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.     

Anthropogenic nitrogen deposition in boreal forests has a minor impact on the global carbon cycle

It is proposed that increases in anthropogenic reactive nitrogen (N_r) 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 N_r 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 N_r 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 N_r deposition.     

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.     

The Sensitivity of Moss-Associated Nitrogen Fixation towards Repeated Nitrogen Input

Nitrogen (N₂) fixation is a major source of available N in ecosystems that receive low amounts of atmospheric N deposition. In boreal forest and subarctic tundra, the feather moss Hylocomium splendens is colonized by N₂ fixing cyanobacteria that could contribute fundamentally to increase the N pool in these ecosystems. However, N₂ fixation in mosses is inhibited by N input. Although this has been shown previously, the ability of N₂ fixation to grow less sensitive towards repeated, increased N inputs remains unknown. Here, we tested if N₂ fixation in H. splendens can recover from increased N input depending on the N load (0, 5, 20, 80, 320 kg N ha^-1 yr^-1) after a period of N deprivation, and if sensitivity towards increased N input can decrease after repeated N additions. Nitrogen fixation in the moss was inhibited by the highest N addition, but was promoted by adding 5 kg N ha^-1 yr^-1, and increased in all treatments during a short period of N deprivation. The sensitivity of N₂ fixation towards repeated N additions seem to decrease in the 20 and 80 kg N additions, but increased in the highest N addition (320 kg N ha^-1 yr^-1). Recovery of N in leachate samples increased with increasing N loads, suggesting low retention capabilities of mosses if N input is above 5 kg N ha^-1 yr^-1. Our results demonstrate that the sensitivity towards repeated N additions is likely to decrease if N input does not exceed a certain threshold.     

The impact of simulated chronic nitrogen deposition on the biomass and N2-fixation activity of two boreal feather moss–cyanobacteria associations

Bryophytes achieve substantial biomass and play several key functional roles in boreal forests that can influence how carbon (C) and nitrogen (N) cycling respond to atmospheric deposition of reactive nitrogen (N_r). They associate with cyanobacteria that fix atmospheric N₂, and downregulation of this process may offset anthropogenic N_r inputs to boreal systems. Bryophytes also promote soil C accumulation by thermally insulating soils, and changes in their biomass influence soil C dynamics. Using a unique large-scale (0.1 ha forested plots), long-term experiment (16 years) in northern Sweden where we simulated anthropogenic N_r deposition, we measured the biomass and N₂-fixation response of two bryophyte species, the feather mosses Hylocomium splendens and Pleurozium schreberi. Our data show that the biomass declined for both species; however, N₂-fixation rates per unit mass and per unit area declined only for H. splendens. The low and high treatments resulted in a 29% and 54% reduction in total feather moss biomass, and a 58% and 97% reduction in total N₂-fixation rate per unit area, respectively. These results help to quantify the sensitivity of feather moss biomass and N₂ fixation to chronic N_r deposition, which is relevant for modelling ecosystem C and N balances in boreal ecosystems.     

Nitrogen deposition in tropical forests from savanna and deforestation fires

We used satellite‐derived estimates of global fire emissions and a chemical transport model to estimate atmospheric nitrogen (N) fluxes from savanna and deforestation fires in tropical ecosystems. N emissions and reactive N deposition led to a net transport of N equatorward, from savannas and areas undergoing deforestation to tropical forests. Deposition of fire‐emitted N in savannas was only 26% of emissions – indicating a net export from this biome. On average, net N loss from fires (the sum of emissions and deposition) was equivalent to approximately 22% of biological N fixation (BNF) in savannas (4.0 kg N ha^?1 yr^?1) and 38% of BNF in ecosystems at the deforestation frontier (9.3 kg N ha^?1 yr^?1). Net N gains from fires occurred in interior tropical forests at a rate equivalent to 3% of their BNF (0.8 kg N ha^?1 yr^?1). This percentage was highest for African tropical forests in the Congo Basin (15%; 3.4 kg N ha^?1 yr^?1) owing to equatorward transport from frequently burning savannas north and south of the basin. These results provide evidence for cross‐biome atmospheric fluxes of N that may help to sustain productivity in some tropical forest ecosystems on millennial timescales. Anthropogenic fires associated with slash and burn agriculture and deforestation in the southern part of the Amazon Basin and across Southeast Asia have substantially increased N deposition in these regions in recent decades and may contribute to increased rates of carbon accumulation in secondary forests and other N‐limited ecosystems.     

