Plant carbon allocation drives turnover of old soil organic matter in permafrost tundra soils

Carbon cycle feedbacks from permafrost ecosystems are expected to accelerate global climate change. Shifts in vegetation productivity and composition in permafrost regions could influence soil organic carbon (SOC) turnover rates via rhizosphere (root zone) priming effects (RPEs), but these processes are not currently accounted for in model predictions. We use a radiocarbon (bomb‐¹⁴C) approach to test for RPEs in two Arctic tall shrubs, alder (Alnus viridis (Chaix) DC.) and birch (Betula glandulosa Michx.), and in ericaceous heath tundra vegetation. We compare surface CO₂ efflux rates and ¹⁴C content between intact vegetation and plots in which below‐ground allocation of recent photosynthate was prevented by trenching and removal of above‐ground biomass. We show, for the first time, that recent photosynthate drives mineralization of older (>50 years old) SOC under birch shrubs and ericaceous heath tundra. By contrast, we find no evidence of RPEs in soils under alder. This is the first direct evidence from permafrost systems that vegetation influences SOC turnover through below‐ground C allocation. The vulnerability of SOC to decomposition in permafrost systems may therefore be directly linked to vegetation change, such that expansion of birch shrubs across the Arctic could increase decomposition of older SOC. Our results suggest that carbon cycle models that do not include RPEs risk underestimating the carbon cycle feedbacks associated with changing conditions in tundra regions.     

Vegetation shift from deciduous to evergreen dwarf shrubs in response to selective herbivory offsets carbon losses: evidence from 19 years of warming and simulated herbivory in the subarctic tundra

Selective herbivory of palatable plant species provides a competitive advantage for unpalatable plant species, which often have slow growth rates and produce slowly decomposable litter. We hypothesized that through a shift in the vegetation community from palatable, deciduous dwarf shrubs to unpalatable, evergreen dwarf shrubs, selective herbivory may counteract the increased shrub abundance that is otherwise found in tundra ecosystems, in turn interacting with the responses of ecosystem carbon (C) stocks and CO₂ balance to climatic warming. We tested this hypothesis in a 19‐year field experiment with factorial treatments of warming and simulated herbivory on the dominant deciduous dwarf shrub Vaccinium myrtillus. Warming was associated with a significantly increased vegetation abundance, with the strongest effect on deciduous dwarf shrubs, resulting in greater rates of both gross ecosystem production (GEP) and ecosystem respiration (ER) as well as increased C stocks. Simulated herbivory increased the abundance of evergreen dwarf shrubs, most importantly Empetrum nigrum ssp. hermaphroditum, which led to a recent shift in the dominant vegetation from deciduous to evergreen dwarf shrubs. Simulated herbivory caused no effect on GEP and ER or the total ecosystem C stocks, indicating that the vegetation shift counteracted the herbivore‐induced C loss from the system. A larger proportion of the total ecosystem C stock was found aboveground, rather than belowground, in plots treated with simulated herbivory. We conclude that by providing a competitive advantage to unpalatable plant species with slow growth rates and long life spans, selective herbivory may promote aboveground C stocks in a warming tundra ecosystem and, through this mechanism, counteract C losses that result from plant biomass consumption.     

Anthropogenic nitrogen deposition enhances carbon sequestration in boreal soils

It is proposed that carbon (C) sequestration in response to reactive nitrogen (N_r) deposition in boreal forests accounts for a large portion of the terrestrial sink for anthropogenic CO₂ emissions. While studies have helped clarify the magnitude by which N_r deposition enhances C sequestration by forest vegetation, there remains a paucity of long‐term experimental studies evaluating how soil C pools respond. We conducted a long‐term experiment, maintained since 1996, consisting of three N addition levels (0, 12.5, and 50 kg N ha^?1 yr^?1) in the boreal zone of northern Sweden to understand how atmospheric N_r deposition affects soil C accumulation, soil microbial communities, and soil respiration. We hypothesized that soil C sequestration will increase, and soil microbial biomass and soil respiration will decrease, with disproportionately large changes expected compared to low levels of N addition. Our data showed that the low N addition treatment caused a non‐significant increase in the organic horizon C pool of ~15% and a significant increase of ~30% in response to the high N treatment relative to the control. The relationship between C sequestration and N addition in the organic horizon was linear, with a slope of 10 kg C kg^?1 N. We also found a concomitant decrease in total microbial and fungal biomasses and a ~11% reduction in soil respiration in response to the high N treatment. Our data complement previous data from the same study system describing aboveground C sequestration, indicating a total ecosystem sequestration rate of 26 kg C kg^?1 N. These estimates are far lower than suggested by some previous modeling studies, and thus will help improve and validate current modeling efforts aimed at separating the effect of multiple global change factors on the C balance of the boreal region.     

