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Biochar management has been proposed as a possible tool to mitigate anthropogenic CO2 emissions, and thus far its impacts in forested environments remain poorly understood. We conducted a large-scale, replicated field experiment using 0.05-ha plots in the boreal region in northern Sweden to evaluate how soil and vegetation properties and processes responded to biochar application and the disturbance associated with burying biochar in the soil. We employed a randomized block design, where biochar and soil mixing treatments were established in factorial combination (i.e., control, soil mixing only, biochar only, and biochar and soil mixing; n = 6 plots of each). After two growing seasons, we found that biochar application enhanced net soil N mineralization rates and soil [ABSTRACT FROM AUTHOR]

It is proposed that increases in anthropogenic reactive nitrogen (Nr) deposition may cause temperate and boreal forests to sequester a globally significant quantity of carbon (C); however, long-term data from boreal forests describing how C sequestration responds to realistic levels of chronic Nr deposition are scarce. Using a long-term (14-year) stand-scale (0.1 ha) N addition experiment (three levels: 0, 12.5, and 50 kg N ha−1 yr−1) in the boreal zone of northern Sweden, we evaluated how chronic N additions altered N uptake and biomass of understory communities, and whether changes in understory communities explained N uptake and C sequestration by trees. We hypothesized that understory communities (i.e. mosses and shrubs) serve as important sinks for low-level N additions, with the strength of these sinks weakening as chronic N addition rates increase, due to shifts in species composition. We further hypothesized that trees would exhibit nonlinear increases in N acquisition, and subsequent C sequestration as N addition rates increased, due to a weakening understory N sink. Our data showed that understory biomass was reduced by 50% in response to the high N addition treatment, mainly due to reduced moss biomass. A 15N labeling experiment showed that feather mosses acquired the largest fraction of applied label, with this fraction decreasing as the chronic N addition level increased. Contrary to our hypothesis, the proportion of label taken up by trees was equal (ca. 8%) across all three N addition treatments. The relationship between N addition and C sequestration in all vegetation pools combined was linear, and had a slope of 16 kg C kg−1 N. While canopy retention of Nr deposition may cause C sequestration rates to be slightly different than this estimate, our data suggest that a minor quantity of annual anthropogenic CO2 emissions are sequestered into boreal forests as a result of Nr deposition. [ABSTRACT FROM AUTHOR]

Productivity in boreal ecosystems is primarily limited by available soil nitrogen (N), and there is substantial interest in understanding whether deposition of anthropogenically derived reactive nitrogen (Nr) results in greater N availability to woody vegetation, which could result in greater carbon (C) sequestration. One factor that may limit the acquisition of Nr by woody plants is the presence of bryophytes, which are a significant C and N pool, and a location where associative cyanobacterial N-fixation occurs. Using a replicated stand-scale N-addition experiment (five levels: 0, 3, 6, 12, and 50 kgNha-1 yr-1; n56) in the boreal zone of northern Sweden, we tested the hypothesis that sequestration of Nr into bryophyte tissues, and downregulation of N-fixation would attenuate Nr inputs, and thereby limit anthropogenic Nr acquisition by woody plants. Our data showed that N-fixation per unit moss mass and per unit area sharply decreased with increasing N addition. Additionally, the tissue N concentrations of Pleuorzium schreberi increased and its biomass decreased with increasing N addition. This response to increasing N addition caused the P. schreberi N pool to be stable at all but the highest N addition rate, where it significantly decreased. The combined effects of changed N-fixation and P. schreberi biomass N accounted for 56.7% of cumulative Nr additions at the lowest Nr addition rate, but only a minor fraction for all other treatments. This 'bryophyte effect' can in part explain why soil inorganic N availability and acquisition by woody plants (indicated by their δ15N signatures) remained unchanged up to N addition rates of 12 kg ha-1 yr-1 or greater. Finally, we demonstrate that approximately 71.8% of the boreal forest experiences Nr deposition rates at or below 3 kg ha-1 yr-1, suggesting that bryophytes likely limit woody plant acquisition of ambient anthropogenic Nr inputs throughout a majority of the boreal forest. [ABSTRACT FROM AUTHOR]