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.     

Tree growth and soil acidification in response to 30 years of experimental nitrogen loading on boreal forest

Relations among nitrogen load, soil acidification and forest growth have been evaluated based on short‐term (<15 years) experiments, or on surveys across gradients of N deposition that may also include variations in edaphic conditions and other pollutants, which confound the interpretation of effects of N per se. We report effects on trees and soils in a uniquely long‐term (30 years) experiment with annual N loading on an un‐polluted boreal forest. Ammonium nitrate was added to replicated (N=3) 0.09 ha plots at two doses, N1 and N2, 34 and 68 kg N ha^?1 yr^?1, respectively. A third treatment, N3, 108 kg N ha^?1 yr^?1, was terminated after 20 years, allowing assessment of recovery during 10 years. Tree growth initially responded positively to all N treatments, but the longer term response was highly rate dependent with no gain in N3, a gain of 50 m³ ha^?1 stemwood in N2 and a gain of 100 m³ ha^?1 stemwood in excess of the control (N0) in N1. High N treatments caused losses of up to 70% of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺) in the mineral soil, along with decreases in pH and increases in exchangeable Al³⁺. In contrast, the organic mor‐layer (forest floor) in the N‐treated plots had similar amounts per hectare of exchangeable base cations as in the N0 treatment. Magnesium was even higher in the mor of N‐treated plots, providing evidence of up‐lift by the trees from the mineral soil. Tree growth did not correlate with the soil Ca/Al ratio (a suggested predictor of effects of soil acidity on tree growth). A boron deficiency occurred on N‐treated plots, but was corrected at an early stage. Extractable NH₄⁺ and NO₃^?were high in mor and mineral soils of on‐going N treatments, while NH₄+ was elevated in the mor only in N3 plots. Ten years after termination of N addition in the N3 treatment, the pH had increased significantly in the mineral soil; there were also tendencies of higher soil base status and concentrations of base cations in the foliage. Our data suggest the recovery of soil chemical properties, notably pH, may be quicker after removal of the N‐load than predicted. Our long‐term experiment demonstrated the fundamental importance of the rate of N application relative to the total amount of N applied, in particular with regard to tree growth and C sequestration. Hence, experiments adding high doses of N over short periods do not mimic the long‐term effects of N deposition at lower rates.     

Nitrogen fixation by white clover in pastures grazed by dairy cows: Temporal variation and effects of nitrogen fertilization

Effects of rate of nitrogen (N) fertilizer and stocking rate on production and N₂ fixation by white clover (Trifolium repens L.) grown with perennial ryegrass (Lolium perenne L.) were determined over 5 years in farmlets near Hamilton, New Zealand. Three farmlets carried 3.3 dairy cows ha^–1 and received urea at 0, 200 or 400 kg N ha^–1 yr^–1 in 8–10 split applications. A fourth farmlet received 400 kg N ha^–1 yr^–1 and had 4.4 cows ha^–1.There was large variation in annual clover production and total N₂ fixation, which in the 0 N treatment ranged from 9 to 20% clover content in pasture and from 79 to 212 kg N fixed ha^–1 yr^–1. Despite this variation, total pasture production in the 0 N treatment remained at 75–85% of that in the 400 N treatments in all years, due in part to the moderating effect of carry-over of fixed N between years.Fertilizer N application decreased the average proportion of clover N derived from N₂ fixation (P_N; estimated by ¹⁵N dilution) from 77% in the 0 N treatment to 43–48% in the 400 N treatments. The corresponding average total N₂ fixation decreased from 154 kg N ha^–1 yr^–1 to 39–53 kg N ha^–1 yr^–1. This includes N₂ fixation in clover tissue below grazing height estimated at 70% of N₂ fixation in above grazing height tissue, based on associated measurements, and confirmed by field N balance calculations. Effects of N fertilizer on clover growth and N₂ fixation were greatest in spring and summer. In autumn, the 200 N treatment grew more clover than the 0 N treatment and N₂ fixation was the same. This was attributed to more severe grazing during summer in the 0 N treatment, resulting in higher surface soil temperatures and a deleterious effect on clover stolons.In the 400 N treatments, a 33% increase in cow stocking rate tended to decrease P_N from 48 to 43% due to more N cycling in excreta, but resulted in up to 2-fold more clover dry matter and N₂ fixation because lower pasture mass reduced grass competition, particularly during spring.     