Soil carbon and belowground carbon balance of a short‐rotation coppice: assessments from three different approaches

Uncertainty in soil carbon (C) fluxes across different land‐use transitions is an issue that needs to be addressed for the further deployment of perennial bioenergy crops. A large‐scale short‐rotation coppice (SRC) site with poplar (Populus) and willow (Salix) was established to examine the land‐use transitions of arable and pasture to bioenergy. Soil C pools, output fluxes of soil CO₂, CH₄, dissolved organic carbon (DOC) and volatile organic compounds, as well as input fluxes from litter fall and from roots, were measured over a 4‐year period, along with environmental parameters. Three approaches were used to estimate changes in the soil C. The largest C pool in the soil was the soil organic carbon (SOC) pool and increased after four years of SRC from 10.9 to 13.9 kg C m^?2. The belowground woody biomass (coarse roots) represented the second largest C pool, followed by the fine roots (Fr). The annual leaf fall represented the largest C input to the soil, followed by weeds and Fr. After the first harvest, we observed a very large C input into the soil from high Fr mortality. The weed inputs decreased as trees grew older and bigger. Soil respiration averaged 568.9 g C m^?2 yr^?1. Leaching of DOC increased over the three years from 7.9 to 14.5 g C m^?2. The pool‐based approach indicated an increase of 3360 g C m^?2 in the SOC pool over the 4‐year period, which was high when compared with the ?27 g C m^?2 estimated by the flux‐based approach and the ?956 g C m^?2 of the combined eddy‐covariance + biometric approach. High uncertainties were associated to the pool‐based approach. Our results suggest using the C flux approach for the assessment of the short‐/medium‐term SOC balance at our site, while SOC pool changes can only be used for long‐term C balance assessments.     

CO2 exchange in three Canadian High Arctic ecosystems: response to long-term experimental warming

Carbon dioxide exchange, soil C and N, leaf mineral nutrition and leaf carbon isotope discrimination (LCID‐Δ) were measured in three High Arctic tundra ecosystems over 2 years under ambient and long‐term (9 years) warmed (～2°C) conditions. These ecosystems are located at Alexandra Fiord (79°N) on Ellesmere Island, Nunavut, and span a soil water gradient; dry, mesic, and wet tundra. Growing season CO₂ fluxes (i.e., net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (R_e)) were measured using an infrared gas analyzer and winter C losses were estimated by chemical absorption. All three tundra ecosystems lost CO₂ to the atmosphere during the winter, ranging from 7 to 12 g CO₂‐C m^?2 season^?1 being highest in the wet tundra. The period during the growing season when mesic tundra switch from being a CO₂ source to a CO₂ sink was increased by 2 weeks because of warming and increases in GEP. Warming during the summer stimulated dry tundra GEP more than R_e and thus, NEE was consistently greater under warmed as opposed to ambient temperatures. In mesic tundra, warming stimulated GEP with no effect on R_e increasing NEE by ～10%, especially in the first half of the summer. During the ～70 days growing season (mid‐June–mid‐August), the dry and wet tundra ecosystems were net CO₂‐C sinks (30 and 67 g C m^?2 season^?1, respectively) and the mesic ecosystem was a net C source (58 g C m^?2 season^?1) to the atmosphere under ambient temperature conditions, due in part to unusual glacier melt water flooding that occurred in the mesic tundra. Experimental warming during the growing season increased net C uptake by ～12% in dry tundra, but reduced net C uptake by ～20% in wet tundra primarily because of greater rates of R_e as opposed to lower rates of GEP. Mesic tundra responded to long‐term warming with ～30% increase in GEP with almost no change in R_e reducing this tundra type to a slight C source (17 g C m^?2 season^?1). Warming caused LCID of Dryas integrafolia plants to be higher in dry tundra and lower in Salix arctic plants in mesic and wet tundra. Our findings indicate that: (1) High Arctic ecosystems, which occur in similar mesoclimates, have different net CO₂ exchange rates with the atmosphere; (2) long‐term warming can increase the net CO₂ exchange of High Arctic tundra by stimulating GEP, but it can also reduce net CO₂ exchange in some tundra types during the summer by stimulating R_e to a greater degree than stimulating GEP; (3) after 9 years of experimental warming, increases in soil carbon and nitrogen are detectable, in part, because of increases in deciduous shrub cover, biomass, and leaf litter inputs; (4) dry tundra increases in GEP, in response to long‐term warming, is reflected in D. integrifolia LCID; and (5) the differential carbon exchange responses of dry, mesic, and wet tundra to similar warming magnitudes appear to depend, in part, on the hydrologic (soil water) conditions. Annual net ecosystem CO₂‐C exchange rates ranged from losses of 64 g C m^?2 yr^?1 to gains of 55 g C m^?2 yr^?1. These magnitudes of positive NEE are close to the estimates of NPP for these tundra types in Alexandra Fiord and in other High Arctic locations based on destructive harvests.     

