Nitrogen oxide gas emissions from temperate forest soils receiving long-term nitrogen inputs

From spring 2000 through fall 2001, we measured nitric oxide (NO) and nitrous oxide (N₂O) fluxes in two temperate forest sites in Massachusetts, USA that have been treated since 1988 with different levels of nitrogen (N) to simulate elevated rates of atmospheric N deposition. Plots within a pine stand that were treated with either 50 or 150 kg N ha^?1 yr^?1 above background displayed consistently elevated NO fluxes (100–200 µg N m^?2 h^?1) compared to control plots, while only the higher N treatment plot within a mixed hardwood stand displayed similarly elevated NO fluxes. Annual NO emissions estimated from monthly sampling accounted for 3.0–3.7% of N inputs to the high‐N plots and 8.3% of inputs to the Pine low‐N plot. Nitrous oxide fluxes in the N‐treated plots were generally < 10% of NO fluxes. Net nitrification rates (NRs) and NO production rates measured in the laboratory displayed patterns that were consistent with field NO fluxes. Total N oxide gas flux was positively correlated with contemporaneous measurements of NR and concentration. Acetylene inhibited both nitrification and NO production, indicating that autotrophic nitrification was responsible for the elevated NO production. Soil pH was negatively correlated with N deposition rate. Low levels (3–11 µg N kg^?1) of nitrite () were detected in mineral soils from both sites. Kinetic models describing NO production as a function of the protonated form of (nitrous acid [HNO₂]) adequately described the mineral soil data. The results indicate that atmospheric deposition may generate losses of gaseous NO from forest soils by promoting nitrification, and that the response may vary significantly between forest types under similar climatic regimes. The lowering of pH resulting from nitrification and/or directly from deposition may also play a role by promoting reactions involving HNO₂.     

Throughfall exclusion and leaf litter addition drive higher rates of soil nitrous oxide emissions from a lowland wet tropical forest

Tropical forests are a significant global source of the greenhouse gas nitrous oxide (N₂O). Predicted environmental changes for this biome highlight the need to understand how simultaneous changes in precipitation and labile carbon (C) availability may affect soil N₂O production. We conducted a small‐scale throughfall and leaf litter manipulation in a lowland tropical forest in southwestern Costa Rica to test how potential changes in both water and litter derived labile C inputs to soils may alter N₂O emissions. Experimentally reducing throughfall in this wet tropical forest significantly increased soil emissions of N₂O, and our data suggest that at least part of this response was driven by an increase in the concentration of dissolved organic carbon [DOC] inputs delivered from litter to soil under the drier conditions. Furthermore, [DOC] was significantly correlated with N₂O emissions across both throughfall and litterfall manipulation plots, despite the fact that native NO₃^? pools in this site were generally small. Our results highlight the importance of understanding not only the potential direct effects of changing precipitation on soil biogeochemistry, but also the indirect effects resulting from interactions between the hydrologic, C and N cycles. Finally, over all sampling events we observed lower mean N₂O emissions (<1 ng N₂O‐N cm^?2 h^?1) than reported for many other lowland tropical forests, perhaps reflecting a more general pattern of increasing relative N constraints to biological activity as one moves from drier to wetter portions of the lowland tropical forest biome.     

Nitrous oxide emissions from silage maize fields under different mineral nitrogen fertilizer and slurry applications

Intensive dairy farming systems are a large source of emission of the greenhouse gas nitrous oxide (N₂O), because of high nitrogen (N) application rates to grasslands and silage maize fields. The objective of this study was to compare measured N₂O emissions from two different soils to default N₂O emission factors, and to look at alternative emission factors based on (i) the N uptake in the crop and (ii) the N surplus of the system, i.e., N applied minus N uptake by the crop. Twelve N fertilization regimes were implemented on a sandy soil (typic endoaquoll) and a clay soil (typic endoaquept) in the Netherlands, and N₂O emissions were measured throughout the growing season. Highest cumulative fluxes of 1.92 and 6.81 kg N₂O-N ha^–1 for the sandy soil and clay soil were measured at the highest slurry application rate of 250 kg N ha^–1. Background emissions from unfertilized soils were 0.14 and 1.52 kg N₂O-N ha^–1 for the sandy soil and the clay soil, respectively. Emission factors for the sandy soil averaged 0.08, 0.51 and 0.26% of the N applied via fertilizer, slurry, and combinations of both. For the clay soil, these numbers were 1.18, 1.21 and 1.69%, respectively. Surplus N was linearly related to N₂O emission for both the sandy soil (R²=0.60) and the clay soil (R²=0.40), indicating a possible alternative emission factor. We concluded that, in our study, N₂O emission was not linearly related to N application rates, and varied with type and application rate of fertilizer. Finally, the relatively high emission from the clay soil indicates that background emissions might have to be taken into account in N₂O budgets.     

