Accumulation of pollutant nitrogen in calcareous and acidic grasslands: Evidence from N flux and 15N tracer studies

Semi-natural calcareous and acidic grasslands are known to be sensitive to increased atmospheric N deposition. However, the fate of pollutant N within these systems is unknown. This paper reports on the first studies to determine the fate of added N within a calcareous and an acidic grassland subject to long-term simulated enhanced N deposition. Intact soil/turf cores were removed from field plots treated for six years with enhanced N deposition (ambient +0, +35 and +140 kg N ha^–1 year^–1). Cores were inserted into lysimeters and output fluxes of N were monitored in detail. Complete N budgets—calculated from the N flux data—showed considerable accumulation of N within the treated grasslands, up to 76% and 38% of pollutant N in the calcareous and acidic grasslands respectively. In the second study, the short-term (21 day) fate of pollutant N was determined by tracing ¹⁵N labelled ammonium nitrate (+35 kg N ha^–1 year^–1) though the acidic and calcareous lysimeters into plant, soil and leachate pools. Up to 91% and 59% of ¹⁵N was recovered in soils and vegetation of the calcareous and acidic grasslands respectively, with negligible amounts recovered in soil extractable ammonium and nitrate (<0.3%) and in leachate (<0.02%). This rapid short-term immobilisation of pollutant N supports the long-term accumulation of the element calculated from the N flux study.     

Nitrogen Budget of a Spruce Forest Ecosystem After Six-year Addition of Ammonium Sulphate in Southwest Sweden

A nitrogen (N) budget was constructed for a period of 6 years (1988–1993) in a Norway spruce stand with current deposition of 19 kg N and 22 kg S ha⁻¹ year⁻¹. The stand was fertilized annually by addition of 100 kg N and 114 kg S ha⁻¹ (NS). Above and below ground biomass, litterfall, fine- root litter production, soil solution and net mineralization were measured to estimate pools, fluxes and accumulation of nitrogen. The average needle litterfall in control (C) and NS plots in 1993 was 2.2 and 2.5 ton ha⁻¹ year⁻¹, respectively. The fine root litter production prior to treatment (1987) was 4.4 ton ha⁻¹ year⁻¹ and after treatment (1993) it was 4.5 and 3.9 ton ha⁻¹ year⁻¹ in C and NS plots, respectively. Net N mineralization in the soil profile down to 50 cm was estimated to be 86 and 115 kg ha⁻¹ year⁻¹ in C and NS plots, respectively in 1992. During the treatment period the uptake of N in the needle biomass in C and NS plots was 29 and 77 kg ha⁻¹ year⁻¹, respectively. No N was accumulated in needles of C plot where the NS plots accumulated 34 kg ha⁻¹ year⁻¹. Of the annually added inorganic N to NS plots 47% was accumulated in the above and below ground biomass and 37% in the soil. N fluxes via fine-root litter production in the C plots were much higher (54 kg ha⁻¹ year⁻¹) than that via litterfall (29 kg ha⁻¹ year⁻¹). The corresponding values in the NS plots were 65 and 43 kg ha⁻¹ year⁻¹, respectively. Most of the net N mineralization occurred in the FH layer and upper mineral soil. It is concluded that fine root litter and litterfall play an important role in the cycling of N. Despite a high N uptake the losses of N in litterfall and fine root litter resulted in an incorporation of N in soil organic matter.     

An automated wet deposition system to compare the effects of reduced and oxidised N on ombrotrophic bog species: Practical considerations

Critical N loads for ombrotrophic bogs, which often contain rare and N-sensitive plants (especially those in lower plant groups: lichens, mosses and liverworts), are based on very few experimental data from measured, low background N deposition areas. Additionally the relative effects of reduced versus oxidised N are largely unknown. This paper describes an automated field exposure system (30 km S. of Edinburgh, Scotland) for treating ombrotrophic bog vegetation with fine droplets of oxidised N (NaNO₃) and reduced N (NH₄Cl). Whim Moss exists in an area of low ambient N deposition (ca. 8 kg N ha^?1 y^?1), the sources and quantification of which are described. The wet N treatment system is run continuously, and is controlled/activated by wind speed and rainfall to provide a unique simulation of “real worl” treatment patterns (no rain=no treatment). Simulated precipitation is supplied at ionic concentrations below 4 mM in rainwater collected on site. Treatments provide a replicated dose response to 16, 32 and 64 kg N ha^?1 y^?1 adjusted for ambient deposition (8 kg N ha^?1 y^?1). The 16 and 64 kg N ha^?1 y^?1 are duplicated with a P+K supplement. Baseline soil chemistry and foliar nutrient status was established for all 44 plots for Calluna vulgaris, Sphagnum capillifolium, Hypnum jutlandicum and Cladonia portentosa.     