Carbon distribution of a well- and poorly-drained black spruce fire chronosequence

The objective of this study was to quantify carbon (C) distribution for boreal black spruce (Picea mariana (Mill.) BSP) stands comprising a fire chronosequence in northern Manitoba, Canada. The experimental design included seven well‐drained (dry) and seven poorly‐drained (wet) stands that burned between 1998 and 1850. Vegetation C pools (above‐ground + below‐ground) steadily increased from 1.3 to 83.3 t C ha^?1 for the dry chronosequence, and from 0.6 to 37.4 t C ha^?1 for the wet chronosequence. The detritus C pools (woody debris + forest floor) varied from 10.3 to 96.0 t C ha^?1 and from 12.6 to 77.4 t C ha^?1 for the dry and wet chronosequence, respectively. Overstorey biomass, mean annual biomass increment (MAI), woody debris mass, and litterfall were significantly greater (α = 0.05) for the dry stands than for the wet stands, but the bryophyte, understorey, and forest floor C pools were significantly less for the dry than for the wet stands. The root mass ratio decreased with stand age until 37 years after fire, was fairly constant thereafter, and was not significantly affected by soil drainage. The C pools of the overstorey and bryophyte tended to increase with stand age. Foliage biomass, litterfall, and MAI (for the dry stands) peaked at 71 years after fire and declined in the oldest stands. The results from this study illustrate that the effects of disturbance and edaphic conditions must be accounted for in boreal forest C inventories and C models. The appropriateness of using chronosequences to examine effects of wildfire on ecosystem C distribution is discussed.     

Exposure to nitrogen does not eliminate N2 fixation in the feather moss Pleurozium schreberi (Brid.) Mitt.

Background and aims

The feather moss Pleurozium schreberi (Brid.) Mitt. is colonized by cyanobacteria, which fix substantial amounts of atmospheric nitrogen (N) in pristine and N-poor ecosystems. Cyanobacterial N₂ fixation is inhibited by N deposition. However, the threshold of N input that leads to the inhibition of N₂ fixation has not been adequately investigated. Further, the ability of N₂ fixation to recover in mosses from high N deposition areas has not been studied to date.

Methods

We conducted two laboratory studies in which we (1) applied a range of concentrations of N as NH₄NO₃ to mosses from low N-deposition areas, and (2) we deprived mosses from a high N-deposition area of N to test their ability to recover N₂ fixation.

Results

Higher addition rates (up to 10 kg N ha^?1) did not systematically inhibit N₂ fixation in P. schreberi. Conversely, upon weeks of N deprivation of mosses from a high N environment, N₂ fixation rates increased.

Conclusions

The threshold of total N deposition above which N₂ fixation in P. schreberi is inhibited is likely to be > 10 kg N ha^?1. Further, cyanobacteria are able to recover from high N inputs and are able to fix atmospheric N₂ after a period of N deprivation.     