Denitrification and nitrous oxide emissions from riparian forests soils exposed to prolonged nitrogen runoff

Compared to upland forests, riparian forest soils have greater potential to remove nitrate (NO₃) from agricultural runoff through denitrification. It is unclear, however, whether prolonged exposure of riparian soils to nitrogen (N) loading will affect the rate of denitrification and its end products. This research assesses the rate of denitrification and nitrous oxide (N₂O) emissions from riparian forest soils exposed to prolonged nutrient runoff from plant nurseries and compares these to similar forest soils not exposed to nutrient runoff. Nursery runoff also contains high levels of phosphate (PO₄). Since there are conflicting reports on the impact of PO₄ on the activity of denitrifying microbes, the impact of PO₄ on such activity was also investigated. Bulk and intact soil cores were collected from N-exposed and non-exposed forests to determine denitrification and N₂O emission rates, whereas denitrification potential was determined using soil slurries. Compared to the non-amended treatment, denitrification rate increased 2.7- and 3.4-fold when soil cores collected from both N-exposed and non-exposed sites were amended with 30 and 60 μg NO₃-N g⁻¹ soil, respectively. Net N₂O emissions were 1.5 and 1.7 times higher from the N-exposed sites compared to the non-exposed sites at 30 and 60 μg NO₃-N g⁻¹ soil amendment rates, respectively. Similarly, denitrification potential increased 17 times in response to addition of 15 μg NO₃-N g⁻¹ in soil slurries. The addition of PO₄ (5 μg PO₄-P g⁻¹) to soil slurries and intact cores did not affect denitrification rates. These observations suggest that prolonged N loading did not affect the denitrification potential of the riparian forest soils; however, it did result in higher N₂O emissions compared to emission rates from non-exposed forest soils.     

Estimation of nitrous oxide, nitric oxide and ammonia emissions from croplands in East, Southeast and South Asia

Agricultural activities have greatly altered the global nitrogen (N) cycle and produced nitrogenous gases of environmental significance. More than half of all chemical N fertilizer produced globally is used in crop production in East, Southeast and South Asia, where rice is central to nutrition. Emissions of nitrous oxide (N₂O), nitric oxide (NO) and ammonia (NH₃) from croplands in this region were estimated by considering background emission and emissions resulting from N added to croplands, including chemical N, animal manure, biologically fixed N and N in crop residues returned to fields. Background emission fluxes of N₂O and NO from croplands were estimated to be 1.22 and 0.57 kg N ha^?1 yr^?1, respectively. Separate fertilizer‐induced emission factors were estimated for upland fields and rice fields. Total N₂O emission from croplands in the study region was estimated to be 1.19 Tg N yr^?1, with 43% contributed by background emissions. The average fertilizer‐induced N₂O emission, however, accounts for only 0.93% of the applied N, which is less than the default IPCC value of 1.25%, because of the low emission factor from paddy fields. Total NO emission was 591 Gg N yr^?1 in the study region, with 40% from background emissions. The average fertilizer‐induced NO emission factor was 0.48%. Total NH₃ emission was estimated to be 11.8 Tg N yr^?1. The use of urea and ammonium bicarbonate and the cultivation of rice led to a high average NH₃ loss rate from chemical N fertilizer in the study region. Emissions were displayed at a 0.5° × 0.5° resolution with the use of a global landuse database.     

