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The impacts of global environmental change on productivity in northern latitudes will be contingent on nitrogen (N) availability. In circumpolar boreal ecosystems, nonvascular plants (i.e., bryophytes) and associated N2‐fixing diazotrophs provide one of the largest known N inputs but are rarely accounted for in Earth system models. Instead, most models link N2‐fixation with the functioning of vascular plants. Neglecting nonvascular N2‐fixation may be contributing toward high uncertainty that currently hinders model predictions in northern latitudes, where nonvascular N2‐fixing plants are more common. Adequately accounting for nonvascular N2‐fixation and its drivers could subsequently improve predictions of future N availability and ultimately, productivity, in northern latitudes. Here, we review empirical evidence of boreal nonvascular N2‐fixation responses to global change factors (elevated CO2, N deposition, warming, precipitation, and shading by vascular plants), and compare empirical findings with model predictions of N2‐fixation using nine Earth system models. The majority of empirical studies found positive effects of CO2, warming, precipitation, or light on nonvascular N2‐fixation, but N deposition strongly downregulated N2‐fixation in most empirical studies. Furthermore, we found that the responses of N2‐fixation to elevated CO2 were generally consistent between models and very limited empirical data. In contrast, empirical‐model comparisons suggest that all models we assessed, and particularly those that scale N2‐fixation with net primary productivity or evapotranspiration, may be overestimating N2‐fixation under increasing N deposition. Overestimations could generate erroneous predictions of future N stocks in boreal ecosystems unless models adequately account for the drivers of nonvascular N2‐fixation. Based on our comparisons, we recommend that models explicitly treat nonvascular N2‐fixation and that field studies include more targeted measurements to improve model structures and parameterization. [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]

Trophic cascades in which predators regulate densities of organisms at lower trophic levels are important drivers of population dynamics, but effects of trophic cascades on ecosystem-level fluxes and processes, and the conditions under which top-down control is important, remain unresolved. We manipulated the structure of a food web in boreal feather mosses and found that moss-inhabiting microfauna exerted top-down control of N2-fixation by moss-associated cyanobacteria. However, the presence of higher trophic levels alleviated this top-down control, likely through feeding on bacterivorous microfauna. These effects of food-web structure on cyanobacterial N2-fixation were dependent on global change factors and strongly suppressed under N fertilisation. Our findings illustrate how food web interactions and trophic cascades can regulate N cycling in boreal ecosystems, where carbon uptake is generally strongly N-limited, and shifting trophic control of N cycling under global change is therefore likely to impact ecosystem functioning. [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]

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]