Dry deposition of ammonia gas drives species change faster than wet deposition of ammonium ions: evidence from a long‐term field manipulation

Although the effects of atmospheric nitrogen deposition on species composition are relatively well known, the roles of the different forms of nitrogen, in particular gaseous ammonia (NH₃), have not been tested in the field. Since 2002, we have manipulated the form of N deposition to an ombrotrophic bog, Whim, on deep peat in southern Scotland, with low ambient N (wet + dry = 8 kg N ha^?1 yr^?1) and S (4 kg S ha^?1 yr^?1) deposition. A gradient of ammonia (NH₃, dry N), from 70 kg N ha^?1 yr^?1 down to background, 3–4 kg N ha^?1 yr^?1 was generated by free air release. Wet ammonium (NH₄⁺, wet N) was provided to replicate plots in a fine rainwater spray (NH₄Cl at +8, +24, +56 kg N ha^?1 yr^?1). Automated treatments are coupled to meteorological conditions, in a globally unique, field experiment. Ammonia concentrations were converted to NH₃‐N deposition (kg N ha^?1) using a site/vegetation specific parameterization. Within 3 years, exposure to relatively modest deposition of NH₃, 20–56 kg NH₃‐N ha^?1 yr^?1 led to dramatic reductions in species cover, with almost total loss of Calluna vulgaris, Sphagnum capillifolium and Cladonia portentosa. These effects appear to result from direct foliar uptake and interaction with abiotic and biotic stresses, rather than via effects on the soil. Additional wet N by contrast, significantly increased Calluna cover after 5 years at the 56 kg N dose, but reduced cover of Sphagnum and Cladonia. Cover reductions caused by wet N were significantly different from and much smaller than those caused by equivalent dry N doses. The effects of gaseous NH₃ described here, highlight the potential for ammonia to destroy acid heathland and peat bog ecosystems. Separating the effects of gaseous ammonia and wet ammonium deposition, for a peat bog, has significant implications for regulatory bodies and conservation agencies.     

High retention of 15N‐labeled nitrogen deposition in a nitrogen saturated old‐growth tropical forest

The effects of increased reactive nitrogen (N) deposition in forests depend largely on its fate in the ecosystems. However, our knowledge on the fates of deposited N in tropical forest ecosystems and its retention mechanisms is limited. Here, we report the results from the first whole ecosystem ¹⁵N labeling experiment performed in a N‐rich old‐growth tropical forest in southern China. We added ¹⁵N tracer monthly as ¹⁵NH₄¹⁵NO₃ for 1 year to control plots and to N‐fertilized plots (N‐plots, receiving additions of 50 kg N ha^?1 yr^?1 for 10 years). Tracer recoveries in major ecosystem compartments were quantified 4 months after the last addition. Tracer recoveries in soil solution were monitored monthly to quantify leaching losses. Total tracer recovery in plant and soil (N retention) in the control plots was 72% and similar to those observed in temperate forests. The retention decreased to 52% in the N‐plots. Soil was the dominant sink, retaining 37% and 28% of the labeled N input in the control and N‐plots, respectively. Leaching below 20 cm was 50 kg N ha^?1 yr^?1 in the control plots and was close to the N input (51 kg N ha^?1 yr^?1), indicating N saturation of the top soil. Nitrogen addition increased N leaching to 73 kg N ha^?1 yr^?1. However, of these only 7 and 23 kg N ha^?1 yr^?1 in the control and N‐plots, respectively, originated from the labeled N input. Our findings indicate that deposited N, like in temperate forests, is largely incorporated into plant and soil pools in the short term, although the forest is N‐saturated, but high cycling rates may later release the N for leaching and/or gaseous loss. Thus, N cycling rates rather than short‐term N retention represent the main difference between temperate forests and the studied tropical forest.     