Nitrous oxide and methane fluxes from soils of the Orinoco savanna under different land uses

The study investigates the effect of land‐use change on nitrous oxide (N₂O) and methane (CH₄) fluxes from soil, in savanna ecosystems of the Orinoco region (Venezuela). Gas fluxes were measured by closed static chambers, in the wet and dry season, in representative systems of land management of the region: a cultivated pasture, an herbaceous savanna, a tree savanna and a woodland (control site). Higher N₂O emissions were observed in the cultivated pasture and in the herbaceous savanna compared with the tree savanna and the woodland, and differences were mainly related to fine soil particle content and soil volumetric water content measured in the studied sites. Overall N₂O emissions were quite low in all sites (0–1.58 mg N₂O‐N m^?2 day^?1). The cultivated pasture and the woodland savanna were on average weak CH₄ sinks (?0.05±0.07 and ?0.08±0.05 mg CH₄ m^?2 day^?1, respectively), whereas the herbaceous savanna and the tree savanna showed net CH₄ production (0.23±0.05 and 0.19±0.05 mg CH₄ m^?2 day^?1, respectively). Variations of CH₄ fluxes were mainly driven by variation of soil water‐filled pore space (WFPS), and a shift from net CH₄ consumption to net CH₄ production was observed at around 30% WFPS. Overall, the data suggest that conversion of woodland savanna to managed landscape could alter both CH₄ and N₂O fluxes; however, the magnitude of such variation depends on the soil characteristics and on the type of land management before conversion.     

Night and day: short‐term variation in nitrogen chemistry and nitrous oxide emissions from streams

1. Diel variation in metabolism contributes to variation in oxygen (O₂) concentrations in streams. This variation in O₂ and other parameters (e.g. pH) can in turn affect the rates of microbial nitrogen (N) processing, concentrations of nitrogenous solutes and production of the greenhouse gas nitrous oxide (N₂O). We investigated diel variability in emissions of N₂O and the magnitude of short‐term variability in N solutes across 10 streams. 2. Nitrous oxide fluxes varied on average 2.3‐fold over diel cycles. Concentrations would be underestimated by sampling around noon, but N₂O fluxes would not show a consistent bias. Time‐weighted mean daily N₂O flux was strongly related to nitrate concentration (r² = 0.58). Diel patterns in N₂O and dissolved N species were often complex (rather than simple sinusoidal curves), probably reflecting complex underlying processes. 3. Reliance on samples obtained around noon would overestimate daily mean nitrate concentrations by 5% and underestimate ammonium by 32% (average bias across all streams and dates). 4. Dissolved organic N did not show consistent day–night variation. However, the magnitude of diel variability was similar to that observed for dissolved inorganic N. Organic and inorganic N concentrations were often similar. Both appear to be dynamic components of stream N budgets. 5. The Intergovernmental Panel on Climate Change (IPCC) relies upon an emission factor to estimate indirect agricultural N₂O emissions from streams and ground water. The measured emission factor (defined as the ratio of concentrations of N₂O‐N to ‐N) was typically below the recently revised IPCC default figure. Measured values varied on average 1.8‐fold over approximately 24‐h periods and were slightly higher at night than by day. The emission factor was actually highest in streams that were net sinks for N₂O, highlighting a conceptual problem in the current IPCC method. 6. Typical sampling programmes rely on daytime‐only sampling, which might cause bias in results. In our study streams, the bias was generally small. Diel variation in nitrate concentrations was related to mean temperature; variation in ammonium and N₂O concentrations was greatest at low concentrations of nitrite and ammonium.     

Soil-Atmosphere Exchange of N<Subscript>2</Subscript>O and NO in Near-Natural Savanna and Agricultural Land in Burkina Faso (W. Africa)

