Tree planting in organic soils does not result in net carbon sequestration on decadal timescales.

Tree planting is increasingly being proposed as a strategy to combat climate change through carbon (C) sequestration in tree biomass. However, total ecosystem C storage that includes soil organic C (SOC) must be considered to determine whether planting trees for climate change mitigation results in increased C storage. We show that planting two native tree species (Betula pubescens and Pinus sylvestris), of widespread Eurasian distribution, onto heather (Calluna vulgaris) moorland with podzolic and peaty podzolic soils in Scotland, did not lead to an increase in net ecosystem C stock 12 or 39 years after planting. Plots with trees had greater soil respiration and lower SOC in organic soil horizons than heather control plots. The decline in SOC cancelled out the increment in C stocks in tree biomass on decadal timescales. At all four experimental sites sampled, there was no net gain in ecosystem C stocks 12-39 years after afforestation-indeed we found a net ecosystem C loss in one of four sites with deciduous B. pubescens stands; no net gain in ecosystem C at three sites planted with B. pubescens; and no net gain at additional stands of P. sylvestris. We hypothesize that altered mycorrhizal communities and autotrophic C inputs have led to positive 'priming' of soil organic matter, resulting in SOC loss, constraining the benefits of tree planting for ecosystem C sequestration. The results are of direct relevance to current policies, which promote tree planting on the assumption that this will increase net ecosystem C storage and contribute to climate change mitigation. Ecosystem-level biogeochemistry and C fluxes must be better quantified and understood before we can be assured that large-scale tree planting in regions with considerable pre-existing SOC stocks will have the intended policy and climate change mitigation outcomes.

Rhizosphere allocation by canopy-forming species dominates soil CO2 efflux in a subarctic landscape.

In arctic ecosystems, climate change has increased plant productivity. As arctic carbon (C) stocks predominantly are located belowground, the effects of greater plant productivity on soil C storage will significantly determine the net sink/source potential of these ecosystems, but vegetation controls on soil CO2 efflux remain poorly resolved. In order to identify the role of canopy-forming species in belowground C dynamics, we conducted a girdling experiment with plots distributed across 1 km2 of treeline birch (Betula pubescens) forest and willow (Salix lapponum) patches in northern Sweden and quantified the contribution of canopy vegetation to soil CO2 fluxes and belowground productivity. Girdling birches reduced total soil CO2 efflux in the peak growing season by 53%, which is double the expected amount, given that trees contribute only half of the total leaf area in the forest. Root and mycorrhizal mycelial production also decreased substantially. At peak season, willow shrubs contributed 38% to soil CO2 efflux in their patches. Our findings indicate that C, recently fixed by trees and tall shrubs, makes a substantial contribution to soil respiration. It is critically important that these processes are taken into consideration in the context of a greening arctic because productivity and ecosystem C sequestration are not synonymous.

Exploring drivers of litter decomposition in a greening Arctic: results from a transplant experiment across a treeline.

Decomposition of plant litter is a key control over carbon (C) storage in the soil. The biochemistry of the litter being produced, the environment in which the decomposition is taking place, and the community composition and metabolism of the decomposer organisms exert a combined influence over decomposition rates. As deciduous shrubs and trees are expanding into tundra ecosystems as a result of regional climate warming, this change in vegetation represents a change in litter input to tundra soils and a change in the environment in which litter decomposes. To test the importance of litter biochemistry and environment in determining litter mass loss, we reciprocally transplanted litter between heath (Empetrum nigrum), shrub (Betula nana), and forest (Betula pubescens) at a sub-Arctic treeline in Sweden. As expansion of shrubs and trees promotes deeper snow, we also used a snow fence experiment in a tundra heath environment to understand the importance of snow depth, relative to other factors, in the decomposition of litter. Our results show that B. pubescens and B. nana leaf litter decomposed at faster rates than E. nigrum litter across all environments, while all litter species decomposed at faster rates in the forest and shrub environments than in the tundra heath. The effect of increased snow on decomposition was minimal, leading us to conclude that microbial activity over summer in the productive forest and shrub vegetation is driving increased mass loss compared to the heath. Using B. pubescens and E. nigrum litter, we demonstrate that degradation of carbohydrate-C is a significant driver of mass loss in the forest. This pathway was less prominent in the heath, which is consistent with observations that tundra soils typically have high concentrations of "labile" C. This experiment suggests that further expansion of shrubs and trees may stimulate the loss of undecomposed carbohydrate C in the tundra.