A long‐term nitrogen fertilizer gradient has little effect on soil organic matter in a high‐intensity maize production system

Global maize production alters an enormous soil organic C (SOC) stock, ultimately affecting greenhouse gas concentrations and the capacity of agroecosystems to buffer climate variability. Inorganic N fertilizer is perhaps the most important factor affecting SOC within maize‐based systems due to its effects on crop residue production and SOC mineralization. Using a continuous maize cropping system with a 13 year N fertilizer gradient (0–269 kg N ha^?1 yr^?1) that created a large range in crop residue inputs (3.60–9.94 Mg dry matter ha^?1 yr^?1), we provide the first agronomic assessment of long‐term N fertilizer effects on SOC with direct reference to N rates that are empirically determined to be insufficient, optimum, and excessive. Across the N fertilizer gradient, SOC in physico‐chemically protected pools was not affected by N fertilizer rate or residue inputs. However, unprotected particulate organic matter (POM) fractions increased with residue inputs. Although N fertilizer was negatively linearly correlated with POM C/N ratios, the slope of this relationship decreased from the least decomposed POM pools (coarse POM) to the most decomposed POM pools (fine intra‐aggregate POM). Moreover, C/N ratios of protected pools did not vary across N rates, suggesting little effect of N fertilizer on soil organic matter (SOM) after decomposition of POM. Comparing a N rate within 4% of agronomic optimum (208 kg N ha^?1 yr^?1) and an excessive N rate (269 kg N ha^?1 yr^?1), there were no differences between SOC amount, SOM C/N ratios, or microbial biomass and composition. These data suggest that excessive N fertilizer had little effect on SOM and they complement agronomic assessments of environmental N losses, that demonstrate N₂O and NO₃ emissions exponentially increase when agronomic optimum N is surpassed.     

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.     

Nitrogen to phosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests

Increased atmospheric carbon dioxide (CO₂) concentrations and nitrogen (N) deposition induced by human activities have greatly influenced the stoichiometry of N and phosphorus (P). We used model forest ecosystems in open‐top chambers to study the effects of elevated CO₂ (ca. 700 μmol mol^?1) alone and together with N addition (100 kg N ha^?1 yr^?1) on N to P (N : P) ratios in leaves, stems and roots of five tree species, including four non‐N₂ fixers and one N₂ fixer, in subtropical China from 2006 to 2009. Elevated CO₂ decreased or had no effects on N : P ratios in plant tissues of tree species. N addition, especially under elevated CO₂, lowered N : P ratios in the N₂ fixer, and this effect was significant in the stems and the roots. However, only one species of the non‐N₂ fixers showed significantly lower N : P ratios under N addition in 2009, and the others were not affected by N addition. The reductions of N : P ratios in response to elevated CO₂ and N addition were mainly associated with the increases in P concentrations. Our results imply that elevated CO₂ and N addition could facilitate tree species to mitigate P limitation by more strongly influencing P dynamics than N in the subtropical forests.     

Nitrogen input together with ecosystem nitrogen enrichment predict nitrate leaching from European forests

The IFEF database (Indicators of Forest Ecosystem Functioning), consisting of nitrogen deposition, nitrate leaching fluxes, and soil and ecosystem characteristics, is analysed to evaluate the C/N ratio in the organic horizon as an indicator of nitrate leaching. One hundred and eighty one forests are examined, from countries across Europe ranging from boreal to Mediterranean regions, encompassing broadleaf and coniferous sites and plot and catchment studies. N input in throughfall ranges from less than 1 kg N ha^?1 y^?1 in northern Norway and Finland to greater than 60 kg N ha^?1 y^?1 in the Netherlands and Czech Republic. The amount of NO₃^– leached covers a smaller range, between 1 and 40 kg N ha^?1 y^?1. Nitrate leaching is strongly dependent on the amount of nitrogen deposited in throughfall (N input) and simply adding the C/N ratio in the organic horizon to a regression equation does not improve this relationship. However, when the data are stratified based on C/N ratios less than or equal to 25 and greater than 25, highly significant relationships (P < 0.05) are observed between N input and NO₃^– leached. The slope of the relationship for those sites where C/N ratio is ≤ 25 (′nitrogen enriched′ sites) is twice that for those sites where C/N ratio is > 25. These empirical relationships may be used to identify which forested ecosystems are likely to show elevated rates of nitrate leaching under predicted future nitrogen deposition scenarios. Elevated NO₃^– leaching also shows a relationship with soil pH, with high rates of NO₃^– leaching only observed at sites with a pH < 4.5 and N inputs > 30 kg N ha^?1 y^?1. Tree age and species have no significant impact on the ecosystem response to N input at a regional scale.     