Soil nitrous oxide and methane fluxes are low from a bioenergy crop (canola) grown in a semi-arid climate

Understanding nitrous oxide (N₂O) and methane (CH₄) fluxes from agricultural soils in semi‐arid climates is necessary to fully assess greenhouse gas emissions from bioenergy cropping systems, and to improve our knowledge of global terrestrial gaseous exchange. Canola is grown globally as a feedstock for biodiesel production, however, resulting soil greenhouse gas fluxes are rarely reported for semi‐arid climates. We measured soil N₂O and CH₄ fluxes from a rain‐fed canola crop in a semi‐arid region of south‐western Australia for 1 year on a subdaily basis. The site included N fertilized (75 kg N ha^?1 yr^?1) and nonfertilized plots. Daily N₂O fluxes were low (?1.5 to 4.7 g N₂O‐N ha^?1 day^?1) and culminated in an annual loss of 128 g N₂O‐N ha^?1 (standard error, 12 g N₂O‐N ha^?1) from N fertilized soil and 80 g N₂O‐N ha^?1 (standard error, 11 g N₂O‐N ha^?1) from nonfertilized soil. Daily CH₄ fluxes were also low (?10.3 to 11.9 g CH₄‐C ha^?1 day^?1), and did not differ with treatments, with an average annual net emission of 6.7 g CH₄–C ha^?1 (standard error, 20 g CH₄–C ha^?1). Greatest daily N₂O fluxes occurred when the soil was fallow, and following a series of summer rainfall events. Summer rainfall increased soil water contents and available N, and occurred when soil temperatures were >25 °C, and when there was no active plant growth to compete with soil microorganisms for mineralized N; conditions known to promote N₂O production. The proportion of N fertilizer emitted as N₂O, after correction for emissions from the no N fertilizer treatment, was 0.06%; 17 times lower than IPCC default value for the application of synthetic N fertilizers to land (1.0%). Soil greenhouse gas fluxes from bioenergy crop production in semi‐arid regions are likely to have less influence on the net global warming potential of biofuel production than in temperate climates.     

Effects of experimental nitrogen additions on plant diversity in an old‐growth tropical forest

Response of plant biodiversity to increased availability of nitrogen (N) has been investigated in temperate and boreal forests, which are typically N‐limited, but little is known in tropical forests. We examined the effects of artificial N additions on plant diversity (species richness, density and cover) of the understory layer in an N saturated old‐growth tropical forest in southern China to test the following hypothesis: N additions decrease plant diversity in N saturated tropical forests primarily from N‐mediated changes in soil properties. Experimental additions of N were administered at the following levels from July 2003 to July 2008: no addition (Control); 50 kg N ha^?1 yr^?1 (Low‐N); 100 kg N ha^?1 yr^?1 (Medium‐N), and 150 kg N ha^?1 yr^?1 (High‐N). Results showed that no understory species exhibited positive growth response to any level of N addition during the study period. Although low‐to‐medium levels of N addition (≤100 kg N ha^?1 yr^?1) generally did not alter plant diversity through time, high levels of N addition significantly reduced species diversity. This decrease was most closely related to declines within tree seedling and fern functional groups, as well as to significant increases in soil acidity and Al mobility, and decreases in Ca availability and fine‐root biomass. This mechanism for loss of biodiversity provides sharp contrast to competition‐based mechanisms suggested in studies of understory communities in other forests. Our results suggest that high‐N additions can decrease plant diversity in tropical forests, but that this response may vary with rate of N addition.     

Soil‐derived trace gas fluxes from different energy crops – results from a field experiment in Southwest Germany