Using stable isotopes to estimate travel times in a data-sparse Arctic catchment: Challenges and possible solutions.

Use of isotopes to quantify the temporal dynamics of the transformation of precipitation into run-off has revealed fundamental new insights into catchment flow paths and mixing processes that influence biogeochemical transport. However, catchments underlain by permafrost have received little attention in isotope-based studies, despite their global importance in terms of rapid environmental change. These high-latitude regions offer limited access for data collection during critical periods (e.g., early phases of snowmelt). Additionally, spatio-temporal variable freeze-thaw cycles, together with the development of an active layer, have a time variant influence on catchment hydrology. All of these characteristics make the application of traditional transit time estimation approaches challenging. We describe an isotope-based study undertaken to provide a preliminary assessment of travel times at Siksik Creek in the western Canadian Arctic. We adopted a model-data fusion approach to estimate the volumes and isotopic characteristics of snowpack and meltwater. Using samples collected in the spring/summer, we characterize the isotopic composition of summer rainfall, melt from snow, soil water, and stream water. In addition, soil moisture dynamics and the temporal evolution of the active layer profile were monitored. First approximations of transit times were estimated for soil and streamwater compositions using lumped convolution integral models and temporally variable inputs including snowmelt, ice thaw, and summer rainfall. Comparing transit time estimates using a variety of inputs revealed that transit time was best estimated using all available inflows (i.e., snowmelt, soil ice thaw, and rainfall). Early spring transit times were short, dominated by snowmelt and soil ice thaw and limited catchment storage when soils are predominantly frozen. However, significant and increasing mixing with water in the active layer during the summer resulted in more damped steam water variation and longer mean travel times (~1.5 years). The study has also highlighted key data needs to better constrain travel time estimates in permafrost catchments.

Draft Genome Sequence of Methylocella silvestris TVC, a Facultative Methanotroph Isolated from Permafrost.

Permafrost environments play a crucial role in global carbon and methane cycling. We report here the draft genome sequence of Methylocella silvestris TVC, a new facultative methanotroph strain, isolated from the Siksik Creek catchment in the continuous permafrost zone of Inuvik (Northwest Territories, Canada).

Slowed Biogeochemical Cycling in Sub-arctic Birch Forest Linked to Reduced Mycorrhizal Growth and Community Change after a Defoliation Event.

Sub-arctic birch forests (Betula pubescens Ehrh. ssp. czerepanovii) periodically suffer large-scale defoliation events caused by the caterpillars of the geometrid moths Epirrita autumnata and Operophtera brumata. Despite their obvious influence on ecosystem primary productivity, little is known about how the associated reduction in belowground C allocation affects soil processes. We quantified the soil response following a natural defoliation event in sub-arctic Sweden by measuring soil respiration, nitrogen availability and ectomycorrhizal fungi (EMF) hyphal production and root tip community composition. There was a reduction in soil respiration and an accumulation of soil inorganic N in defoliated plots, symptomatic of a slowdown of soil processes. This coincided with a reduction of EMF hyphal production and a shift in the EMF community to lower autotrophic C-demanding lineages (for example, /russula-lactarius). We show that microbial and nutrient cycling processes shift to a slower, less C-demanding state in response to canopy defoliation. We speculate that, amongst other factors, a reduction in the potential of EMF biomass to immobilise excess mineral nitrogen resulted in its build-up in the soil. These defoliation events are becoming more geographically widespread with climate warming, and could result in a fundamental shift in sub-arctic ecosystem processes and properties. EMF fungi may be important in mediating the response of soil cycles to defoliation and their role merits further investigation.

The Role of Microbial Community Composition in Controlling Soil Respiration Responses to Temperature.