It is well established that application of biochar to soils can promote soil fertility, which ultimately may enhance plant growth. While many mechanisms have been proposed to explain this, one specific mechanism, the "microbial refugia hypothesis," suggests that biochar may provide physical protection for soil microbe from soil microfauna that otherwise exert top‐down control on microbial biomass and activity. We tested the microbial refugia hypothesis by incubating two boreal soils with and without biochar derived from a wood mixture of boreal tree species (Picea abies and Pinus sylvestris), and with and without soil nematodes. We measured phospholipid fatty acids (PLFA) as a relative measure of microbial biomass, and several variables indicative of microbial activity, including extractable nutrient concentrations (NH4+, NO3−, and PO4−), heterotrophic N2‐fixation, and soil respiration. Contrary to our expectations, we found that biochar by itself did not stimulate microbial biomass or activity. Furthermore, we found that nematode addition to soil stimulated rather than depressed the biomass of several bacterial PLFA groups. Finally, interactive effects between the nematode treatment and biochar never worked in a way that supported the microbial refugia hypothesis. Our findings suggest that a typical boreal biochar applied to boreal soils may not have the same stimulatory effect on microbial biomass and activity that has been shown in some other ecosystems, and that enhanced plant growth in response to biochar addition sometimes observed in boreal environments is likely due to other mechanisms, such as direct nutrient supply from biochar or amelioration of soil pH. [ABSTRACT FROM AUTHOR]

Ecosystems in the far north, including arctic and boreal biomes, are a globally significant pool of carbon ( C). Global change is proposed to influence both C uptake and release in these ecosystems, thereby potentially affecting whether they act as C sources or sinks. Bryophytes (i.e., mosses) serve a variety of key functions in these systems, including their association with nitrogen ( N2)-fixing cyanobacteria, as thermal insulators of the soil, and producers of recalcitrant litter, which have implications for both net primary productivity ( NPP) and heterotrophic respiration. While ground-cover bryophytes typically make up a small proportion of the total biomass in northern systems, their combined physical structure and N2-fixing capabilities facilitate a disproportionally large impact on key processes that control ecosystem C and N cycles. As such, the response of bryophyte-cyanobacteria associations to global change may influence whether and how ecosystem C balances are influenced by global change. Here, we review what is known about their occurrence and N2-fixing activity, and how bryophyte systems will respond to several key global change factors. We explore the implications these responses may have in determining how global change influences C balances in high northern latitudes. [ABSTRACT FROM AUTHOR]

Biochar soil amendment may provide the forestry sector with a formidable tool to simultaneously sequester carbon (C) in the soil and aboveground by enhancing plant productivity, yet several key uncertainties remain. Crucially, empirical evidence of long‐term effects of biochar management on vegetation and on greenhouse gas emissions in forest ecosystems is scarce. Using a large field experiment in a young managed boreal forest in northern Sweden, we investigated the effects of biochar (applied either on the soil surface or mixed in the soil 8–9 years prior to this study) on supply rates of soil nutrients, on survival and growth of planted Pinus sylvestris, on community composition of the understory vegetation, and on forest floor fluxes of N2O, CH4, and CO2. We found that biochar promoted P. sylvestris survival only when biochar was applied on the soil surface. Conversely, biochar enhanced P. sylvestris growth overall, resulting in a 19% increase in C stored in biomass. Biochar also altered the composition of the understory vegetation, especially when mixed into the soil, and promoted a more resource‐conservative community (i.e., with more ericaceous shrubs and less graminoids and forbs). Meanwhile, supply rates of the main soil nutrients were largely unaffected by biochar. Finally, we found that biochar did not alter overall N2O and CO2 emissions and CH4 uptake from the forest floor. Our findings show that biochar amendment increased the net C input to the system, since, besides directly increasing soil C stocks, biochar enhanced biomass growth without increasing soil C losses. Therefore, our study suggests that biochar could potentially be used for emissions abatement in intensively managed boreal forests. [ABSTRACT FROM AUTHOR]