In a combined field and laboratory study in the southwest of Burkina Faso, we quantified soil-atmosphere N₂O and NO exchange. N₂O emissions were measured during two field campaigns throughout the growing seasons 2005 and 2006 at five different experimental sites, that is, a natural savanna site and four agricultural sites planted with sorghum (n = 2), cotton and peanut. The agricultural fields were not irrigated and not fertilized. Although N₂O exchange mostly fluctuated between −2 and 8 μg N₂O–N m⁻² h⁻¹, peak N₂O emissions of 10–35 μg N₂O–N m⁻² h⁻¹ during the second half of June 2005, and up to 150 μg N₂O–N m⁻² h⁻¹ at the onset of the rainy season 2006, were observed at the native savanna site, whereas the effect of the first rain event on N₂O emissions at the crop sites was low or even not detectable. Additionally, a fertilizer experiment was conducted at a sorghum field that was divided into three plots receiving different amounts of N fertilizer (plot A: 140 kg N ha⁻¹; plot B: 52.5 kg N ha⁻¹; plot C: control). During the first 3 weeks after fertilization, only a minor increase in N₂O emissions at the two fertilized plots was detected. After 24 days, however, N₂O emission rates increased exponentially at plot A up to a mean of 80 μg N₂O–N m⁻² h⁻¹, whereas daily mean values at plot B reached only 19 μg N₂O–N m⁻² h⁻¹, whereas N₂O flux rates at plot C remained unchanged. The calculated annual N₂O emission of the nature reserve site amounted to 0.52 kg N₂O–N ha⁻¹ a⁻¹ in 2005 and to 0.67 kg N₂O–N ha⁻¹ a⁻¹ in 2006, whereas the calculated average annual N₂O release of the crop sites was only 0.19 kg N₂O–N ha⁻¹ a⁻¹ and 0.20 kg N₂O–N ha⁻¹ a⁻¹ in 2005 and 2006, respectively. In a laboratory study, potential N₂O and NO formation under different soil moisture regimes were determined. Single wetting of dry soil to medium soil water content with subsequent drying caused the highest increase in N₂O and NO emissions with maximum fluxes occurring 1 day after wetting. The stimulating effect lasted for 3–4 days. A weaker stimulation of N₂O and NO fluxes was detected during daily wetting of soil to medium water content, whereas no significant stimulating effect of single or daily wetting to high soil water content (>67% WHC_max) was observed. This study demonstrates that the impact of land-use change in West African savanna on N trace gas emissions is smaller—with the caveat that there could have been potentially higher N₂O and NO emissions during the initial conversion—than the effect of timing and distribution of rainfall and of the likely increase in nitrogen fertilization in the future.     

Snow depth, soil freezing, and fluxes of carbon dioxide, nitrous oxide and methane in a northern hardwood forest

Soil–atmosphere fluxes of trace gases (especially nitrous oxide (N₂O)) can be significant during winter and at snowmelt. We investigated the effects of decreases in snow cover on soil freezing and trace gas fluxes at the Hubbard Brook Experimental Forest, a northern hardwood forest in New Hampshire, USA. We manipulated snow depth by shoveling to induce soil freezing, and measured fluxes of N₂O, methane (CH₄) and carbon dioxide (CO₂) in field chambers monthly (bi-weekly at snowmelt) in stands dominated by sugar maple or yellow birch. The snow manipulation and measurements were carried out in two winters (1997/1998 and 1998/1999) and measurements continued through 2000. Fluxes of CO₂ and CH₄ showed a strong seasonal pattern, with low rates in winter, but N₂O fluxes did not show strong seasonal variation. The snow manipulation induced soil freezing, increased N₂O flux and decreased CH₄ uptake in both treatment years, especially during winter. Annual N₂O fluxes in sugar maple treatment plots were 207 and 99 mg N m⁻² yr⁻¹ in 1998 and 1999 vs. 105 and 42 in reference plots. Tree species had no effect on N₂O or CO₂ fluxes, but CH₄ uptake was higher in plots dominated by yellow birch than in plots dominated by sugar maple. Our results suggest that winter fluxes of N₂O are important and that winter climate change that decreases snow cover will increase soil:atmosphere N₂O fluxes from northern hardwood forests.     

Substantial labile carbon stocks and microbial activity in deeply weathered soils below a tropical wet forest

Contrary to large areas in Amazonia of tropical moist forests with a pronounced dry season, tropical wet forests in Costa Rica do not depend on deep roots to maintain an evergreen forest canopy through the year. At our Costa Rican tropical wet forest sites, we found a large carbon stock in the subsoil of deeply weathered Oxisols, even though only 0.04–0.2% of the measured root biomass (>2 mm diameter) to 3 m depth was below 2 m. In addition, we demonstrate that 20% or more of this deep soil carbon (depending on soil type) can be mobilized after forest clearing for pasture establishment. Microbial activity between 0.3 and 3 m depth contributed about 50% to the microbial activity in these soils, confirming the importance of the subsoil in C cycling. Depending on soil type, forest clearing for pasture establishment led from no change to a slight addition of carbon in the topsoil (0–0.3 m depth). However, this effect was countered by a substantial loss of C stocks in the subsoil (1–3 m depth). Our results show that large stocks of relatively labile carbon are not limited to areas with a prolonged dry season, but can also be found in deeply weathered soils below tropical wet forests. Forest clearing in such areas may produce unexpectedly high C losses from the subsoil.     

