Rivers transport large amounts of allochthonous organic matter (OM) to the ocean every year, but there are still fundamental gaps in how allochthonous OM is processed in the marine environment. Here, we estimated the relative contribution of allochthonous OM (allochthony) to the biomass of benthic and pelagic consumers in a shallow coastal ecosystem in the northern Baltic Sea. We used deuterium as a tracer of allochthony and assessed both temporal variation (monthly from May to August) and spatial variation (within and outside river plume). We found variability in allochthony in space and time and across species, with overall higher values for zoobenthos (26.2 +/- 20.9%) than for zooplankton (0.8 +/- 0.3%). Zooplankton allochthony was highest in May and very low during the other months, likely as a result of high inputs of allochthonous OM during the spring flood that fueled the pelagic food chain for a short period. In contrast, zoobenthos allochthony was only lower in June and remained high during the other months. Allochthony of zoobenthos was generally higher close to the river mouth than outside of the river plume, whereas it did not vary spatially for zooplankton. Last, zoobenthos allochthony was higher in deeper than in shallower areas, indicating that allochthonous OM might be more important when autochthonous resources are limited. Our results suggest that climate change predictions of increasing inputs of allochthonous OM to coastal ecosystems may affect basal energy sources supporting coastal food webs.

Deposition of feces is a key mechanism by which herbivores influence soil nutrient cycling and plant production, but the knowledge about its importance for plant production and community structure is still rudimental since experimental evidence is scarce. We thus performed a 7-year long reindeer feces manipulation experiment in two tundra vegetation types with contrasting nutrient availability and analyzed effects on plant community composition and soil nutrient availability. Despite feces being fairly nutrient poor, feces manipulation had strong effect on both the nutrient-poor heath and the nutrient-rich meadow. The strongest effect was detected when feces were added at high density, with a substantial increase in total vascular plant productivity and graminoids in the two communities. Doubling natural deposition of reindeer feces enhanced primary production and the growth of deciduous shrubs in the heath. By contrast, removal of feces decreased only the production of graminoids and deciduous shrubs in the heath. Although the response to feces addition was faster in the nutrient-rich meadow, after 7 years it was more pronounced in the nutrient-poor heath. The effect of feces manipulation on soil nutrient availability was low and temporarily variable. Our study provides experimental evidence for a central role of herbivore feces in regulating primary production when herbivores are abundant enough. Deposition of feces alone does, however, not cause dramatic vegetation shifts; to drive unproductive heath to a productive grass dominated state, herbivore trampling, and grazing are probably also needed.

Low nitrogen (N) availability in the Arctic and Subarctic constrains plant productivity, resulting in low litter inputs to soil. Increased N availability and litter inputs as a result of climate change, therefore, have the potential to impact the functioning of these ecosystems. We examined plant and microbial responses to chronic inorganic N (5 g m−2 year−1) and/or litter (90 g m−2 year−1), supplied during three growing seasons. We also compared the response to more extreme additions, where the total cumulative additions of N (that is, 15 g m−2) and litter (that is, 270 g m−2) were concentrated into a single growth season. Plant productivity was stimulated by N additions and was higher in the extreme addition plots than those with chronic annual additions. Microbial community structure also differed between the chronic and extreme plots, and there was a significant relationship between plant and microbial community structures. Despite differences in microbial structure, the field treatments had no effect on microbial growth or soil C mineralization. However, gross N mineralization was higher in the N addition plots. This led to a lower ratio of soil C mineralization to gross N mineralization, indicating microbial targeting of N-rich organic matter (“microbial N-mining”), likely driven by the increased belowground C-inputs due to stimulated plant productivity. Surprisingly, aboveground litter addition also decreased ratio of soil C mineralization to gross N mineralization. Together, these results suggest that elevated N availability will induce strong responses in tundra ecosystems by promoting plant productivity, driving changes in above- and belowground community structures, and accelerating gross N mineralization. In contrast, increased litter inputs will have subtle effects, primarily altering the ratio between C and N derived from soil organic matter.