Rising global temperatures may increase the rates of soil organic matter decomposition by heterotrophic microorganisms, potentially accelerating climate change further by releasing additional carbon dioxide (CO2) to the atmosphere. However, the possibility that microbial community responses to prolonged warming may modify the temperature sensitivity of soil respiration creates large uncertainty in the strength of this positive feedback. Both compensatory responses (decreasing temperature sensitivity of soil respiration in the long-term) and enhancing responses (increasing temperature sensitivity) have been reported, but the mechanisms underlying these responses are poorly understood. In this study, microbial biomass, community structure and the activities of dehydrogenase and ß-glucosidase enzymes were determined for 18 soils that had previously demonstrated either no response or varying magnitude of enhancing or compensatory responses of temperature sensitivity of heterotrophic microbial respiration to prolonged cooling. The soil cooling approach, in contrast to warming experiments, discriminates between microbial community responses and the consequences of substrate depletion, by minimising changes in substrate availability. The initial microbial community composition, determined by molecular analysis of soils showing contrasting respiration responses to cooling, provided evidence that the magnitude of enhancing responses was partly related to microbial community composition. There was also evidence that higher relative abundance of saprophytic Basidiomycota may explain the compensatory response observed in one soil, but neither microbial biomass nor enzymatic capacity were significantly affected by cooling. Our findings emphasise the key importance of soil microbial community responses for feedbacks to global change, but also highlight important areas where our understanding remains limited.

Biogeochemistry of "pristine" freshwater stream and lake systems in the western Canadian Arctic.

Climate change poses a substantial threat to the stability of the Arctic terrestrial carbon (C) pool as warmer air temperatures thaw permafrost and deepen the seasonally-thawed active layer of soils and sediments. Enhanced water flow through this layer may accelerate the transport of C and major cations and anions to streams and lakes. These act as important conduits and reactors for dissolved C within the terrestrial C cycle. It is important for studies to consider these processes in small headwater catchments, which have been identified as hotspots of rapid mineralisation of C sourced from ancient permafrost thaw. In order to better understand the role of inland waters in terrestrial C cycling we characterised the biogeochemistry of the freshwater systems in a c. 14 km2 study area in the western Canadian Arctic. Sampling took place during the snow-free seasons of 2013 and 2014 for major inorganic solutes, dissolved organic and inorganic C (DOC and DIC, respectively), carbon dioxide (CO2) and methane (CH4) concentrations from three water type groups: lakes, polygonal pools and streams. These groups displayed differing biogeochemical signatures, indicative of contrasting biogeochemical controls. However, none of the groups showed strong signals of enhanced permafrost thaw during the study seasons. The mean annual air temperature in the region has increased by more than 2.5 °C since 1970, and continued warming will likely affect the aquatic biogeochemistry. This study provides important baseline data for comparison with future studies in a warming Arctic.

Quantifying landscape-level methane fluxes in subarctic Finland using a multiscale approach.

Quantifying landscape-scale methane (CH4 ) fluxes from boreal and arctic regions, and determining how they are controlled, is critical for predicting the magnitude of any CH4 emission feedback to climate change. Furthermore, there remains uncertainty regarding the relative importance of small areas of strong methanogenic activity, vs. larger areas with net CH4 uptake, in controlling landscape-level fluxes. We measured CH4 fluxes from multiple microtopographical subunits (sedge-dominated lawns, interhummocks and hummocks) within an aapa mire in subarctic Finland, as well as in drier ecosystems present in the wider landscape, lichen heath and mountain birch forest. An intercomparison was carried out between fluxes measured using static chambers, up-scaled using a high-resolution landcover map derived from aerial photography and eddy covariance. Strong agreement was observed between the two methodologies, with emission rates greatest in lawns. CH4 fluxes from lawns were strongly related to seasonal fluctuations in temperature, but their floating nature meant that water-table depth was not a key factor in controlling CH4 release. In contrast, chamber measurements identified net CH4 uptake in birch forest soils. An intercomparison between the aerial photography and satellite remote sensing demonstrated that quantifying the distribution of the key CH4 emitting and consuming plant communities was possible from satellite, allowing fluxes to be scaled up to a 100 km(2) area. For the full growing season (May to October), ~ 1.1-1.4 g CH4  m(-2) was released across the 100 km(2) area. This was based on up-scaled lawn emissions of 1.2-1.5 g CH4  m(-2) , vs. an up-scaled uptake of 0.07-0.15 g CH4  m(-2) by the wider landscape. Given the strong temperature sensitivity of the dominant lawn fluxes, and the fact that lawns are unlikely to dry out, climate warming may substantially increase CH4 emissions in northern Finland, and in aapa mire regions in general.