Isotopic evidence for the occurrence of biological nitrification and nitrogen deposition processing in forest canopies

This study examines the role of tree canopies in processing atmospheric nitrogen (N_dep) for four forests in the United Kingdom subjected to different N_dep: Scots pine and beech stands under high N_dep (HN, 13–19 kg N ha^?1 yr^?1), compared to Scots pine and beech stands under low N_dep (LN, 9 kg N ha^?1 yr^?1). Changes of NO₃‐N and NH₄‐N concentrations in rainfall (RF) and throughfall (TF) together with a quadruple isotope approach, which combines δ¹⁸O, Δ¹⁷O and δ¹⁵N in NO₃^? and δ¹⁵N in NH₄⁺, were used to assess N transformations by the canopies. Generally, HN sites showed higher NH₄‐N and NO₃‐N concentrations in RF compared to the LN sites. Similar values of δ¹⁵N‐NO₃^? and δ¹⁸O in RF suggested similar source of atmospheric NO₃^? (i.e. local traffic), while more positive values for δ¹⁵N‐NH₄⁺ at HN compared to LN likely reflected the contribution of dry NH_x deposition from intensive local farming. The isotopic signatures of the N‐forms changed after interacting with tree canopies. Indeed, ¹⁵N‐enriched NH₄⁺ in TF compared to RF at all sites suggested that canopies played an important role in buffering dry N_dep also at the low N_dep site. Using two independent methods, based on δ¹⁸O and Δ¹⁷O, we quantified for the first time the proportion of NO₃^? in TF, which derived from nitrification occurring in tree canopies at the HN site. Specifically, for Scots pine, all the considered isotope approaches detected biological nitrification. By contrast for the beech, only using the mixing model with Δ¹⁷O, we were able to depict the occurrence of nitrification within canopies. Our study suggests that tree canopies play an active role in the N cycling within forest ecosystems. Processing of N_dep within canopies should not be neglected and needs further exploration, with the combination of multiple isotope tracers, with particular reference to Δ¹⁷O.     

Can intensification reduce emission intensity of biofuel through optimized fertilizer use? Theory and the case of oil palm in Indonesia

Closing yield gaps through higher fertilizer use increases direct greenhouse gas emissions but shares the burden over a larger production volume. Net greenhouse gas (GHG) footprints per unit product under agricultural intensification vary depending on the context, scale and accounting method. Life cycle analysis of footprints includes attributable emissions due to (i) land conversion (‘fixed cost’); (ii) external inputs used (‘variable cost’); (iii) crop production (‘agronomic efficiency’); and (iv) postharvest transport and processing (‘proportional’ cost). The interplay between fixed and variable costs results in a nuanced opportunity for intermediate levels of intensification to minimize footprints. The fertilizer level that minimizes the footprint may differ from the economic optimum. The optimization problem can be solved algebraically for quadratic crop fertilizer response equations. We applied this theory to data of palm oil production and fertilizer use from 23 plantations across the Indonesian production range. The current EU threshold requiring at least 35% emission saving for biofuel use can never be achieved by palm oil if produced: (i) on peat soils, or (ii) on mineral soils where the C debt due to conversion is larger than 20 Mg C ha^?1, if the footprint is calculated using an emission ratio of N₂O–N/N fertilizer of 4%. At current fertilizer price levels in Indonesia, the economically optimized N fertilizer rate is 344–394 kg N ha^?1, while the reported mean N fertilizer rate is 141 kg N ha^?1 yr^?1 and rates of 74–277 kg N ha^?1 would minimize footprints, for a N₂O–N/N fertilizer ratio of 4–1%, respectively. At a C debt of 30 Mg C ha^?1, these values are 200–310 kg N ha^?1. Sustainable weighting of ecology and economics would require a higher fertilizer/yield price ratio, depending on C debt. Increasing production by higher fertilizer use from current 67% to 80% of attainable yields would not decrease footprints in current production conditions.     