A Comparison of Loch Chemistry from 1955 and 1999 in the Cairngorms,N.E. Scotland

The Cairngorms in north-east Scotland is remote from pollutant sources although it currently receives ca. 10 kg ha¹ yr¹ S and ca. 11 kg ha¹ yr¹ N deposition from the atmosphere.In 1955, 15 lochs (lakes) at a range of altitudes were sampled and analysed for major ion concentrations. A new survey of these and an additional 23 lochs and their catchment soils was conducted in 1999 to determine the impact of acid deposition, and the changes in loch chemistry since the 1955 survey. The bedrock geology of this region has a strong influence on the loch chemistry. Surface waters were generally more acidic in high altitude areas due to predominantly poorly buffered, thin alpine soils developed on granitic parent material (mean acid neutralising capacity (ANC) for 23 lochs = 30 eq L¹). At lower altitudes where the geology is dominated by Dalradian metamorphic rocks surface waters are comparatively base rich and have higher ANC (mean ANC for 15 lochs = 157 eq L¹). Surface water nitrate concentrations show a negative relationship with soil C:N status, in that higher nitrate only occurs at low soil C:N ratios. A comparison of data for 1955 and 1999 shows that sulphate concentrations are significantly lower (67.8 and 47.5 eq L¹, respectively), and pH has improved (pH 5.6 and 5.9) in response to decreased S deposition since the mid 1970s. However, mean nitrate concentrations were found to increase from 2.48 >eq L¹ in 1955 to 5.65 eq L¹ in 1999. Differences in the sampling and laboratory methods from 1955 and 1999 are acknowledged in the interpretation of data.     

Linking field experiments to long-term simulation of impacts of nitrogen deposition on heathlands and moorlands

The results from three long-term field manipulation studies of the impacts of increased nitrogen deposition (0–120 kg N ha^?1 yr^?1) on lowland and upland heathlands in the UK were compared, to test if common responses are observed. Consistent increases in Calluna foliar N content and decreases in litter C:N ratios were found across all sites, while increases in N leaching were not observed at any site over the range 0–80 kg ha^?1 yr^?1. However, the response of Calluna biomass did vary between sites, possibly reflecting site differences in nutrient status and management histories. Five versions of a simulation model of heathland responses to N were developed, each reflecting different assumptions about the fate and turnover of soil N. Model outputs supported the deduction from mass balance calculations at two of the field sites that N additions have resulted in an increase in immobilisation; the latter was needed to prevent the model overestimating measured N leaching. However, this version of the model significantly underestimated Calluna biomass. Model versions, which included uptake of organic N by Callunaand re-mobilisation of N from the soil organic store provided some improvement in the fit between modelled and field biomass data, but re-mobilisation also led to an overestimation of N leaching. Quantification of these processes and their response to increased N deposition are therefore critical to interpreting experimental data and predicting the long-term impacts of atmospheric deposition on heathlands and moorlands.     

Applying metabolic fingerprinting to ecology: The use of Fourier-transform infrared spectroscopy for the rapid screening of plant responses to N deposition

The potential for metabolic fingerprinting via Fourier-transform infrared (FT-IR) spectroscopy to provide a novel approach for the detection of plant biochemical responses to N deposition is examined. An example of spectral analysis using shoot samples taken from an open top chamber (OTC) experiment simulating wet ammonium deposition is given. Sample preparation involved oven drying and homogenisation via mill grinding. Slurries of a consistent dilution were then prepared prior to FT-IR analysis. Spectra from control, 8 and 16 kg N ha^?1 yr^?1 treatments were then subjected to cross-validated discriminant function analysis. Ordination diagrams showed clear separation between the three N treatments examined. The potential for using Calluna vulgaris (L.) Hull as a bioindicator of N deposition is further evident from these results. The results also clearly demonstrate the power of FT-IR in discriminating between subtle phenotypic alterations in overall plant biochemistry as affected by ammonium pollution.     