Rapid carbon turnover beneath shrub and tree vegetation is associated with low soil carbon stocks at a subarctic treeline.

Climate warming at high northern latitudes has caused substantial increases in plant productivity of tundra vegetation and an expansion of the range of deciduous shrub species. However significant the increase in carbon (C) contained within above-ground shrub biomass, it is modest in comparison with the amount of C stored in the soil in tundra ecosystems. Here, we use a 'space-for-time' approach to test the hypothesis that a shift from lower-productivity tundra heath to higher-productivity deciduous shrub vegetation in the sub-Arctic may lead to a loss of soil C that out-weighs the increase in above-ground shrub biomass. We further hypothesize that a shift from ericoid to ectomycorrhizal systems coincident with this vegetation change provides a mechanism for the loss of soil C. We sampled soil C stocks, soil surface CO2 flux rates and fungal growth rates along replicated natural transitions from birch forest (Betula pubescens), through deciduous shrub tundra (Betula nana) to tundra heaths (Empetrum nigrum) near Abisko, Swedish Lapland. We demonstrate that organic horizon soil organic C (SOCorg ) is significantly lower at shrub (2.98 ± 0.48 kg m(-2) ) and forest (2.04 ± 0.25 kg m(-2) ) plots than at heath plots (7.03 ± 0.79 kg m(-2) ). Shrub vegetation had the highest respiration rates, suggesting that despite higher rates of C assimilation, C turnover was also very high and less C is sequestered in the ecosystem. Growth rates of fungal hyphae increased across the transition from heath to shrub, suggesting that the action of ectomycorrhizal symbionts in the scavenging of organically bound nutrients is an important pathway by which soil C is made available to microbial degradation. The expansion of deciduous shrubs onto potentially vulnerable arctic soils with large stores of C could therefore represent a significant positive feedback to the climate system.

Temperature sensitivity of soil respiration rates enhanced by microbial community response.

Soils store about four times as much carbon as plant biomass, and soil microbial respiration releases about 60 petagrams of carbon per year to the atmosphere as carbon dioxide. Short-term experiments have shown that soil microbial respiration increases exponentially with temperature. This information has been incorporated into soil carbon and Earth-system models, which suggest that warming-induced increases in carbon dioxide release from soils represent an important positive feedback loop that could influence twenty-first-century climate change. The magnitude of this feedback remains uncertain, however, not least because the response of soil microbial communities to changing temperatures has the potential to either decrease or increase warming-induced carbon losses substantially. Here we collect soils from different ecosystems along a climate gradient from the Arctic to the Amazon and investigate how microbial community-level responses control the temperature sensitivity of soil respiration. We find that the microbial community-level response more often enhances than reduces the mid- to long-term (90 days) temperature sensitivity of respiration. Furthermore, the strongest enhancing responses were observed in soils with high carbon-to-nitrogen ratios and in soils from cold climatic regions. After 90 days, microbial community responses increased the temperature sensitivity of respiration in high-latitude soils by a factor of 1.4 compared to the instantaneous temperature response. This suggests that the substantial carbon stores in Arctic and boreal soils could be more vulnerable to climate warming than currently predicted.

Global assessment of experimental climate warming on tundra vegetation: heterogeneity over space and time.

Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

Ecological dynamics across the Arctic associated with recent climate change.