Nitrogen (N) deposition can enhance carbon (C) capture and storage in northern coniferous forests but it may also enhance the demand for phosphorus (P). While it is well established that long‐term N enrichment can decrease decomposition and enhance the accumulation of C in soils, it remains uncertain if a higher demand and acquisition of P influence soil C.We studied microbial phosphorus mobilization and growth within a long‐term N enrichment experiment in a Norway spruce forest, where N deposition was simulated by adding 0, 12.5 or 50 kg N ha−1 year−1 for 21 years (n = 12), by incubating microbial ingrowth cores with needles and humus with low and high P content, and with sand with and without mineral apatite P.Long‐term N enrichment had no effect on microbial P mobilization in needles and humus and did not enhance the positive effect that apatite had on fungal growth. However, it consistently strengthened the retention of C in the soil by decreasing decomposition of needle and humus, both with low and high P content, and by increasing fungal growth in sand‐filled ingrowth cores. Furthermore, we did not find any evidence that higher microbial P mobilization in response to N enrichment affected soil C storage.These results show that long‐term N enrichment in relatively young soils dominated by coniferous trees and ectomycorrhizal fungi can have relatively small impact on microbial P mobilization from organic sources and on the potential to mobilize P from minerals, and subsequently that elevated P demand due to N enrichment is unlikely to lead to a reduction in the soil C accumulation rate. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Nitrogen (N) deposition can change the carbon (C) sink of northern coniferous forests by changing the balance between net primary production and soil respiration. We used a field experiment in an N poor Pinus sylvestris forest where five levels of N (0, 3, 6, 12, and 50 kg N ha−1 yr−1, n = 6) had been added annually for 12–13 years to investigate how litter C inputs and soil respiration, divided into its autotrophic and heterotrophic sources, respond to different rates of N input, and its subsequent effect on soil C storage. The highest N addition rate (50 kg N ha−1 yr−1) stimulated soil C accumulation in the organic layer by 22.3 kg C kg−1 N added, increased litter inputs by 46%, and decreased soil respiration per mass unit of soil C by 31.2%, mainly by decreasing autotrophic respiration. Lower N addition rates (≤ 12 kg N ha−1 yr−1) had no effect on litter inputs or soil respiration. These results support previous studies reporting on increased litter inputs coupled to impeded soil C mineralization, contributing to enhancing the soil C sink when N is supplied at high rates, but add observations for lower N addition rates more realistic for N deposition. In doing so, we show that litter production in N poor northern coniferous forests can be relatively unresponsive to low N deposition levels, that stimulation of microbial activity at low N additions is unlikely to reduce the soil C sink, and that high levels of N deposition enhance the soil C sink by increasing litter inputs and decreasing soil respiration. [ABSTRACT FROM AUTHOR]

There is evidence that anthropogenic nitrogen (N) deposition enhances carbon (C) sequestration in boreal forest soils. However, it is unclear how free‐living saprotrophs (bacteria and fungi, SAP) and ectomycorrhizal (EM) fungi responses to N addition impact soil C dynamics. Our aim was to investigate how SAP and EM communities are impacted by N enrichment and to estimate whether these changes influence decay of litter and humus. We conducted a long‐term experiment in northern Sweden, maintained since 2004, consisting of ambient, low N additions (0, 3, 6, and 12 kg N ha−1 year−1) simulating current N deposition rates in the boreal region, as well as a high N addition (50 kg N ha−1 year−1). Our data showed that long‐term N enrichment impeded mass loss of litter, but not of humus, and only in response to the highest N addition treatment. Furthermore, our data showed that EM fungi reduced the mass of N and P in both substrates during the incubation period compared to when only SAP organisms were present. Low N additions had no effect on microbial community structure, while the high N addition decreased fungal and bacterial biomasses and altered EM fungi and SAP community composition. Actinomycetes were the only bacterial SAP to show increased biomass in response to the highest N addition. These results provide a mechanistic understanding of how anthropogenic N enrichment can influence soil C accumulation rates and suggest that current N deposition rates in the boreal region (≤12 kg N ha−1 year−1) are likely to have a minor impact on the soil microbial community and the decomposition of humus and litter. [ABSTRACT FROM AUTHOR]