Annual wood production in a tropical rain forest in NE Costa Rica linked to climatic variation but not to increasing CO2

Increased atmospheric [CO₂] could theoretically lead to increased forest productivity (‘CO₂ fertilization’). This mechanism was hypothesized as a possible explanation for biomass increases reported from tropical forests in the last 30+ years. We used unique long‐term records of annually measured stands (eighteen 0.5 ha plots, 10 years) and focal tree species (six species, 24 years) to assess the effects of rainfall, temperature, and atmospheric [CO₂] on annual wood production in a neotropical rain forest. Our study area was a meso‐scale section (600 ha) of old‐growth Tropical Wet Forest in NE Costa Rica. Using the repeated remeasurements we directly assessed the relative effects of interannual climatic variation and increasing atmospheric [CO₂] on wood production. A remarkably simple two‐factor model explained 91% of the interannual variance in stand‐level tree growth; the statistically independent factors were total dry season rainfall (positive effect, r²=0.85) and night‐time temperature (negative effect, r²=0.42). Stand‐level tree mortality increased significantly with night‐time temperature. After accounting for dry season rainfall and night‐time temperature, there was no effect of annual [CO₂] on tree growth in either the stand or focal species data. Tree growth in this Tropical Wet Forest was surprisingly sensitive to the current range of dry season conditions and to variations in mean annual night‐time temperature of 1–2°. Our results suggest that wood production in the lowland rainforests of NE Costa Rica (and by extension in other tropical regions) may be severely reduced in future climates that are only slightly drier and/or warmer.     

Temperate forest responses to carbon dioxide, temperature and nitrogen: a model analysis

The ITE Edinburgh Forest Model, which describes diurnal and seasonal changes in the pools and fluxes of C, N and water in a fully coupled forest–soil system, was parametrized to simulate a managed conifer plantation in upland Britain. The model was used to examine (i) the transient effects on forest growth of an IS92a scenario of increasing [CO₂] and temperature over two future rotations, and (ii) the equilibrium (sustainable) effects of all combinations of increases in [CO₂] from 350 to 550 and 750 μmol mol^?1, mean annual temperature from 7.5 to 8.5 and 9.5°C and annual inputs of 20 or 40 kg N ha^?1. Changes in underlying processes represented in the model were then used to explain the responses. Eight conclusions were supported by the model for this forest type and climate.

• 1 Increasing temperatures above 3°C alone may cause forest decline owing to water stress.
• 2 Elevated [CO₂] can protect trees from water stress that they may otherwise suffer in response to increased temperature.
• 3 In N-limiting conditions, elevated [CO₂] can increase allocation to roots with little increase in leaf area, whereas in N-rich conditions elevated [CO₂] may not increase allocation to roots and generally increases leaf area.
• 4 Elevated [CO₂] can decrease water use by forests in N-limited conditions and increase water use in N-rich conditions.
• 5 Elevated [CO₂] can increase forest productivity even in N-limiting conditions owing to increased N acquisition and use efficiency.
• 6 Rising temperatures (along with rising [CO₂]) may increase or decrease forest productivity depending on the supply of N and changes in water stress.
• 7 Gaseous losses of N from the soil can increase or decrease in response to elevated [CO₂] and temperature.
• 8 Projected increases in [CO₂] and temperature (IS92a) are likely to increase net ecosystem productivity and hence C sequestration in temperate forests.