Variable sensitivity of plant communities in Iceland to experimental warming

Facing an increased threat of rapid climate change in cold‐climate regions, it is important to understand the sensitivity of plant communities both in terms of degree and direction of community change. We studied responses to 3–5 years of moderate experimental warming by open‐top chambers in two widespread but contrasting tundra communities in Iceland. In a species‐poor and nutrient‐deficient moss heath, dominated by Racomitrium lanuginosum, mean daily air temperatures at surface were 1–2°C higher in the warmed plots than the controls whereas soil temperatures tended to be lower in the warmed plots throughout the season. In a species‐rich dwarf shrub heath on relatively rich soils at a cooler site, dominated by Betula nana and R. lanuginosum, temperature changes were in the same direction although more moderate. In the moss heath, there were no detectable community changes while significant changes were detected in the dwarf shrub heath: the abundance of deciduous and evergreen dwarf shrubs significantly increased (>50%), bryophytes decreased (18%) and canopy height increased (100%). Contrary to some other studies of tundra communities, we detected no changes in species richness or other diversity measures in either community and the abundance of lichens did not change. It is concluded that the sensitivity of Icelandic tundra communities to climate warming varies greatly depending on initial conditions in terms of species diversity, dominant species, soil and climatic conditions as well as land‐use history.     

Complex effects of mammalian grazing on extramatrical mycelial biomass in the Scandes forest‐tundra ecotone

Mycorrhizal associations are widespread in high‐latitude ecosystems and are potentially of great importance for global carbon dynamics. Although large herbivores play a key part in shaping subarctic plant communities, their impact on mycorrhizal dynamics is largely unknown. We measured extramatrical mycelial (EMM) biomass during one growing season in 16‐year‐old herbivore exclosures and unenclosed control plots (ambient), at three mountain birch forests and two shrub heath sites, in the Scandes forest‐tundra ecotone. We also used high‐throughput amplicon sequencing for taxonomic identification to investigate differences in fungal species composition. At the birch forest sites, EMM biomass was significantly higher in exclosures (1.36 ± 0.43 g C/m²) than in ambient conditions (0.66 ± 0.17 g C/m²) and was positively influenced by soil thawing degree‐days. At the shrub heath sites, there was no significant effect on EMM biomass (exclosures: 0.72 ± 0.09 g C/m²; ambient plots: 1.43 ± 0.94). However, EMM biomass was negatively related to Betula nana abundance, which was greater in exclosures, suggesting that grazing affected EMM biomass positively. We found no significant treatment effects on fungal diversity but the most abundant ectomycorrhizal lineage/cortinarius, showed a near‐significant positive effect of herbivore exclusion (p = .08), indicating that herbivory also affects fungal community composition. These results suggest that herbivory can influence fungal biomass in highly context‐dependent ways in subarctic ecosystems. Considering the importance of root‐associated fungi for ecosystem carbon balance, these findings could have far‐reaching implications.     

