI, 1.5 to 3.4) times more C under aerobic conditions. These results imply that permafrost ecosystems thawing under aerobic conditions and releasing CO 2 will strengthen the permafrost C feedback more than waterlogged systems releasing CO 2 and CH 4 for a given amount of C.

Permafrost (perennially frozen) soils store vast amounts of organic carbon (C) and nitrogen (N) that are vulnerable to mobilization as dissolved organic carbon (DOC) and dissolved organic and inorganic nitrogen (DON, DIN) upon thaw. Such releases will affect the biogeochemistry of permafrost regions, yet little is known about the chemical composition and source variability of active-layer (seasonally frozen) and permafrost soil DOC, DON and DIN. We quantified DOC, total dissolved N (TDN), DON, and DIN leachate yields from deep active-layer and near-surface boreal Holocene permafrost soils in interior Alaska varying in soil C and N content and radiocarbon age to determine potential release upon thaw. Soil cores were collected at three sites distributed across the Alaska boreal region in late winter, cut in 15 cm thick sections, and deep active-layer and shallow permafrost sections were thawed and leached. Leachates were analyzed for DOC, TDN, nitrate (NO3−), and ammonium (NH4+) concentrations, dissolved organic matter optical properties, and DOC biodegradability. Soils were analyzed for C, N, and radiocarbon (14C) content. Soil DOC, TDN, DON, and DIN yields increased linearly with soil C and N content, and decreased with increasing radiocarbon age. These relationships were significantly different for active-layer and permafrost soils such that for a given soil C or N content, or radiocarbon age, permafrost soils released more DOC and TDN (mostly as DON) per gram soil than active-layer soils. Permafrost soil DOC biodegradability was significantly correlated with soil Δ14C and DOM optical properties. Our results demonstrate that near-surface Holocene permafrost soils preserve greater relative potential DOC and TDN yields than overlying seasonally frozen soils that are exposed to annual leaching and decomposition. While many factors control the fate of DOC and TDN, the greater relative yields from newly thawed Holocene permafrost soils will have the largest potential impact in areas dominated by organic-rich soils.

Permafrost soils are large reservoirs of potentially labile carbon (C). Understanding the dynamics of C release from these soils requires us to account for the impact of wildfires, which are increasing in frequency as the climate changes. Boreal wildfires contribute to global emission of greenhouse gases (GHG—CO 2 , CH 4 and N 2 O) and indirectly result in the thawing of near-surface permafrost. In this study, we aimed to define the impact of fire on soil microbial communities and metabolic potential for GHG fluxes in samples collected up to 1 m depth from an upland black spruce forest near Nome Creek, Alaska. We measured geochemistry, GHG fluxes, potential soil enzyme activities and microbial community structure via 16SrRNA gene and metagenome sequencing. We found that soil moisture, C content and the potential for respiration were reduced by fire, as were microbial community diversity and metabolic potential. There were shifts in dominance of several microbial community members, including a higher abundance of candidate phylum AD3 after fire. The metagenome data showed that fire had a pervasive impact on genes involved in carbohydrate metabolism, methanogenesis and the nitrogen cycle. Although fire resulted in an immediate release of CO 2 from surface soils, our results suggest that the potential for emission of GHG was ultimately reduced at all soil depths over the longer term. Because of the size of the permafrost C reservoir, these results are crucial for understanding whether fire produces a positive or negative feedback loop contributing to the global C cycle.

Background Winter carbon loss in northern ecosystems is estimated to be greater than the average growing season carbon uptake and is primarily driven by microbial decomposers. Viruses modulate microbial carbon cycling via induced mortality and metabolic controls, but it is unknown whether viruses are active under winter conditions (anoxic and sub-freezing temperatures). Results We used stable isotope probing (SIP) targeted metagenomics to reveal the genomic potential of active soil microbial populations under simulated winter conditions, with an emphasis on viruses and virus-host dynamics. Arctic peat soils from the Bonanza Creek Long-Term Ecological Research site in Alaska were incubated under sub-freezing anoxic conditions with H 2 18 O or natural abundance water for 184 and 370 days. We sequenced 23 SIP-metagenomes and measured carbon dioxide (CO 2 ) efflux throughout the experiment. We identified 46 bacterial populations (spanning 9 phyla) and 243 viral populations that actively took up 18 O in soil and respired CO 2 throughout the incubation. Active bacterial populations represented only a small portion of the detected microbial community and were capable of fermentation and organic matter degradation. In contrast, active viral populations represented a large portion of the detected viral community and one third were linked to active bacterial populations. We identified 86 auxiliary metabolic genes and other environmentally relevant genes. The majority of these genes were carried by active viral populations and had diverse functions such as carbon utilization and scavenging that could provide their host with a fitness advantage for utilizing much-needed carbon sources or acquiring essential nutrients. Conclusions Overall, there was a stark difference in the identity and function of the active bacterial and viral community compared to the unlabeled community that would have been overlooked with a non-targeted standard metagenomic analysis. Our results illustrate that substantial active virus-host interactions occur in sub-freezing anoxic conditions and highlight viruses as a major community-structuring agent that likely modulates carbon loss in peat soils during winter, which may be pivotal for understanding the future fate of arctic soils' vast carbon stocks. 7CT7oo5s6pZcfh9yAbPeef Video abstract