     

Soil nitrogen cycling and nitrous oxide flux in a Rocky Mountain Douglas-fir forest: effects of fertilization,irrigation and carbon addition

Nitrous oxide fluxes and soil nitrogen transformations were measured in experimentally-treated high elevation Douglas-fir forests in northwestern New Mexico, USA. On an annual basis, forests that were fertilized with 200 kg N/ha emitted an average of 0.66 kg/ha of N₂O-N, with highest fluxes occurring in July and August when soils were both warm and wet. Control, irrigated, and woodchip treated plots were not different from each other, and annual average fluxes ranged from 0.03 to 0.23 kg/ha. Annual net nitrogen mineralization and nitrate production were estimated in soil and forest floor usingin situ incubations; fertilized soil mineralized 277 kg ha⁻¹ y⁻¹ in contrast to 18 kg ha⁻¹ y⁻¹ in control plots. Relative recovery of¹⁵NH₄-N applied to soil in laboratory incubations was principally in the form of NO³-N in the fertilized soils, while recovery was mostly in microbial biomass-N in the other treatments. Fertilization apparently added nitrogen that exceeded the heterotrophic microbial demand, resulting in higher rates of nitrate production and higher nitrous oxide fluxes. Despite the elevated nitrous oxide emission resulting from fertilization, we estimate that global inputs of nitrogen into forests are not currently contributing significantly to the increasing concentrations of nitrous oxide in the atmosphere.     

Implications of pre- and post-fertilizing changes in growth and nitrogen pools following multiple applications of nitrogen fertilizer to a Pinus radiata stand over 12 years

Around the world large tracts of forest, previously available for production, have been reserved for nature conservation. This means that wood supply must be met from a reduced land base, including land of low productivity. In addition there are likely to be increasing demands on the use of managed forests for sequestering C as one means of reducing the build up of atmospheric CO2. One way for the forest industry to meet the demands of increased production would be through the use of fertilizers. Substantially increased growth from fertilizer N application has been measured in many cases while, in the northern hemisphere, atmospheric N deposition has been associated with increased growth in some forests.The possibility of using fertilizer N to increase growth, and the effect on the forest and soil, was studied in a research trial area in north-east Tasmania, Australia. Nitrogen was applied for 12 years to a 16 year old P. radiata stand in a low rainfall zone. Growth and foliar nutrient concentrations were measured to age 34 years, to determine change after attainment of steady state growth at age 25 years, and following cessation of fertilizing at age 29 years. Biomass sampling was carried out at ages 25 years, 29 years, and 34 years.Growth at the steady peak rate achieved in the fertilizer plots, of 31 m3 ha-1 periodic annual increment, was accompanied by changes in the N nutrient pools. Surface applied N built up in the surface litter layer while this layer was increasing, from 15 t ha-1 to nearly 50 t ha-1, and fertilizer was being applied between ages 25 and 29 years. Decline of total N in the soil, between ages 25 and 29 years, indicated continued uptake from that source even though fertilizer was being applied. Subsequent increases in soil N, through leaching from the litter layer, followed cessation of fertilizing.Following cessation of annual applications of N fertilizer, growth rates declined by less than 15% and remained substantially (>150%) above the control growth rates for 4 years. Over a 3 year period foliar N concentrations declined steadily, to the same as control plots, although mass of needles was not reduced 5 years after cessation of fertilizer applications and remained significantly above that of the control plots.Without fertilizers, N for new foliage must be supplied from internal recycling and from mineralization of litter and soil organic matter. There were net losses of N from the foliage and the wood, estimated at around 12 kg N ha-1 yr-1, for the fertilized trees for the 5 years following cessation of fertilizing. This indicated retention and recycling within the tree of a considerable proportion of the 104 kg N ha-1 in the foliage, at age 29 years. As rates of mineralization of soil N were likely to be low, this retention of N within the tree was important in maintaining growth rates.In the forest ecosystems investigated, with low initial content of soil organic matter and N, fertilizer N produced improved tree health and substantially increased growth, thus providing the opportunity to manage this forest for increased wood production or C sequestration. Although there was a substantial build up of N and C in the litter on the fertilized treatment, incorporation of N and C into the mineral soil was slow. Over 12 years 1.34 t N ha-1 had been applied, comparable to 40 years of atmospheric deposition at 30 kg N ha-1. There were no signs of detrimental effects from this application.     