Soil carbon balance following conversion of grassland to oil palm

Oil palm (Elaeis guineensis Jacq.) crops are expanding rapidly in the tropics, with implications for the global carbon cycle. Little is currently known about soil organic carbon (SOC) dynamics following conversion to oil palm and virtually nothing for conversion of grassland. We measured changes in SOC stocks following conversion of tropical grassland to oil palm plantations in Papua New Guinea using a chronosequence of plantations planted over a 25‐year period. We further used carbon isotopes to quantify the loss of grassland‐derived and gain in oil palm‐derived SOC over this period. The grassland and oil palm soils had average SOC stocks of 10.7 and 12.0 kg m^?2, respectively, across all the study sites, to a depth of 1.5 m. In the 0–0.05 m depth interval, 0.79 kg m^?2 of SOC was gained from oil palm inputs over 25 years and approximately the same amount of the original grass‐derived SOC was lost. For the whole soil profile (0–1.5 m), 3.4 kg m^?2 of SOC was gained from oil palm inputs with no significant losses of grass‐derived SOC. The grass‐derived SOC stocks were more resistant to decrease than SOC reported in other studies. Black carbon produced in grassfires could partially but not fully account for the persistence of the original SOC stocks. Oil palm‐derived SOC accumulated more slowly where soil nitrogen contents where high. Forest soils in the same region had smaller carbon stocks than the grasslands. In the majority of cases, conversion of grassland to oil palm plantations in this region resulted in net sequestration of soil organic carbon.     

Impact of agricultural management practices on soil organic carbon: simulation of Australian wheat systems

Quantifying soil organic carbon (SOC) dynamics at a high spatial and temporal resolution in response to different agricultural management practices and environmental conditions can help identify practices that both sequester carbon in the soil and sustain agricultural productivity. Using an agricultural systems model (the Agricultural Production Systems sIMulator), we conducted a high spatial resolution and long‐term (122 years) simulation study to identify the key management practices and environmental variables influencing SOC dynamics in a continuous wheat cropping system in Australia's 96 million ha cereal‐growing regions. Agricultural practices included five nitrogen application rates (0–200 kg N ha^?1 in 50 kg N ha^?1 increments), five residue removal rates (0–100% in 25% increments), and five residue incorporation rates (0–100% in 25% increments). We found that the change in SOC during the 122‐year simulation was influenced by the management practices of residue removal (linearly negative) and fertilization (nonlinearly positive) – and the environmental variables of initial SOC content (linearly negative) and temperature (nonlinearly negative). The effects of fertilization were strongest at rates up to 50 kg N ha^?1, and the effects of temperature were strongest where mean annual temperatures exceeded 19 °C. Reducing residue removal and increasing fertilization increased SOC in most areas except Queensland where high rates of SOC decomposition caused by high temperature and soil moisture negated these benefits. Management practices were particularly effective in increasing SOC in south‐west Western Australia – an area with low initial SOC. The results can help target agricultural management practices for increasing SOC in the context of local environmental conditions, enabling farmers to contribute to climate change mitigation and sustaining agricultural production.     

Methane oxidation in contrasting soil types: responses to experimental warming with implication for landscape‐integrated CH4 budget

Arctic ecosystems are characterized by a wide range of soil moisture conditions and thermal regimes and contribute differently to the net methane (CH₄) budget. Yet, it is unclear how climate change will affect the capacity of those systems to act as a net source or sink of CH₄. Here, we present results of in situ CH₄ flux measurements made during the growing season 2014 on Disko Island (west Greenland) and quantify the contribution of contrasting soil and landscape types to the net CH₄ budget and responses to summer warming. We compared gas flux measurements from a bare soil and a dry heath, at ambient conditions and increased air temperature, using open‐top chambers (OTCs). Throughout the growing season, bare soil consumed 0.22 ± 0.03 g CH₄‐C m^?2 (8.1 ± 1.2 g CO₂‐eq m^?2) at ambient conditions, while the dry heath consumed 0.10 ± 0.02 g CH₄‐C m^?2 (3.9 ± 0.6 g CO₂‐eq m^?2). These uptake rates were subsequently scaled to the entire study area of 0.15 km², a landscape also consisting of wetlands with a seasonally integrated methane release of 0.10 ± 0.01 g CH₄‐C m^?2 (3.7 ± 1.2 g CO₂‐eq m^?2). The result was a net landscape sink of 12.71 kg CH₄‐C (0.48 tonne CO₂‐eq) during the growing season. A nonsignificant trend was noticed in seasonal CH₄ uptake rates with experimental warming, corresponding to a 2% reduction at the bare soil, and 33% increase at the dry heath. This was due to the indirect effect of OTCs on soil moisture, which exerted the main control on CH₄ fluxes. Overall, the net landscape sink of CH₄ tended to increase by 20% with OTCs. Bare and dry tundra ecosystems should be considered in the net CH₄ budget of the Arctic due to their potential role in counterbalancing CH₄ emissions from wetlands – not the least when taking the future climatic scenarios of the Arctic into account.     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