Willow coppice, energy maize and Miscanthus were evaluated regarding their soil‐derived trace gas emission potential involving a nonfertilized and a crop‐adapted slow‐release nitrogen (N) fertilizer scheme. The N application rate was 80 kg N ha^?1 yr^?1 for the perennial crops and 240 kg N ha^?1 yr^?1 for the annual maize. A replicated field experiment was conducted with 1‐year measurements of soil fluxes of CH₄, CO₂ and N₂O in weekly intervals using static chambers. The measurements revealed a clear seasonal trend in soil CO₂ emissions, with highest emissions being found for the N‐fertilized Miscanthus plots (annual mean: 50 mg C m^?² h^?1). Significant differences between the cropping systems were found in soil N₂O emissions due to their dependency on amount and timing of N fertilization. N‐fertilized maize plots had highest N₂O emissions by far, which accumulated to 3.6 kg N₂O ha^?1 yr^?1. The contribution of CH₄ fluxes to the total soil greenhouse gas subsumption was very small compared with N₂O and CO₂. CH₄ fluxes were mostly negative indicating that the investigated soils mainly acted as weak sinks for atmospheric CH₄. To identify the system providing the best ratio of yield to soil N₂O emissions, a subsumption relative to biomass yields was calculated. N‐fertilized maize caused the highest soil N₂O emissions relative to dry matter yields. Moreover, unfertilized maize had higher relative soil N₂O emissions than unfertilized Miscanthus and willow. These results favour perennial crops for bioenergy production, as they are able to provide high yields with low N₂O emissions in the field.     

Old trees contribute bio-available nitrogen through canopy bryophytes

Symbiotic cyanobacteria??bryophyte associations on the forest floor are shown to contribute significantly to stand-level nitrogen budgets through the process of biological nitrogen fixation (BNF), but few studies have considered the role of canopy bryophytes. Given the high biomass of epiphytic bryophytes in many tree species of the North American temperate rain forest, we suggest that canopy bryophytes may contribute substantially to stand-level N dynamics. We confirm the presence of cyanobacteria and measure rates of BNF at three heights (0, 15 and 30 m) in Sitka spruce trees across three watershed estuaries of Clayoquot Sound, British Columbia, Canada. This study is the first to report BNF by cyanobacteria associated with epiphytic and forest floor bryophytes in the coastal temperate rain forest of North America. Cyanobacteria density was significantly greater in epiphytic bryophytes compared to mosses on the forest floor, and rates of BNF were highest at 30 m in the canopy. The majority of total stand-level BNF (0.76 kg N · ha^-1 · yr^-1) occurs in the canopy, rather than on the forest floor (0.26 kg N · ha^-1 · yr^-1). We suggest that BNF by cyanobacterial-bryophyte associations in the canopy of coastal temperate rain forests is a unique source of ecosystem N, which is dependent on large, old trees with high epiphytic bryophyte biomass.     

On the tracks of Nitrogen deposition effects on temperate forests at their southern European range – an observational study from Italy

We studied forest monitoring data collected at permanent plots in Italy over the period 2000–2009 to identify the possible impact of nitrogen (N) deposition on soil chemistry, tree nutrition and growth. Average N throughfall (N‐NO₃+N‐NH₄) ranged between 4 and 29 kg ha^?1 yr^?1, with Critical Loads (CLs) for nutrient N exceeded at several sites. Evidence is consistent in pointing out effects of N deposition on soil and tree nutrition: topsoil exchangeable base cations (BCE) and pH decreased with increasing N deposition, and foliar nutrient N ratios (especially N : P and N : K) increased. Comparison between bulk openfield and throughfall data suggested possible canopy uptake of N_, levelling out for bulk deposition >4–6 kg ha^?1 yr^?1. Partial Least Square (PLS) regression revealed that ‐ although stand and meteorological variables explained the largest portion of variance in relative basal area increment (BAI_rel 2000–2009) ‐ N‐related predictors (topsoil BCE, C : N, pH; foliar N‐ratios; N deposition) nearly always improved the BAI_rel model in terms of variance explained (from 78.2 to 93.5%) and error (from 2.98 to 1.50%). N deposition was the strongest predictor even when stand, management and atmosphere‐related variables (meteorology and tropospheric ozone) were accounted for. The maximal annual response of BAI_rel was estimated at 0.074–0.085% for every additional kgN. This corresponds to an annual maximal relative increase of 0.13–0.14% of carbon sequestered in the above‐ground woody biomass for every additional kgN, i.e. a median value of 159 kgC per kgN ha^?1 yr^?1 (range: 50–504 kgC per kgN, depending on the site). Positive growth response occurred also at sites where signals of possible, perhaps recent N saturation were detected. This may suggest a time lag for detrimental N effects, but also that, under continuous high N input, the reported positive growth response may be not sustainable in the long‐term.     

