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 e ect 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.High-latitude ecosystems store almost twice as much C in soils than what is contained in the atmosphere 7,8 . As the global climate warms, northern high latitudes are experiencing rapid increases in temperature 9 that have the potential to not only increase C emissions from previously frozen C in permafrost and the active layer 10 but also to indirectly affect the C cycle through changes in regional and local hydrology. Warmer temperatures increase thawing of icerich permafrost and the melting of ground ice, which causes the land surface to collapse into the space that was previously filled by ice resulting in thermokarst terrain 11 . Permafrost thawing can also gradually increase active layer thickness (seasonally thawed ground), causing poorly drained soil conditions in lowlands or drier conditions in uplands where natural drainage can increase 3 . On the other hand, permafrost thaw and collapse can cause soils to become waterlogged where anaerobic conditions prevail and C is released in the form of CO 2 and CH 4 . One major uncertainty in determining the climate forcing impact of permafrost ecosystems is understanding the relative magnitudes of the effects of shifting subsurface hydrology versus increasing temperatures on greenhouse gas release in permafrost ecosystems.In addition to soil temperature and moisture, the chemical composition (for example, carbon to nitrogen ratio) 12 , physical protection by soil minerals, microbial community dynamics, and other environmental controls, such as pH and nutrient availability, also impact the amount of C released to the atmosphere 13 . While temperature and soil moisture (that is, oxygen availability) a...

DNA is inherently limited by its four natural nucleotides. Efforts to expand the genetic alphabet, by addition of an unnatural base pair, promise to expand the biotechnological applications available for DNA as well as being an essential first step towards expansion of the genetic code. We have conducted two independent screens of hydrophobic unnatural nucleotides to identify novel candidate base pairs that are well recognized by a natural DNA polymerase. From a pool of 3600 candidate base pairs, both screens identified the same base pair, dSICS:dMMO2, which we report here. Using a series of related analogs, we performed a detailed structure-activity relationship analysis, which allowed us to identify the essential functional groups on each nucleobase. From the results of these studies, we designed an optimized base pair, d5SICS:dMMO2, which is efficiently and selectively synthesized by Kf within the context of natural DNA.

Rising temperatures in the Arctic can affect soil organic matter (SOM) decomposition directly and indirectly, by increasing plant primary production and thus the allocation of plant-derived organic compounds into the soil. Such compounds, for example root exudates or decaying fine roots, are easily available for microorganisms, and can alter the decomposition of older SOM (“priming effect”). We here report on a SOM priming experiment in the active layer of a permafrost soil from the central Siberian Arctic, comparing responses of organic topsoil, mineral subsoil, and cryoturbated subsoil material (i.e., poorly decomposed topsoil material subducted into the subsoil by freeze–thaw processes) to additions of 13C-labeled glucose, cellulose, a mixture of amino acids, and protein (added at levels corresponding to approximately 1% of soil organic carbon). SOM decomposition in the topsoil was barely affected by higher availability of organic compounds, whereas SOM decomposition in both subsoil horizons responded strongly. In the mineral subsoil, SOM decomposition increased by a factor of two to three after any substrate addition (glucose, cellulose, amino acids, protein), suggesting that the microbial decomposer community was limited in energy to break down more complex components of SOM. In the cryoturbated horizon, SOM decomposition increased by a factor of two after addition of amino acids or protein, but was not significantly affected by glucose or cellulose, indicating nitrogen rather than energy limitation. Since the stimulation of SOM decomposition in cryoturbated material was not connected to microbial growth or to a change in microbial community composition, the additional nitrogen was likely invested in the production of extracellular enzymes required for SOM decomposition. Our findings provide a first mechanistic understanding of priming in permafrost soils and suggest that an increase in the availability of organic carbon or nitrogen, e.g., by increased plant productivity, can change the decomposition of SOM stored in deeper layers of permafrost soils, with possible repercussions on the global climate.

Cryoturbation, the burial of topsoil material into deeper soil horizons by repeated freeze-thaw events, is an important storage mechanism for soil organic matter (SOM) in permafrost-affected soils. Besides abiotic conditions, microbial community structure and the accessibility of SOM to the decomposer community are hypothesized to control SOM decomposition and thus have a crucial role in SOM accumulation in buried soils. We surveyed the microbial community structure in cryoturbated soils from nine soil profiles in the northeastern Siberian tundra using high-throughput sequencing and quantification of bacterial, archaeal and fungal marker genes. We found that bacterial abundances in buried topsoils were as high as in unburied topsoils. In contrast, fungal abundances decreased with depth and were significantly lower in buried than in unburied topsoils resulting in remarkably low fungal to bacterial ratios in buried topsoils. Fungal community profiling revealed an associated decrease in presumably ectomycorrhizal (ECM) fungi. The abiotic conditions (low to subzero temperatures, anoxia) and the reduced abundance of fungi likely provide a niche for bacterial, facultative anaerobic decomposers of SOM such as members of the Actinobacteria, which were found in significantly higher relative abundances in buried than in unburied topsoils. Our study expands the knowledge on the microbial community structure in soils of Northern latitude permafrost regions, and attributes the delayed decomposition of SOM in buried soils to specific microbial taxa, and particularly to a decrease in abundance and activity of ECM fungi, and to the extent to which bacterial decomposers are able to act as their functional substitutes.

Most heterotrophic organisms feed on substrates that are poor in nutrients compared to their demand, leading to elemental imbalances that may constrain their growth and function. Flexible carbon (C)-use efficiency (CUE, C used for growth over C taken up) can represent a strategy to reduce elemental imbalances. Here, we argue that metabolic regulation has evolved to maximise the organism growth rate along gradients of nutrient availability and translated this assumption into an optimality model that links CUE to substrate and organism stoichiometry. The optimal CUE is predicted to decrease with increasing substrate C-to-nutrient ratio, and increase with nutrient amendment. These predictions are generally confirmed by empirical evidence from a new database of c. 2200 CUE estimates, lending support to the hypothesis that CUE is optimised across levels of organisation (microorganisms and animals), in aquatic and terrestrial systems, and when considering nitrogen or phosphorus as limiting nutrients.