We compared terrestrial net primary production (NPP) and terrestrial export of dissolved organic carbon (DOC) with lake water heterotrophic bacterial activity in 12 headwater lake catchments along an altitude gradient in subarctic Sweden. Modelled NPP declined strongly with altitude and annual air temperature decreases along the altitude gradient (6°C between the warmest and the coldest catchment). Estimated terrestrial DOC export to the lakes was closely correlated to NPP. Heterotrophic bacterial production (BP) and respiration (BR) were mainly based on terrestrial organic carbon and strongly correlated with the terrestrial DOC export. Excess respiration over PP of the pelagic system was similar to net emission of CO2 in the lakes. BR and CO2 emission made up considerably higher shares of the terrestrial DOC input in warm lakes than in cold lakes, implying that respiration and the degree of net heterotrophy in the lakes were dependant not only on terrestrial export of DOC, but also on characteristics in the lakes which changed along the gradient and affected the bacterial metabolization of allochthonous DOC. The study showed close links between terrestrial primary production, terrestrial DOC export and bacterial activity in lakes and how these relationships were dependant on air temperature. Increases in air temperature in high latitude unproductive systems might have considerable consequences for lake water productivity and release of CO2 to the atmosphere, which are ultimately determined by terrestrial primary production.

Partial pressure (pCO(2)) and flux to the atmosphere of carbon dioxide (CO2) were studied in northern alpine and forest lakes along a gradient of dissolved organic carbon (DOC) content (0.4-9.9 mg L-1). Sixteen lakes were each sampled three times over the course of the ice-free season, and an additional 35 lakes were sampled once at midsummer. pCO(2) data were acquired in the field by a headspace equilibration technique. Most lakes were supersaturated with CO2 along the entire DOC gradient, with relatively small seasonal differences. pCO(2) was positively correlated to DOC content, reflecting a close dependence between allochthonous DOC in-put and heterotrophic respiration in the lakes. Fluxes of CO2 to the atmosphere were estimated from the pCO(2) measurements. Benthic respiration was indicated to be important for CO2 emission in lakes with high DOC concentrations. In lakes with low DOC concentrations, pelagic mineralization alone was sufficient to account for a large part of the estimated fluxes.

Tundra vegetation is responding rapidly to on-going climate warming. The changes in plant abundance and chemistry might have cascading effects on tundra food webs, but an integrated understanding of how the responses vary between habitats and across environmental gradients is lacking. We assessed responses in plant abundance and plant chemistry to warmer climate, both at species and community levels, in two different habitats. We used a long-term and multisite warming (OTC) experiment in the Scandinavian forest–tundra ecotone to investigate (i) changes in plant community composition and (ii) responses in foliar nitrogen, phosphorus, and carbon-based secondary compound concentrations in two dominant evergreen dwarf-shrubs (Empetrum hermaphroditum and Vaccinium vitis-idaea) and two deciduous shrubs (Vaccinium myrtillus and Betula nana). We found that initial plant community composition, and the functional traits of these plants, will determine the responsiveness of the community composition, and thus community traits, to experimental warming. Although changes in plant chemistry within species were minor, alterations in plant community composition drive changes in community-level nutrient concentrations. In view of projected climate change, our results suggest that plant abundance will increase in the future, but nutrient concentrations in the tundra field layer vegetation will decrease. These effects are large enough to have knock-on consequences for major ecosystem processes like herbivory and nutrient cycling. The reduced food quality could lead to weaker trophic cascades and weaker top down control of plant community biomass and composition in the future. However, the opposite effects in forest indicate that these changes might be obscured by advancing treeline forests.