Ecosystem nitrogen fixation throughout the snow‐free period in subarctic tundra: effects of willow and birch litter addition and warming

Nitrogen (N) fixation in moss‐associated cyanobacteria is one of the main sources of available N for N‐limited ecosystems such as subarctic tundra. Yet, N₂ fixation in mosses is strongly influenced by soil moisture and temperature. Thus, temporal scaling up of low‐frequency in situ measurements to several weeks, months or even the entire growing season without taking into account changes in abiotic conditions cannot capture the variation in moss‐associated N₂ fixation. We therefore aimed to estimate moss‐associated N₂ fixation throughout the snow‐free period in subarctic tundra in field experiments simulating climate change: willow (Salix myrsinifolia) and birch (Betula pubescens spp. tortuosa) litter addition, and warming. To achieve this, we established relationships between measured in situ N₂ fixation rates and soil moisture and soil temperature and used high‐resolution measurements of soil moisture and soil temperature (hourly from May to October) to model N₂ fixation. The modelled N₂ fixation rates were highest in the warmed (2.8 ± 0.3 kg N ha^?1) and birch litter addition plots (2.8 ± 0.2 kg N ha^?1), and lowest in the plots receiving willow litter (1.6 ± 0.2 kg N ha^?1). The control plots had intermediate rates (2.2 ± 0.2 kg N ha^?1). Further, N₂ fixation was highest during the summer in the warmed plots, but was lowest in the litter addition plots during the same period. The temperature and moisture dependence of N₂ fixation was different between the climate change treatments, indicating a shift in the N₂ fixer community. Our findings, using a combined empirical and modelling approach, suggest that a longer snow‐free period and increased temperatures in a future climate will likely lead to higher N₂ fixation rates in mosses. Yet, the consequences of increased litter fall on moss‐associated N₂ fixation due to shrub expansion in the Arctic will depend on the shrub species’ litter traits.     

Traceable components of terrestrial carbon storage capacity in biogeochemical models

Biogeochemical models have been developed to account for more and more processes, making their complex structures difficult to be understood and evaluated. Here, we introduce a framework to decompose a complex land model into traceable components based on mutually independent properties of modeled biogeochemical processes. The framework traces modeled ecosystem carbon storage capacity (X_ss) to (i) a product of net primary productivity (NPP) and ecosystem residence time (τ_E). The latter τ_E can be further traced to (ii) baseline carbon residence times (τ′_E), which are usually preset in a model according to vegetation characteristics and soil types, (iii) environmental scalars (ξ), including temperature and water scalars, and (iv) environmental forcings. We applied the framework to the Australian Community Atmosphere Biosphere Land Exchange (CABLE) model to help understand differences in modeled carbon processes among biomes and as influenced by nitrogen processes. With the climate forcings of 1990, modeled evergreen broadleaf forest had the highest NPP among the nine biomes and moderate residence times, leading to a relatively high carbon storage capacity (31.5 kg cm^?2). Deciduous needle leaf forest had the longest residence time (163.3 years) and low NPP, leading to moderate carbon storage (18.3 kg cm^?2). The longest τ_E in deciduous needle leaf forest was ascribed to its longest τ′_E (43.6 years) and small ξ (0.14 on litter/soil carbon decay rates). Incorporation of nitrogen processes into the CABLE model decreased X_ss in all biomes via reduced NPP (e.g., ?12.1% in shrub land) or decreased τ_E or both. The decreases in τ_E resulted from nitrogen‐induced changes in τ′_E (e.g., ?26.7% in C₃ grassland) through carbon allocation among plant pools and transfers from plant to litter and soil pools. Our framework can be used to facilitate data model comparisons and model intercomparisons via tracking a few traceable components for all terrestrial carbon cycle models. Nevertheless, more research is needed to develop tools to decompose NPP and transient dynamics of the modeled carbon cycle into traceable components for structural analysis of land models.     