Diazotrophy in Alluvial Meadows of Subarctic River Systems

There is currently limited understanding of the contribution of biological N₂ fixation (diazotrophy) to the N budget of large river systems. This natural source of N in boreal river systems may partially explain the sustained productivity of river floodplains in Northern Europe where winter fodder was harvested for centuries without fertilizer amendments. In much of the world, anthropogenic pollution and river regulation have nearly eliminated opportunities to study natural processes that shaped early nutrient dynamics of large river systems; however, pristine conditions in northern Fennoscandia allow for the retrospective evaluation of key biochemical processes of historical significance. We investigated biological N₂ fixation (diazotrophy) as a potential source of nitrogen fertility at 71 independent floodplain sites along 10 rivers and conducted seasonal and intensive analyses at a subset of these sites. Biological N₂ fixation occurred in all floodplains, averaged 24.5 kg N ha⁻¹ yr⁻¹ and was down regulated from over 60 kg N ha⁻¹ yr⁻¹ to 0 kg N ha⁻¹ yr⁻¹ by river N pollution. A diversity of N₂-fixing cyanobacteria was found to colonize surface detritus in the floodplains. The data provide evidence for N₂ fixation to be a fundamental source of new N that may have sustained fertility at alluvial sites along subarctic rivers. Such data may have implications for the interpretation of ancient agricultural development and the design of contemporary low-input agroecosystems.     

Carbon cycling and storage in world forests: biome patterns related to forest age

Forest age, which is affected by stand‐replacing ecosystem disturbances (such as forest fires, harvesting, or insects), plays a distinguishing role in determining the distribution of carbon (C) pools and fluxes in different forested ecosystems. In this synthesis, net primary productivity (NPP), net ecosystem productivity (NEP), and five pools of C (living biomass, coarse woody debris, organic soil horizons, soil, and total ecosystem) are summarized by age class for tropical, temperate, and boreal forest biomes. Estimates of variability in NPP, NEP, and C pools are provided for each biome‐age class combination and the sources of variability are discussed. Aggregated biome‐level estimates of NPP and NEP were higher in intermediate‐aged forests (e.g., 30–120 years), while older forests (e.g., >120 years) were generally less productive. The mean NEP in the youngest forests (0–10 years) was negative (source to the atmosphere) in both boreal and temperate biomes (?0.1 and –1.9 Mg C ha^?1 yr^?1, respectively). Forest age is a highly significant source of variability in NEP at the biome scale; for example, mean temperate forest NEP was ?1.9, 4.5, 2.4, 1.9 and 1.7 Mg C ha^?1 yr^?1 across five age classes (0–10, 11–30, 31–70, 71–120, 121–200 years, respectively). In general, median NPP and NEP are strongly correlated (R²=0.83) across all biomes and age classes, with the exception of the youngest temperate forests. Using the information gained from calculating the summary statistics for NPP and NEP, we calculated heterotrophic soil respiration (R_h) for each age class in each biome. The mean R_h was high in the youngest temperate age class (9.7 Mg C ha^?1 yr^?1) and declined with age, implying that forest ecosystem respiration peaks when forests are young, not old. With notable exceptions, carbon pool sizes increased with age in all biomes, including soil C. Age trends in C cycling and storage are very apparent in all three biomes and it is clear that a better understanding of how forest age and disturbance history interact will greatly improve our fundamental knowledge of the terrestrial C cycle.     