Permafrost thaw releases nutrients and metals from previously frozen soils and these nutrients may affect important biogeochemical processes including methane (CH4) production and oxidation. Here we assessed how concentrations of nutrients, solutes, and metals varied across four plant communities undergoing permafrost thaw and if these geochemical characteristics affected rates of CH4 production and oxidation. We tested nutrient limitation in CH4 production and oxidation by experimentally adding nitrogen (N), phosphorus (P) and a permafrost leachate to peat across these four plant communities. The upper 20 cm of permafrost contained 715 ± 298 mg m−2 of extractable inorganic N and 20 ± 6 mg m−2 of resin-extractable phosphorus (Presin), for a N:P ratio of 36:1. These low amounts of Presin coincide with high acid-digestible aluminum (Al), iron (Fe), and P concentrations in the permafrost soil and suggest that P may accumulate via sorption and constrain easily available forms of P for plants and microbes. Permafrost leachate additions decreased potential CH4 production rates up to 80% and decreased CH4 oxidation rates by 66%, likely due to inhibitory effects of N in the permafrost. In contrast, organic and inorganic P additions increased CH4 oxidation rates up to 36% in the tall graminoid fen, a community where phosphate availability was low and CH4 production was high. Our results suggest that (1) inorganic N is available immediately from permafrost thaw, while (2) P availability is controlled by sorption properties, and (3) plant community, nutrient stoichiometry, and metal availability modulate how permafrost thaw affects CH4 production and oxidation.

Nutrient availability limits productivity of arctic ecosystems, and this constraint means that the amount of nitrogen (N) in plant canopies is an exceptionally strong predictor of vegetation productivity. However, climate change is predicted to increase nutrient availability leading to increases in carbon sequestration and shifts in community structure to more productive species. Despite tight coupling of productivity with canopy nutrients at the vegetation scale, it remains unknown how species/shoot level foliar nutrients couple to growth, or how climate change may influence foliar nutrients–productivity relationships to drive changes in ecosystem carbon gain and community structure. We investigated the influence of climate change on arctic plant growth relationships to shoot level foliar N and phosphorus (P) in three dominant subarctic dwarf shrubs using an 18-year warming and nutrient addition experiment. We found a tight coupling between total leaf N and P per shoot, leaf area and shoot extension. Furthermore, a steeper shoot length-leaf N relationship in deciduous species (Vaccinium myrtillus and Vaccinium uliginosum) under warming manipulations suggests a greater capacity for nitrogen to stimulate growth under warmer conditions in these species. This mechanism may help drive the considerable increases in deciduous shrub cover observed already in some arctic regions. Overall, our work provides the first evidence at the shoot level of tight coupling between foliar N and P, leaf area and growth i.e. consistent across species, and provides mechanistic insight into how interspecific differences in alleviation of nutrient limitation will alter community structure and primary productivity in a warmer Arctic.

Nitrogen (N) is a critical resource for plant growth in tundra ecosystems, and species differences in the timing of N uptake may be an important feature regulating community composition and ecosystem productivity. We added ¹⁵N-labelled glycine to a subarctic heath tundra dominated by dwarf shrubs, mosses and graminoids in fall, and investigated its partitioning among ecosystem components at several time points (October, November, April, May, June) through to the following spring/early summer. Soil microbes had acquired 65 ± 7% of the ¹⁵N tracer by October, but this pool decreased through winter to 37 ± 7% by April indicating significant microbial N turnover prior to spring thaw. Only the evergreen dwarf shrubs showed active ¹⁵N acquisition before early May indicating that they had the highest potential of all functional groups for acquiring nutrients that became available in early spring. The faster-growing deciduous shrubs did not resume ¹⁵N acquisition until after early May indicating that they relied more on nitrogen made available later during the spring/early summer. The graminoids and mosses had no significant increases in ¹⁵N tracer recovery or tissue ¹⁵N tracer concentrations after the first harvest in October. However, the graminoids had the highest root ¹⁵N tracer concentrations of all functional groups in October indicating that they primarily relied on N made available during summer and fall. Our results suggest a temporal differentiation among plant functional groups in the post-winter resumption of N uptake with evergreen dwarf shrubs having the highest potential for early N uptake, followed by deciduous dwarf shrubs and graminoids.

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.