Long‐term deepened snow promotes tundra evergreen shrub growth and summertime ecosystem net CO2 gain but reduces soil carbon and nutrient pools

Arctic climate warming will be primarily during winter, resulting in increased snowfall in many regions. Previous tundra research on the impacts of deepened snow has generally been of short duration. Here, we report relatively long‐term (7–9 years) effects of experimentally deepened snow on plant community structure, net ecosystem CO₂ exchange (NEE), and soil biogeochemistry in Canadian Low Arctic mesic shrub tundra. The snowfence treatment enhanced snow depth from 0.3 to ~1 m, increasing winter soil temperatures by ~3°C, but with no effect on summer soil temperature, moisture, or thaw depth. Nevertheless, shoot biomass of the evergreen shrub Rhododendron subarcticum was near‐doubled by the snowfences, leading to a 52% increase in aboveground vascular plant biomass. Additionally, summertime NEE rates, measured in collars containing similar plant biomass across treatments, were consistently reduced ~30% in the snowfenced plots due to decreased ecosystem respiration rather than increased gross photosynthesis. Phosphate in the organic soil layer (0–10 cm depth) and nitrate in the mineral soil layer (15–25 cm depth) were substantially reduced within the snowfences (47–70 and 43%–73% reductions, respectively, across sampling times). Finally, the snowfences tended (p = .08) to reduce mineral soil layer C% by 40%, but with considerable within‐ and among plot variation due to cryoturbation across the landscape. These results indicate that enhanced snow accumulation is likely to further increase dominance of R. subarcticum in its favored locations, and reduce summertime respiration and soil biogeochemical pools. Since evergreens are relatively slow growing and of low stature, their increased dominance may constrain vegetation‐related feedbacks to climate change. We found no evidence that deepened snow promoted deciduous shrub growth in mesic tundra, and conclude that the relatively strong R. subarcticum response to snow accumulation may explain the extensive spatial variability in observed circumpolar patterns of evergreen and deciduous shrub growth over the past 30 years.     

Modelling carbon balances of coastal arctic tundra under changing climate

Rising air temperatures are believed to be hastening heterotrophic respiration (R_h) in arctic tundra ecosystems, which could lead to substantial losses of soil carbon (C). In order to improve confidence in predicting the likelihood of such loss, the comprehensive ecosystem model ecosys was first tested with carbon dioxide (CO₂) fluxes measured over a tundra soil in a growth chamber under various temperatures and soil‐water contents (θ). The model was then tested with CO₂ and energy fluxes measured over a coastal arctic tundra near Barrow, Alaska, under a range of weather conditions during 1998–1999. A rise in growth chamber temperature from 7 to 15 °C caused large, but commensurate, rises in respiration and CO₂ fixation, and so no significant effect on net CO₂ exchange was modelled or measured. An increase in growth chamber θ from field capacity to saturation caused substantial reductions in respiration but not in CO₂ fixation, and so an increase in net CO₂ exchange was modelled and measured. Long daylengths over the coastal tundra at Barrow caused an almost continuous C sink to be modelled and measured during most of July (2–4 g C m^?2 d^?1), but shortening daylengths and declining air temperatures caused a C source to be modelled and measured by early September (～1 g C m^?2 d^?1). At an annual time scale, the coastal tundra was modelled to be a small C sink (4 g C m^?2 y^?1) during 1998 when average air temperatures were 4 °C above normal, and a larger C sink (16 g C m^?2 y^?1) during 1999 when air temperatures were close to long‐term normals. During 100 years under rising atmospheric CO₂ concentration (C_a), air temperature and precipitation driven by the IS92a emissions scenario, modelled R_h rose commensurately with net primary productivity (NPP) under both current and elevated rates of atmospheric nitrogen (N) deposition, so that changes in soil C remained small. However, methane (CH₄) emissions were predicted to rise substantially in coastal tundra with IS92a‐driven climate change (from ～20 to ～40 g C m^?2 y^?1), causing a substantial increase in the emission of CO₂ equivalents. If the rate of temperature increase hypothesized in the IS92a emissions scenario had been raised by 50%, substantial losses of soil C (～1 kg C m^?2) would have been modelled after 100 years, including additional emissions of CH₄.     