Nitrogen Addition Shapes Soil Phosphorus Availability in Two Reforested Tropical Forests in Southern China

Scant information is available on how soil phosphorus (P) availability responds to atmospheric nitrogen (N) deposition, especially in the tropical zones. This study examined the effect of N addition on soil P availability, and compared this effect between forest sites of contrasting land‐use history. Effects of N addition on soil properties, litterfall production, P release from decomposing litter, and soil P availability were studied in a disturbed (reforested pine forest with previous understory vegetation and litter harvesting) and a rehabilitated (reforested mixed pine/broadleaf forest with no understory vegetation and litter harvesting) tropical forest in southern China. Experimental N‐treatments (above ambient) were the following: Control (no N addition), N50 (50 kg N ha^?1 yr^?1), and N100 (100 kg N ha^?1 yr^?1). Results indicated that N addition significantly decreased soil P availability in the disturbed forest. In the rehabilitated forest, however, soil P availability was significantly increased by N addition. Decreases in soil P availability may be correlated with decreases in rates of P release from decomposing litter in the N‐treated plots, whereas the increase in soil P availability was correlated with an increase in litterfall production. Our results suggest that response of soil P availability to N deposition in the reforested tropical forests in southern China may vary greatly with temporal changes in tree species composition and soil nutrient status, caused by different land‐use practices.     

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.     

Nitrate leaching and soil nitrous oxide emissions diminish with time in a hybrid poplar short‐rotation coppice in southern Germany

Hybrid poplar short‐rotation coppices (SRC) provide feedstocks for bioenergy production and can be established on lands that are suboptimal for food production. The environmental consequences of deploying this production system on marginal agricultural land need to be evaluated, including the investigation of common management practices i.e., fertilization and irrigation. In this work, we evaluated (1) the soil‐atmosphere exchange of carbon dioxide, methane, and nitrous oxide (N₂O); (2) the changes in soil organic carbon (SOC) stocks; (3) the gross ammonification and nitrification rates; and (4) the nitrate leaching as affected by the establishment of a hybrid poplar SRC on a marginal agricultural land in southern Germany. Our study covered one 3‐year rotation period and 2 years after the first coppicing. We combined field and laboratory experiments with modeling. The soil N₂O emissions decreased from 2.2 kg N₂O‐N ha^?1 a^?1 in the year of SRC establishment to 1.1–1.4 kg N₂O‐N ha^?1 a^?1 after 4 years. Likewise, nitrate leaching reduced from 13 to 1.5–8 kg N ha^?1 a^?1. Tree coppicing induced a brief pulse of soil N₂O flux and marginal effects on gross N turnover rates. Overall, the N losses diminished within 4 years by 80% without fertilization (irrespective of irrigation) and by 40% when 40–50 kg N ha^?1 a^?1 were applied. Enhanced N losses due to fertilization and the minor effect of fertilization and irrigation on tree growth discourage its use during the first rotation period after SRC establishment. A SOC accrual rate of 0.4 Mg C ha^?1 a^?1 (uppermost 25 cm, P = 0.2) was observed 5 years after the SRC establishment. Overall, our data suggest that SRC cultivation on marginal agricultural land in the region is a promising option for increasing the share of renewable energy sources due to its net positive environmental effects.     

Productivity, 15N dynamics and water use efficiency in low‐ and high‐input switchgrass systems

Sustainable and environmentally benign switchgrass production systems need to be developed for switchgrass to become a large‐scale dedicated energy crop. An experiment was conducted in California from 2009 to 2011 to determine the sustainability of low‐ and high‐input irrigated switchgrass systems as a function of yield, irrigation requirement, crop N removal, N translocation from aboveground (AG) to belowground (BG) biomass during senescence, and fertilizer ¹⁵N recovery (FNR) in the AG and BG biomass (0–300 cm), and soil (0–300 cm). The low‐input system consisted of a single‐harvest (mid‐fall) irrigated until flowering (early summer), while the high‐input system consisted of a two‐harvest system (early summer and mid‐fall) irrigated throughout the growing season. Three N fertilization rates (0, 100, and 200 kg N ha^?1 yr^?1) were applied as subtreatments in a single application in the spring of each year. A single pulse of ¹⁵N enriched fertilizer was applied in the first year of the study to micro‐plots within the 100 kg N ha^?1 subplots. Average yields across years under optimal N rates (100 and 200 kg ha^?1 yr^?1 for low‐ and high‐input systems, respectively) were 20.7 and 24.8 Mg ha^?1. However, the low input (372 ha mm) required 47% less irrigation than the high‐input system (705 ha mm) and achieved higher irrigation use efficiency. In addition, the low‐input system had 46% lower crop N removal, 53% higher N stored in BG biomass, and a positive N balance, presumably due to 49% of ¹⁵N translocation from AG to BG biomass during senescence. Furthermore, at the end of 3 years, the low‐input system had lower fertilizer ¹⁵N removed by harvest (26%) and higher FNR remaining in the system in BG biomass plus soil (31%) than the high‐input system (45% and 21%, respectively). Based on these findings, low‐input systems are more sustainable than high‐input systems in irrigated Mediterranean climates.     