At the close of the Fourth International Polar Year, we take stock of the ecological consequences of recent climate change in the Arctic, focusing on effects at population, community, and ecosystem scales. Despite the buffering effect of landscape heterogeneity, Arctic ecosystems and the trophic relationships that structure them have been severely perturbed. These rapid changes may be a bellwether of changes to come at lower latitudes and have the potential to affect ecosystem services related to natural resources, food production, climate regulation, and cultural integrity. We highlight areas of ecological research that deserve priority as the Arctic continues to warm.

No evidence for compensatory thermal adaptation of soil microbial respiration in the study of Bradford et al. (2008).

Bradford et al. (2008) conclude that thermal adaptation will reduce the response of soil microbial respiration to rising global temperatures. However, we question both the methods used to calculate mass-specific respiration rates and the interpretation of the results. No clear evidence of thermal adaptation reducing soil microbial activity was produced.

The impact of climate change on ecosystem carbon dynamics at the Scandinavian mountain birch forest-tundra heath ecotone.

Changes in temperature and moisture resulting from climate change are likely to strongly modify the ecosystem carbon sequestration capacity in high-latitude areas, both through vegetation shifts and via direct warming effects on photosynthesis and decomposition. This paper offers a synthesis of research addressing the potential impacts of climate warming on soil processes and carbon fluxes at the forest-tundra ecotone in Scandinavia. Our results demonstrated higher rates of organic matter decomposition in mountain birch forest than in tundra heath soils, with markedly shallower organic matter horizons in the forest. Field and laboratory experiments suggest that increased temperatures are likely to increase CO2 efflux from both tundra and forest soil providing moisture availability does not become limiting for the decomposition process. Furthermore, colonization of tundra heath by mountain birch forest would increase rates of decomposition, and thus CO2 emissions, from the tundra heath soils, which currently store substantial amounts of potentially labile carbon. Mesic soils underlying both forest and tundra heath are currently weak sinks of atmospheric methane, but the strength of this sink could be increased with climate warming and/or drying.

Soil microbial respiration in arctic soil does not acclimate to temperature.

Warming-induced release of CO2 from the large carbon (C) stores in arctic soils could accelerate climate change. However, declines in the response of soil respiration to warming in long-term experiments suggest that microbial activity acclimates to temperature, greatly reducing the potential for enhanced C losses. As reduced respiration rates with time could be equally caused by substrate depletion, evidence for thermal acclimation remains controversial. To overcome this problem, we carried out a cooling experiment with soils from arctic Sweden. If acclimation causes the reduction in soil respiration observed after experimental warming, then it should subsequently lead to an increase in respiration rates after cooling. We demonstrate that thermal acclimation did not occur following cooling. Rather, during the 90 days after cooling, a further reduction in the soil respiration rate was observed, which was only reversed by extended re-exposure to warmer temperatures. We conclude that over the time scale of a few weeks to months, warming-induced changes in the microbial community in arctic soils will amplify the instantaneous increase in the rates of CO2 production and thus enhance C losses potentially accelerating the rate of 21st century climate change.

Plant community responses to experimental warming across the tundra biome.

Recent observations of changes in some tundra ecosystems appear to be responses to a warming climate. Several experimental studies have shown that tundra plants and ecosystems can respond strongly to environmental change, including warming; however, most studies were limited to a single location and were of short duration and based on a variety of experimental designs. In addition, comparisons among studies are difficult because a variety of techniques have been used to achieve experimental warming and different measurements have been used to assess responses. We used metaanalysis on plant community measurements from standardized warming experiments at 11 locations across the tundra biome involved in the International Tundra Experiment. The passive warming treatment increased plant-level air temperature by 1-3 degrees C, which is in the range of predicted and observed warming for tundra regions. Responses were rapid and detected in whole plant communities after only two growing seasons. Overall, warming increased height and cover of deciduous shrubs and graminoids, decreased cover of mosses and lichens, and decreased species diversity and evenness. These results predict that warming will cause a decline in biodiversity across a wide variety of tundra, at least in the short term. They also provide rigorous experimental evidence that recently observed increases in shrub cover in many tundra regions are in response to climate warming. These changes have important implications for processes and interactions within tundra ecosystems and between tundra and the atmosphere.