Global changes in soil stocks of carbon,nitrogen, phosphorus,and sulphur as influenced by long‐term agricultural production

Quantifying changes in stocks of C, N, P, and S in agricultural soils is important not only for managing these soils sustainably as required to feed a growing human population, but for C and N, they are also important for understanding fluxes of greenhouse gases from the soil environment. In a global meta‐analysis, 102 studies were examined to investigate changes in soil stocks of organic C, total N, total P, and total S associated with long‐term land‐use changes. Conversion of native vegetation to cropping resulted in substantial losses of C (?1.6 kg m^?2, ?43%), N (?0.15 kg m^?2, ?42%), P (?0.029 kg m^?2, ?27%), and S (?0.015 kg m^?2, ?33%). The subsequent conversion of conventional cropping systems to no‐till, organic agriculture, or organic amendment systems subsequently increased stocks, but the magnitude of this increase (average of +0.47 kg m^?2 for C and +0.051 kg m^?2 for N) was small relative to the initial decrease. We also examined the conversion of native vegetation to pasture, with changes in C (?11%), N (+4.1%), and P (+25%) generally being modest relative to changes caused by conversion to cropping. The C:N ratio remained relatively constant irrespective of changes in land use, whilst in contrast, the C:S ratio decreased by 21% in soils converted to cropping – this suggesting that biochemical mineralization is of importance for S. The data presented here will assist in the assessment of different agricultural production systems on soil stocks of C, N, P, and S – this information assisting not only in quantifying the effects of existing agricultural production on these stocks, but also allowing for informed decision‐making regarding the potential effects of future land‐use changes.     

Contrasting effects of low and high nitrogen additions on soil CO2 flux components and ectomycorrhizal fungal sporocarp production in a boreal forest

Nitrogen (N) added through atmospheric deposition or as fertilizer to boreal and temperate forests reduces both soil decomposer activity (heterotrophic respiration) and the activity of roots and mycorrhizal fungi (autotrophic respiration). However, these negative effects have been found in studies that applied relatively high levels of N, whereas the responses to ambient atmospheric N deposition rates are still not clear. Here, we compared an unfertilized control boreal forest with a fertilized forest (100 kg N ha^?1 yr^?1) and a forest subject to N‐deposition rates comparable to those in Central Europe (20 kg N ha^?1 yr^?1) to investigate the effects of N addition rate on different components of forest floor respiration and the production of ectomycorrhizal fungal sporocarps. Soil collars were used to partition heterotrophic (R_h) and autotrophic (R_a) respiration, which was further separated into respiration by tree roots (R_tr) and mycorrhizal hyphae (R_m). Total forest floor respiration was twice as high in the low N plot compared to the control, whereas there were no differences between the control and high N plot. There were no differences in R_h respiration among plots. The enhanced forest floor respiration in the low N plot was, therefore, the result of increased R_a respiration, with an increase in R_tr respiration, and a doubling of R_m respiration. The latter was corroborated by a slightly greater ectomycorrhizal (EM) fungal sporocarp production in the low N plot as compared to the control plot. In contrast, EM fungal sporocarp production was nearly eliminated, and R_m respiration severely reduced, in the high N plot, which resulted in significantly lower R_a respiration. We thus found a nonlinear response of the R_a components to N addition rate, which calls for further studies of the quantitative relations among N addition rate, plant photosynthesis and carbon allocation, and the function of EM fungi.     