Nitrogen deposition and forest nitrogen cycling along an urban–rural transect in southern China

There is increasing concern over the impact of atmospheric nitrogen (N) deposition on forest ecosystems in the tropical and subtropical areas. In this study, we quantified atmospheric N deposition and revealed current plant and soil N status in 14 forests along a 150 km urban to rural transect in southern China, with an emphasis on examining whether foliar δ¹⁵N can be used as an indicator of N saturation. Bulk deposition ranged from 16.2 to 38.2 kg N ha^?1 yr^?1, while the throughfall covered a larger range of 11.7–65.1 kg N ha^?1 yr^?1. Foliar N concentration, NO₃^? leaching to stream, and soil NO₃^? concentration were low and NO₃^? production was negligible in some rural forests, indicating that primary production in these forests may be limited by N supply. But all these N variables were enhanced in suburban and urban forests. Across the study transect, throughfall N input was correlated positively with soil nitrification and NO₃^? leaching to stream, and negatively with pH values in soil and stream water. Foliar δ¹⁵N was between ?6.6‰ and 0.7‰, and was negatively correlated with soil NO₃^? concentration and NO₃^? leaching to stream across the entire transect, demonstrating that an increased N supply does not necessarily increase forest δ¹⁵N values. We proposed several potential mechanism that could contribute to the δ¹⁵N pattern, including (1) increased plant uptake of ¹⁵N‐depleted soil NO₃^?, (2) foliage uptake of ¹⁵N‐depleted NH₄⁺, (3) increased utilization of soil inorganic N relative to dissolved organic N, and (4) increased fractionation during plant N uptake under higher soil N availability.     

Simulating switchgrass biomass production across ecoregions using the DAYCENT model

The production potential of switchgrass (Panicum virgatum L.) has not been estimated in a Mediterranean climate on a regional basis and its economic and environmental contribution as a biofuel crop remains unknown. The objectives of the study were to calibrate and validate a biogeochemical model, DAYCENT, and to predict the biomass yield potential of switchgrass across the Central Valley of California. Six common cultivars were calibrated using published data across the US and validated with data generated from four field trials in California (2007–2009). After calibration, the modeled range of yields across the cultivars and various management practices in the US (excluding California) was 2.4–41.2 Mg ha^?1 yr^?1, generally compatible with the observed yield range of 1.3–33.7 Mg ha^?1 yr^?1. Overall, the model was successfully validated in California; the model explained 66–90% of observed yield variation in 2007–2009. The range of modeled yields was 2.0–41.4 Mg ha^?1 yr^?1, which corresponded to the observed range of 1.3–41.1 Mg ha^?1 yr^?1. The response to N fertilizer and harvest frequency on yields were also reasonably validated. The model estimated that Alamo (21–23 Mg ha^?1 yr^?1) and Kanlow (22–24 Mg ha^?1 yr^?1) had greatest yield potential during the years after establishment. The effects of soil texture on modeled yields tended to be consistent for all cultivars, but there were distinct climatic (e.g., annual mean maximum temperature) controls among the cultivars. Our modeled results suggest that early stand maintenance of irrigated switchgrass is strongly dependent on available soil N; estimated yields increased by 1.6–5.5 Mg ha^?1 yr^?1 when residual soil mineral N was sufficient for optimal re‐growth. Therefore, management options of switchgrass for regional biomass production should be ecotype‐specific and ensure available soil N maintenance.     