Soils in northern latitudes store more than twice the amount of carbon (C) currently in the atmosphere and are warming faster than the rest of the globe. Warming has been linked to an expansion of woody vegetation across tundra, raising questions about how these two phenomena interact to modulate C stocks and turnover. We investigated how long-term warming and litter addition have modified microbial processes, soil characteristics, and C and nitrogen (N) stocks. We hypothesized that warming and litter would interact to amplify soil C losses and would be accompanied by increases in microbial activity. Using soil samples from a 16-year warming and litter addition field manipulation, we measured soil C and N stocks, heterotrophic respiration, extracellular enzyme activity, and microbial stoichiometry. We found that warming decreased C and N stocks across the entire soil profile. Depth-specific analyses illustrated that these changes are driven by increasing microbial activity at 5–10 and 10–15 cm depth, and trends toward higher dissolved organic C and N at 5–10 cm depth. This emphasizes the potential for increased leaching losses with warming and additional litter. While litter addition did not change overall C and N stocks, it appears to modify the ecosystem by adding nutrients and C to the soil. Collectively, these findings highlight the vulnerability of northern soils to continued warming with respect to nutrient and C turnover and provide insights into the mechanistic responses of tundra soil to prolonged global change.

Nitrogen (N) fixation is the main source of ‘new’ N for N-limited ecosystems like subarctic and arctic tundra. This crucial ecosystem function is performed by a wide range of N2 fixer (diazotroph) associations that could differ fundamentally in their timing and amount of N release to the soil. To assess the importance of different associative N2 fixers for ecosystem N cycling, we tracked 15N-N2 into four N2-fixer associations (with a legume, lichen, free-living, moss) and into soil, microbial biomass and non-diazotroph-associated plants 3 days and 5 weeks after in situ labelling. In addition, we tracked 13C from 13CO2 labelling to assess if N and C fixation are linked. Three days after labelling, half of the fixed 15N was recovered in the legume soils, indicating a fast release of fixed N2. Within 5 weeks, the free-living N2 fixers released two-thirds of the fixed 15N into the soil, whereas the lichen and moss retained the fixed 15N. Carbon and N2 fixation were linked in the lichen shortly after labelling, in free-living N2 fixers 5 weeks after labelling, and in the moss at both sampling times. The four investigated N2-fixer associations released fixed N2 at different rates into the soil, and non-diazotroph-associated plants have no access to ‘new’ N within several weeks after N2 fixation. Although legumes and free-living N2 fixers are immediate sources of ‘new’ N for N-limited tundra ecosystems, lichens and especially mosses, do not contribute to increase the N pool via N2 fixation in the short term.

Tree mortality from insect infestations can significantly reduce carbon storage in forest soils. In subarctic birch forests (Betula pubescens), ecosystem C cycling is largely affected by recurrent outbreaks of defoliating geometrid moths (Epirrita autumnata, Operophtera brumata). Here, we show that soil C stocks in birch forests across Fennoscandia did not change up to 8 years after moth outbreaks. We found that a decrease in woody fine roots was accompanied by a lower soil CO2 efflux rate and a higher soil N availability following moth outbreaks. We suggest that a high N availability and less ectomycorrhiza likely contributed to lowered heterotrophic respiration and soil enzymatic activity. Based on proxies for decomposition (heterotrophic respiration, phenol oxidase potential activity), we conclude that a decrease in decomposition is a prime cause why soil C stocks of mountain birch forest ecosystems have not changed after moth outbreaks. Compared to disturbed temperate and boreal forests, a CO2-related positive feedback of forest disturbance on climate change might therefore be smaller in subarctic regions.

For the first time in an arctic long-term warming and fertilization experiment, the short-term (days) and longer-term (month and year) nitrogen (N) uptake and allocation in plants, microbes, and soil pools were studied, with ¹⁵N-labeling of an organic nitrogen form, glycine. The long-term warming and fertilization had no marked effect on soil inorganic N content, but both dissolved organic N (DON) and plant biomass did increase after fertilization. Soil microbes initially immobilized most of the added ¹⁵N, but in the following months, they lost two-thirds, while label concentration in plants increased. After a year, however, the ¹⁵N recovered in microbes was still 10-fold higher than that in the plant biomass, showing the high importance of soil microbes in nutrient retention in arctic ecosystems, irrespective of the impact of long-term warming or fertilization. The effects of the treatments on the uptake of label by deciduous shrubs and evergreens paralleled that of their N pool sizes, suggesting that their N uptake potential was unaffected by long-term warming and fertilizer addition. Mosses and herbs had high uptake potential but in fertilized plots they took up less ¹⁵N, that is, they were N saturated. The fraction of ¹⁵N in microbes tended to decrease after fertilization, but this was an effect of higher N pool dilution after 1 month and a year, and not due to lower initial uptake. Although the concentration of soil inorganic N did not change after fertilization, both increased DON and the results of the 15N label addition showed that the N availability in the ecosystem had increased. By contrast, warming had little effect on soil N pools and microbial ¹⁵N uptake, and, hence, had no detectable effects on ¹⁵N accumulation.