Greenhouse gas fluxes from an Australian subtropical cropland under long‐term contrasting management regimes

The long‐term effects of conservation management practices on greenhouse gas fluxes from tropical/subtropical croplands remain to be uncertain. Using both manual and automatic sampling chambers, we measured N₂O and CH₄ fluxes at a long‐term experimental site (1968–present) in Queensland, Australia from 2006 to 2009. Annual net greenhouse gas fluxes (NGGF) were calculated from the 3‐year mean N₂O and CH₄ fluxes and the long‐term soil organic carbon changes. N₂O emissions exhibited clear daily, seasonal and interannual variations, highlighting the importance of whole‐year measurement over multiple years for obtaining temporally representative annual emissions. Averaged over 3 years, annual N₂O emissions from the unfertilized and fertilized soils (90 kg N ha^?1 yr^?1 as urea) amounted to 138 and 902 g N ha^?1, respectively. The average annual N₂O emissions from the fertilized soil were 388 g N ha^?1 lower under no‐till (NT) than under conventional tillage (CT) and 259 g N ha^?1 higher under stubble retention (SR) than under stubble burning (SB). Annual N₂O emissions from the unfertilized soil were similar between the contrasting tillage and stubble management practices. The average emission factors of fertilizer N were 0.91%, 1.20%, 0.52% and 0.77% for the CT‐SB, CT‐SR, NT‐SB and NT‐SR treatments, respectively. Annual CH₄ fluxes from the soil were very small (?200–300 g CH₄ ha^?1 yr^?1) with no significant difference between treatments. The NGGF were 277–350 kg CO₂‐e ha^?1 yr^?1 for the unfertilized treatments and 401–710 kg CO₂‐e ha^?1 yr^?1 for the fertilized treatments. Among the fertilized treatments, N₂O emissions accounted for 52–97% of NGGF and NT‐SR resulted in the lowest NGGF (401 kg CO₂‐e ha^?1 yr^?1 or 140 kg CO₂‐e t^?1 grain). Therefore, NT‐SR with improved N fertilizer management practices was considered the most promising management regime for simultaneously achieving maximal yield and minimal NGGF.     

Long‐term impacts of manure amendments on carbon and greenhouse gas dynamics of rangelands

Livestock manure is applied to rangelands as an organic fertilizer to stimulate forage production, but the long‐term impacts of this practice on soil carbon (C) and greenhouse gas (GHG) dynamics are poorly known. We collected soil samples from manured and nonmanured fields on commercial dairies and found that manure amendments increased soil C stocks by 19.0 ± 7.3 Mg C ha^?1 and N stocks by 1.94 ± 0.63 Mg N ha^?1 compared to nonmanured fields (0–20 cm depth). Long‐term historical (1700–present) and future (present–2100) impacts of management on soil C and N dynamics, net primary productivity (NPP), and GHG emissions were modeled with DayCent. Modeled total soil C and N stocks increased with the onset of dairying. Nitrous oxide (N₂O) emissions also increased by ~2 kg N₂O‐N ha^?1 yr^?1. These emissions were proportional to total N additions and offset 75–100% of soil C sequestration. All fields were small net methane (CH₄) sinks, averaging ?4.7 ± 1.2 kg CH₄‐C ha^?1 yr^?1. Overall, manured fields were net GHG sinks between 1954 and 2011 (?0.74 ± 0.73 Mg CO₂ e ha^?1 yr^?1, CO₂e are carbon dioxide equivalents), whereas nonmanured fields varied around zero. Future soil C pools stabilized 40–60 years faster in manured fields than nonmanured fields, at which point manured fields were significantly larger sources than nonmanured fields (1.45 ± 0.52 Mg CO₂e ha^?1 yr^?1 and 0.51 ± 0.60 Mg CO₂e ha^?1 yr^?1, respectively). Modeling also revealed a large background loss of soil C from the passive soil pool associated with the shift from perennial to annual grasses, equivalent to 29.4 ± 1.47 Tg CO₂e in California between 1820 and 2011. Manure applications increased NPP and soil C storage, but plant community changes and GHG emissions decreased, and eventually eliminated, the net climate benefit of this practice.