Combination of herbivore removal and nitrogen deposition increases upland carbon storage

Ecosystem carbon (C) accrual and storage can be enhanced by removing large herbivores as well as by the fertilizing effect of atmospheric nitrogen (N) deposition. These drivers are unlikely to operate independently, yet their combined effect on aboveground and belowground C storage remains largely unexplored. We sampled inside and outside 19 upland grazing exclosures, established for up to 80 years, across an N deposition gradient (5–24 kg N ha^?1 yr^?1) and found that herbivore removal increased aboveground plant C stocks, particularly in moss, shrubs and litter. Soil C storage increased with atmospheric N deposition, and this was moderated by the presence or absence of herbivores. In exclosures receiving above 11 kg N ha^?1 year^?1, herbivore removal resulted in increased soil C stocks. This effect was typically greater for exclosures dominated by dwarf shrubs (Calluna vulgaris) than by grasses (Molinia caerulea). The same pattern was observed for ecosystem C storage. We used our data to predict C storage for a scenario of removing all large herbivores from UK heathlands. Predictions were made considering herbivore removal only (ignoring N deposition) and the combined effects of herbivore removal and current N deposition rates. Predictions including N deposition resulted in a smaller increase in UK heathland C storage than predictions using herbivore removal only. This finding was driven by the fact that the majority of UK heathlands receive low N deposition rates at which herbivore removal has little effect on C storage. Our findings demonstrate the crucial link between herbivory by large mammals and atmospheric N deposition, and this interaction needs to be considered in models of biogeochemical cycling.     

Comparing annual and perennial crops for bioenergy production – influence on nitrate leaching and energy balance

Production of energy crops is promoted as a means to mitigate global warming by decreasing dependency on fossil energy. However, agricultural production of bioenergy can have various environmental effects depending on the crop and production system. In a field trial initiated in 2008, nitrate concentration in soil water was measured below winter wheat, grass‐clover and willow during three growing seasons. Crop water balances were modelled to estimate the amount of nitrate leached per hectare. In addition, dry matter yields and nitrogen (N) yields were measured, and N balances and energy balances were calculated. In willow, nitrate concentrations were up to approximately 20 mg l^?1 nitrate‐N during the establishment year, but declined subsequently to <5 mg l^?1 nitrate‐N, resulting in an annual N leaching loss of 18, 3 and 0.3 kg ha^?1 yr^?1 N in the first 3 years after planting. A similar trend was observed in grass‐clover where concentrations stabilized at 2–4 mg l^?1 nitrate‐N from the beginning of the second growing season, corresponding to leaching of approximately 5 kg ha^?1 yr^?1 N. In winter wheat, an annual N leaching loss of 36–68 kg ha^?1 yr^?1 was observed. For comparison, nitrate leaching was also measured in an old willow crop established in 1996 from which N leaching ranged from 6 to 27 kg ha^?1 yr^?1. Dry matter yields ranged between 5.9 and 14.8 Mg yr^?1 with lowest yield in the newly established willow and the highest yield harvested in grass‐clover. Grass‐clover gave the highest net energy yield of 244 GJ ha^?1 yr^?1, whereas old willow, winter wheat and first rotation willow gave net energy yields of 235, 180 and 105 GJ ha^?1 yr^?1. The study showed that perennial crops can provide high energy yields and significantly reduce N losses compared to annual crops.     

Regional scale gradients of climate and nitrogen deposition drive variation in ectomycorrhizal fungal communities associated with native Scots pine