Site-dependent N uptake from N-form mixtures by arctic plants,soil microbes and ectomycorrhizal fungi

Soil microbes constitute an important control on nitrogen (N) turnover and retention in arctic ecosystems where N availability is the main constraint on primary production. Ectomycorrhizal (ECM) symbioses may facilitate plant competition for the specific N pools available in various arctic ecosystems. We report here our study on the N uptake patterns of coexisting plants and microbes at two tundra sites with contrasting dominance of the circumpolar ECM shrub Betula nana. We added equimolar mixtures of glycine-N, NH₄⁺–N and NO₃⁻–N, with one N form labelled with ¹⁵N at a time, and in the case of glycine, also labelled with ¹³C, either directly to the soil or to ECM fungal ingrowth bags. After 2 days, the vegetation contained 5.6, 7.7 and 9.1% (heath tundra) and 7.1, 14.3 and 12.5% (shrub tundra) of the glycine-, NH₄⁺- and NO₃⁻–¹⁵N, respectively, recovered in the plant–soil system, and the major part of ¹⁵N in the soil was immobilized by microbes (chloroform fumigation-extraction). In the subsequent 24 days, microbial N turnover transferred about half of the immobilized ¹⁵N to the non-extractable soil organic N pool, demonstrating that soil microbes played a major role in N turnover and retention in both tundra types. The ECM mycelial communities at the two tundras differed in N-form preferences, with a higher contribution of glycine to total N uptake at the heath tundra; however, the ECM mycelial communities at both sites strongly discriminated against NO₃⁻. Betula nana did not directly reflect ECM mycelial N uptake, and we conclude that N uptake by ECM plants is modulated by the N uptake patterns of both fungal and plant components of the symbiosis and by competitive interactions in the soil. Our field study furthermore showed that intact free amino acids are potentially important N sources for arctic ECM fungi and plants as well as for soil microorganisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Declining trend of carbon in Finnish cropland soils in 1974–2009

Soil organic matter not only affects soil properties and productivity but also has an essential role in global carbon (C) cycle. We studied changes in the topsoil C content of Finnish croplands using a dataset produced in nationwide soil monitoring. The monitoring network consisting of fields on both mineral and organic soils was established in 1974 and resampled in 1987, 1998, and 2009. Over the monitoring period from 1974 to 2009, cultivated soils showed a continuous decline in C concentration (g kg^?1). In organic soils, C concentration decreased at a mean rate of 0.2–0.3% yr^?1 relative to the existing C concentration. In mineral soils, the relative decrease was 0.4% yr^?1 corresponding to a C stock (kg m^?2) loss of 220 kg ha^?1 yr^?1. The change in management practices in last decades toward increasing cultivation of annual crops has contributed to soil C losses noted in this study. The results, however, suggest that the C losses result partly from other processes affecting cultivated soils such as climatic change or the continuing long‐term effect of forest clearance. We estimated that Finnish cropland soils store 161 Tg carbon nationwide in the topmost 15 cm of which 117 Tg is in mineral soils. C losses from mineral soils can therefore total up to 0.5 Tg yearly.     

Long‐term increase in snow depth leads to compositional changes in arctic ectomycorrhizal fungal communities

Many arctic ecological processes are regulated by soil temperature that is tightly interconnected with snow cover distribution and persistence. Recently, various climate‐induced changes have been observed in arctic tundra ecosystems, e.g. shrub expansion, resulting in reduction in albedo and greater C fixation in aboveground vegetation as well as increased rates of soil C mobilization by microbes. Importantly, the net effects of these shifts are unknown, in part because our understanding of belowground processes is limited. Here, we focus on the effects of increased snow depth, and as a consequence, increased winter soil temperature on ectomycorrhizal (ECM) fungal communities in dry and moist tundra. We analyzed deep DNA sequence data from soil samples taken at a long‐term snow fence experiment in Northern Alaska. Our results indicate that, in contrast with previously observed responses of plants to increased snow depth at the same experimental site, the ECM fungal community of the dry tundra was more affected by deeper snow than the moist tundra community. In the dry tundra, both community richness and composition were significantly altered while in the moist tundra, only community composition changed significantly while richness did not. We observed a decrease in richness of Tomentella, Inocybe and other taxa adapted to scavenge the soil for labile N forms. On the other hand, richness of Cortinarius, and species with the ability to scavenge the soil for recalcitrant N forms, did not change. We further link ECM fungal traits with C soil pools. If future warmer atmospheric conditions lead to greater winter snow fall, changes in the ECM fungal community will likely influence C emissions and C fixation through altering N plant availability, fungal biomass and soil‐plant C‐N dynamics, ultimately determining important future interactions between the tundra biosphere and atmosphere.