Climate- and crop-responsive emission factors significantly alter estimates of current and future nitrous oxide emissions from fertilizer use

The current Intergovernmental Panel on Climate Change (IPCC) default methodology (tier 1) for calculating nitrous oxide (N₂O) emissions from nitrogen applied to agricultural soils takes no account of either crop type or climatic conditions. As a result, the methodology omits factors that are crucial in determining current emissions, and has no mechanism to assess the potential impact of future climate and land‐use change. Scotland is used as a case study to illustrate the development of a new methodology, which retains the simple structure of the IPCC tier 1 methodology, but incorporates crop‐ and climate‐dependent emission factors (EFs). It also includes a factor to account for the effect of soil compaction because of trampling by grazing animals. These factors are based on recent field studies in Scotland and elsewhere in the UK. Under current conditions, the new methodology produces significantly higher estimates of annual N₂O emissions than the IPCC default methodology, almost entirely because of the increased contribution of grazed pasture. Total emissions from applied fertilizer and N deposited by grazing animals are estimated at 10 662 t N₂O‐N yr^?1 using the newly derived EFs, as opposed to 6 796 t N₂O‐N yr^?1 using the IPCC default EFs. On a spatial basis, emission levels are closer to those calculated using field observations and detailed soil modelling than to estimates made using the IPCC default methodology. This can be illustrated by parts of the western Ayrshire basin, which have previously been calculated to emit 8–9 kg N₂O‐N ha^?1 yr^?1 and are estimated here as 6.25–8.75 kg N₂O‐N ha^?1 yr^?1, while the IPCC default methodology gives a maximum emission level of only 3.75 kg N₂O‐N ha^?1 yr^?1 for the whole area. The new methodology is also applied in conjunction with scenarios for future climate‐ and land‐use patterns, to assess how these emissions may change in the future. The results suggest that by 2080, Scottish N₂O emissions may increase by up to 14%, depending on the climate scenario, if fertilizer and land management practices remain unchanged. Reductions in agricultural land use, however, have the potential to mitigate these increases and, depending on the replacement land use, may even reduce emissions to below current levels.     

Effects of an experimental drought on soil emissions of carbon dioxide, methane, nitrous oxide, and nitric oxide in a moist tropical forest

Changes in precipitation in the Amazon Basin resulting from regional deforestation, global warming, and El Niño events may affect emissions of carbon dioxide (CO₂), methane (CH₄), nitrous oxide (N₂O), and nitric oxide (NO) from soils. Changes in soil emissions of radiatively important gases could have feedback implications for regional and global climates. Here we report results of a large‐scale (1 ha) throughfall exclusion experiment conducted in a mature evergreen forest near Santarém, Brazil. The exclusion manipulation lowered annual N₂O emissions by >40% and increased rates of consumption of atmospheric CH₄ by a factor of >4. No treatment effect has yet been detected for NO and CO₂ fluxes. The responses of these microbial processes after three rainy seasons of the exclusion treatment are characteristic of a direct effect of soil aeration on denitrification, methanogenesis, and methanotrophy. An anticipated second phase response, in which drought‐induced plant mortality is followed by increased mineralization of C and N substrates from dead fine roots and by increased foraging of termites on dead coarse roots, has not yet been detected. Analyses of depth profiles of N₂O and CO₂ concentrations with a diffusivity model revealed that the top 25 cm soil is the site of most of the wet season production of N₂O, whereas significant CO₂ production occurs down to 100 cm in both seasons, and small production of CO₂ occurs to at least 1100 cm depth. The diffusivity‐based estimates of CO₂ production as a function of depth were strongly correlated with fine root biomass, indicating that trends in belowground C allocation may be inferred from monitoring and modeling profiles of H₂O and CO₂.     

Nitrogen Oxide Fluxes and Nitrogen Cycling during Postagricultural Succession and Forest Fertilization in the Humid Tropics