The consequences of warming-induced ‘shrubification’ on Arctic soil carbon storage are receiving increased attention, as the majority of ecosystem carbon in these systems is stored in soils. Soil carbon cycles in these ecosystems are usually tightly coupled with nitrogen availability. Soil microbial responses to ‘shrubification’ may depend on the traits of the shrub species that increase in response to warming. Increase in deciduous shrubs such as Betula nana likely promotes a loss of soil carbon, whereas the opposite may be true if evergreen shrubs such as Empetrum hermaphroditum increase. We analyzed soil organic matter stocks and 13C NMR fractions, microbial CO2 respiration, biomass, extracellular enzyme activities (EEAs), and their association with shrub density in northern Sweden after 20 years of experimental warming using open top chambers (OTCs). Our study sites were located in a tundra heath that stores high soil carbon quantities and where the OTCs had increased deciduous shrubs, and in a mountain birch forest that stores lower soil carbon quantities and where the OTCs had increased evergreen shrubs. We predicted that organic matter stocks should be lower and respiration and EEAs higher inside the OTCs than untreated plots in the tundra, whereas no effect should be detected in the forest. Soil organic matter stocks and 13C NMR fractions remained unaffected at both sites. When expressed as per gram microbial biomass, respiration and EEAs for carbohydrate and chitin degradation were higher inside the OTCs, and contrasting our prediction, this effect was stronger in the forest. Unexpectedly, the OTCs also led to a substantially lower microbial biomass carbon and nitrogen irrespective of habitat. The decline in the microbial biomass counteracted increased activities resulting in no effect of the OTCs on respiration and a lower phenol oxidase activity per gram soil. Microbial biomass nitrogen correlated negatively with evergreen shrub density at both sites, indicating that ‘shrubification’ may have intensified nutrient competition between plants and soil microorganisms. Nutrient limitation could also underlie increased respiration per gram microbial biomass through limiting C assimilation into biomass. We hypothesize that increasing nutrient immobilization into long-lived evergreen shrubs could over time induce microbial nutrient limitation that contributes to a stability of accumulated soil organic matter stocks under climate warming.

Quantifying vegetation structure and function is critical for modeling ecological processes, and an emerging challenge is to apply models at multiple spatial scales. Land surface heterogeneity is commonly characterized using rectangular pixels, whose length scale reflects that of remote sensing measurements or ecological models rather than the spatial scales at which vegetation structure and function varies. We investigated the ‘optimum’ pixel size and shape for averaging leaf area index (LAI) measurements in relatively large (85 m2 estimates on a 600 × 600-m2 grid) and small (0.04 m2 measurements on a 40 × 40-m2 grid) patches of sub-Arctic tundra near Abisko, Sweden. We define the optimum spatial averaging operator as that which preserves the information content (IC) of measured LAI, as quantified by the normalized Shannon entropy (ES,n) and Kullback–Leibler divergence (DKL), with the minimum number of pixels. Based on our criterion, networks of Voronoi polygons created from triangulated irregular networks conditioned on hydrologic and topographic indices are often superior to rectangular shapes for averaging LAI at some, frequently larger, spatial scales. In order to demonstrate the importance of information preservation when upscaling, we apply a simple, validated ecosystem carbon flux model at the landscape level before and after spatial averaging of land surface characteristics. Aggregation errors are minimal due to the approximately linear relationship between flux and LAI, but large errors of approximately 45% accrue if the normalized difference vegetation index (NDVI) is averaged without preserving IC before conversion to LAI due to the nonlinear NDVI-LAI transfer function.