Ectomycorrhizal fungi commonly associate with the roots of forest trees where they enhance nutrient and water uptake, promote seedling establishment and have an important role in forest nutrient cycling. Predicting the response of ectomycorrhizal fungi to environmental change is an important step to maintaining forest productivity in the future. These predictions are currently limited by an incomplete understanding of the relative significance of environmental drivers in determining the community composition of ectomycorrhizal (ECM) fungi at large spatial scales. To identify patterns of community composition in ECM fungi along regional scale gradients of climate and nitrogen deposition in Scotland, fungal communities were analysed from 15 seminatural Scots pine (Pinus sylvestris L.) forests. Fungal taxa were identified by sequencing of the ITS rDNA region using fungal‐specific primers. Nonmetric multidimensional scaling was used to assess the significance of 16 climatic, pollutant and edaphic variables on community composition. Vector fitting showed that there was a strong influence of rainfall and soil moisture on community composition at the species level, and a smaller impact of temperature on the abundance of ectomycorrhizal exploration types. Nitrogen deposition was also found to be important in determining community composition, but only when the forest experiencing the highest deposition (9.8 kg N ha^?1 yr^?1) was included in the analysis. This finding supports previously published critical load estimates for ectomycorrhizal fungi of 5–10 kg N ha^?1 yr^?1. This work demonstrates that both climate and nitrogen deposition can drive gradients of fungal community composition at a regional scale.     

Are the benefits of yield responses to nitrogen fertilizer application in the bioenergy crop Miscanthus × giganteus offset by increased soil emissions of nitrous oxide?

A field trial was carried out on a 15 year old Miscanthus stand, subject to nitrogen fertilizer treatments of 0, 63 and 125 kg‐N ha^?1, measuring N₂O emissions, as well as annual crop yield over a full year. N₂O emission intensity (N₂O emissions calculated as a function of above‐ground biomass) was significantly affected by fertilizer application, with values of 52.2 and 59.4 g N₂O‐N t^?1 observed at 63 and 125 kg‐N ha^?1, respectively, compared to 31.3 g N₂O‐N t^?1 in the zero fertilizer control. A life cycle analyses approach was applied to calculate the increase in yield required to offset N₂O emissions from Miscanthus through fossil fuel substitution in the fuel chain. For the conditions observed during the field trial yield increases of 0.33 and 0.39 t ha^?1 were found to be required to offset N₂O emissions from the 63 kg‐N ha^?1 treatment, when replacing peat and coal, respectively, while increases of 0.71 and 0.83 t ha^?1 were required for the 125 kg‐N ha^?1 treatment, for each fuel. These values are considerably less than the mean above‐ground biomass yield increases observed here of 1.57 and 2.79 t ha^?1 at fertilization rates 63 and 125 kg‐N ha^?1 respectively. Extending this analysis to include a range of fertilizer application rates and N₂O emission factors found increases in yield necessary to offset soil N₂O emissions ranging from 0.26 to 2.54 t ha^?1. These relatively low yield increase requirements indicate that where nitrogen fertilizer application improves yield, the benefits of such a response will not be offset by soil N₂O emissions.     

Major changes in forest carbon and nitrogen cycling caused by declining sulphur deposition

Sulphur (S) and nitrogen (N) deposition are important drivers of the terrestrial carbon (C) and N cycling. We analyzed changes in C and N pools in soil and tree biomass at a highly acidified spruce site in the Czech Republic during a 15 year period. Total S deposition decreased from 5 to 1.1 g m^?2 yr^?1 between 1995 and 2009, whereas bulk N deposition did not change. Over the same period, C and N pools in the Oa horizon declined by 116 g C and 4.2 g N m^?2 yr^?1, a total decrease of 47% and 42%, respectively. This loss of C and N probably originated from organic matter (OM) that had accumulated during the period of high acid deposition when litter decomposition was suppressed. The loss of OM from the Oa horizon coincided with a substantial leaching (1.3 g N m^?2 yr^?1 at 90 cm) in the 1990s to almost no leaching (<0.02 g N m^?2 yr^?1) since 2006. Forest floor net N mineralization also decreased. This had consequences for spruce needle N concentration (from 17.1 to 11.4 mg kg^?1 in current needles), an increase in litterfall C/N ratio (from 51 to 63), and a significant increase in the Oi + Oe horizon C/N ratio (from 23.4 to 27.3) between 1994 and 2009/2010. Higher forest growth and lower canopy defoliation was observed in the 2000s compared to the 1990s. Our results demonstrate that reducing S deposition has had a profound impact on forest organic matter cycling, leading to a reversal of historic ecosystem N enrichment, cessation of nitrate leaching, and a major loss of accumulated organic soil C and N stocks. These results have major implications for our understanding of the controls on both N saturation and C sequestration in forests, and other ecosystems, subjected to current or historic S deposition.     

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.