The effects of changes in tropical land use on soil emissions of nitrous oxide (N₂O) and nitric oxide (NO) are not well understood. We examined emissions of N₂O and NO and their relationships to land use and forest composition, litterfall, soil nitrogen (N) pools and turnover, soil moisture, and patterns of carbon (C) cycling in a lower montane, subtropical wet region of Puerto Rico. Fluxes of N₂O and NO were measured monthly for over 1 year in old (more than 60 years old) pastures, early- and mid-successional forests previously in pasture, and late-successional forests not known to have been in pasture within the tabonuco (Dacryodes excelsa) forest zone. Additional, though less frequent, measures were also made in an experimentally fertilized tabonuco forest. N₂O fluxes exceeded NO fluxes at all sites, reflecting the consistently wet environment. The fertilized forest had the highest N oxide emissions (22.0 kg N · ha⁻¹· y⁻¹). Among the unfertilized sites, the expected pattern of increasing emissions with stand age did not occur in all cases. The mid-successional forest most dominated by leguminous trees had the highest emissions (9.0 kg N · ha⁻¹· y⁻¹), whereas the mid-successional forest lacking legumes had the lowest emissions (0.09 kg N · ha⁻¹· y⁻¹). N oxide fluxes from late-successional forests were higher than fluxes from pastures. Annual N oxide fluxes correlated positively to leaf litter N, net nitrification, potential nitrification, soil nitrate, and net N mineralization and negatively to leaf litter C:N ratio. Soil ammonium was not related to N oxide emissions. Forests with lower fluxes of N oxides had higher rates of C mineralization than sites with higher N oxide emissions. We conclude that (a) N oxide fluxes were substantial where the availability of inorganic N exceeded the requirements of competing biota; (b) species composition resulting from historical land use or varying successional dynamics played an important role in determining N availability; and (c) the established ecosystem models that predict N oxide loss from positive relationships with soil ammonium may need to be modified. Received 22 February 2000; accepted 6 September 2000.     

Laboratory incubations reveal potential responses of soil nitrogen cycling to changes in soil C and N availability in Mojave Desert soils exposed to elevated atmospheric CO2

Elevated atmospheric carbon dioxide (CO₂) has the potential to alter soil carbon (C) and nitrogen (N) cycling in arid ecosystems through changes in net primary productivity. However, an associated feedback exists because any sustained increases in plant productivity will depend upon the continued availability of soil N. We took soils from under the canopies of major shrubs, grasses, and plant interspaces in a Mojave Desert ecosystem exposed to elevated atmospheric CO₂ and incubated them in the laboratory with amendments of labile C and N to determine if elevated CO₂ altered the mechanistic controls of soil C and N on microbial N cycling. Net ammonification increased under shrubs exposed to elevated CO₂, while net nitrification decreased. Elevated CO₂ treatments exhibited greater fluxes of N₂O–N under Lycium spp., but not other microsites. The proportion of microbial/extractable organic N increased under shrubs exposed to elevated CO₂. Heterotrophic N₂‐fixation and C mineralization increased with C addition, while denitrification enzyme activity and N₂O–N fluxes increased when C and N were added in combination. Laboratory results demonstrated the potential for elevated CO₂ to affect soil N cycling under shrubs and supports the hypothesis that energy limited microbes may increase net inorganic N cycling rates as the amount of soil‐available C increases under elevated CO₂. The effect of CO₂ enrichment on N‐cycling processes is mediated by its effect on the plants, particularly shrubs. The potential for elevated atmospheric CO₂ to lead to accumulation of NH₄⁺ under shrubs and the subsequent volatilization of NH₃ may result in greater losses of N from this system, leading to changes in the form and amount of plant‐available inorganic N. This introduces the potential for a negative feedback mechanism that could act to constrain the degree to which plants can increase productivity in the face of elevated atmospheric CO₂.     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

Growth response of Pinus radiata to multiple applications of nitrogen fertilizer and evaluation of the quantity of added nitrogen remaining in the forest system

Over a period of nine years, 922 kg ha^-1 of N was added in eight applications to a 16 year old P. radiata stand in a low rainfall area in north-east Tasmania, Australia. Fertilizing lifted current annual increment from 8.5 m³ha^-1 for the unfertilized plots, to 31 m³ha^-1. Increased growth was associated with improved health of the trees. Biomass measurements showed that there was a large increase in needle retention as well as needle mass on the fertilized trees. Concentration of N was also higher in fertilized trees. Fertilized plots contained 467 kg ha^-1 more N than unfertilized plots. This represented about 50 percent of the N applied. Most of this extra N was in the forest crop and in the surface 10 cm of the soil. There was no increase in organic carbon in the surface soil with the result that the C/N ratio was reduced from a very high 28 to 17.Despite the high growth rates attained in the N fertilized plots, the failure to increase soil organic matter and the loss from the site of much of the applied N indicated that long term improvement of growth of these low rainfall sites was unlikely. Growth would only be maintained with continued N fertilizer additions.