There is evidence that anthropogenic nitrogen (N) deposition enhances carbon (C) sequestration in boreal soils. However, key underlying mechanisms explaining this increase have not been resolved. Two potentially important mechanisms are that aboveground litter production increases, or that litter quality changes in response to N enrichment. As such, our aim was to quantify whether simulated chronic N deposition caused changes in aboveground litter production or quality in a boreal forest. We conducted a long-term (17 years) stand-scale (0.1 ha) forest experiment, consisting of three N addition levels (0, 12.5, and 50 kg N ha-1 yr-1) in northern Sweden, where background N deposition rates are very low. We measured the annual quantity of litter produced for 8 different litter categories, as well as their concentrations of C, N, phosphorus (P), lignin, cellulose and hemi-cellulose. Our results indicate that mosses were the only major litter component showing significant quantitative and qualitative alterations in response to the N additions, indicative of their ability to intercept a substantial portion of the N added. These effects were, however, offset by the other litter fractions where we found no changes in the total litter fluxes, or individual chemical constituents when all litter categories were summed. This study indicates that the current annual litter fluxes cannot explain the increase in soil C that has occurred in our study system in response to simulated chronic N application. These results suggest that other mechanisms are likely to explain the increased soil C accumulation rate we have observed, such as changes in soil microbial activity, or potentially transient changes in aboveground litter inputs that were no longer present at the time of our study. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Summary 1. Despite recent interest in linkages between above- and below-ground communities and their consequences for ecosystem processes, much remains unknown about their responses to long-term ecosystem change. We synthesize multiple lines of evidence from a long-term 'natural experiment' to illustrate how ecosystem retrogression (the decline in ecosystem process rates due to long-term absence of major disturbance) drives vegetation change, and thus above-ground and below-ground carbon (C) sequestration, and communities of consumer biota. 2. Our study system involves 30 islands in Swedish boreal forest that form a 5000-year, fire-driven retrogressive chronosequence. Here, retrogression leads to lower plant productivity and slower decomposition and a community shift from plants with traits associated with resource acquisition to those linked with resource conservation. 3. We present consistent evidence that above-ground ecosystem C sequestration declines, while below-ground and total C storage increases linearly for at least 5000 years following fire absence. This increase is driven primarily by changes in vegetation characteristics, impairment of decomposer organisms and absence of humus combustion. 4. Data from contrasting trophic groups show that during retrogression, biomass or abundance of plants and decomposer biota decreases, while that of above-ground invertebrates and birds increases, due to different organisms accessing resources via distinct energy channels. Meanwhile, diversity measures of vascular plants and above-ground (but not below-ground) consumers respond positively to retrogression. 5. We show that taxonomic richness of plants and above-ground consumers are positively correlated with total ecosystem C storage, suggesting that conserving old-growth forests simultaneously maximizes biodiversity and C sequestration. However, we find little observational or experimental evidence that plant diversity is a major driver of ecosystem C storage on the islands relative to other biotic and abiotic factors. 6. Synthesis.Our study reveals that across contrasting islands differing in exposure to a key extrinsic driver (historical disturbance regime and resulting retrogression), there are coordinated responses of soil fertility, vegetation, consumer communities and ecosystem C sequestration, which all feed back to one another. It also highlights the value of well-replicated natural experiments for tackling questions about above-ground-below-ground linkages over temporal and spatial scales that are otherwise unachievable. [ABSTRACT FROM AUTHOR]