Fate of mutagenic chemicals in soil amended with petroleum and wood preserving sludges

A field study using monoliths (lysimeters) of a sandy clay loam soil was conducted to assess the fate of mutagenic chemicals after refinery and wood preserving bottom sediment sludges were land treated. The Ames Salmonella/microsome assay¹ was used to determine the direct (without metabolic activation, −S9) and indirect (with metabolic activation, + S9) mutagenicity of the wastes, unamended soil, waste amended soils, and leachate. Extracts having a mutagenic ratio (MR) (MR= No. colonies from sample extract/No. colonies from DMSO solvent control) of ⩾ 2 were considered positively mutagenic. Extracts of the wood preserving waste sludge without activation were non-mutagenic (MR < 2) but extracts with activation ( + S9) produced very strong indirect mutagenicity (MR = 7.9). After soil incorporation, the waste amended soil produced very strong direct (MR = 8.9) and indirect (MR = 11.9) mutagenicity by day 180 and remained mutagenic (MR = 5.7, −S9; MR = 3.95, + S9) through day 350. The amount of residue in leachate from the wood preserving waste amended lysimeters was significantly greater (P <0.05) than the unamended soil during the first 90 days after waste application, but was not different after 90 days. The leachate residue from wood preserving waste amended lysimeters in the 90–180-day period produced mutagenic responses both with (MR = 2.24 and 2.51) and without (MR = 2.29) activation. Polynuclear aromatic hydrocarbons were the main constituents identified in the leachate residues that produced a mutagenic response. Soil treatment of the refinery sludge reduced its weak indirect mutagenicity before soil incorporation (MR = 2) to non-mutagenic (MR = 1.4) immediately following soil treatment. The MR of the waste amended soil increased to 1.7 by day 180 but by day 350 decreased to a level equal to that observed at day 0 (MR = 1.4). Leachate from the refinery amended lysimeters had significantly greater (P < 0.05) amounts of organic residue than unamended lysimeters 180 to 350 days after waste application. The leachate from one refinery waste amended lysimeter (90–180 days after waste application) produced a mutagenic response (MR = 3.16). The refinery sludge was detoxified shortly after soil treatment, but the wood preserving sludge required > 350 days to detoxify in the soil environment. The possibility exists that mobile mutagenic chemicals may leach into underlying groundwater from the treatment zone of soils amended with refinery and wood preserving sludges.     

Microbial nitrogen transformation potential in surface run-off leachate from a tropical landfill

Ammonium is one of the major toxic compounds and a critical long-term pollutant in landfill leachate. Leachate from the Jatibarang landfill in Semarang, Indonesia, contains ammonium in concentrations ranging from 376 to 929 mg N L⁻¹. The objective of this study was to determine seasonal variation in the potential for organic nitrogen ammonification, aerobic nitrification, anaerobic nitrate reduction and anaerobic ammonium oxidation (anammox) at this landfilling site. Seasonal samples from leachate collection treatment ponds were used as an inoculum to feed synthetic media to determine potential rates of nitrogen transformations. Aerobic ammonium oxidation potential (<0.06 mg N L⁻¹ h⁻¹) was more than a hundred times lower than the anaerobic nitrogen transformation processes and organic nitrogen ammonification, which were of the same order of magnitude. Anaerobic nitrate oxidation did not proceed beyond nitrite; isolates grown with nitrate as electron acceptor did not degrade nitrite further. Effects of season were only observed for aerobic nitrification and anammox, and were relatively minor: rates were up to three times higher in the dry season. To completely remove the excess ammonium from the leachate, we propose a two-stage treatment system to be implemented. Aeration in the first leachate pond would strongly contribute to aerobic ammonium oxidation to nitrate by providing the currently missing oxygen in the anaerobic leachate and allowing for the growth of ammonium oxidisers. In the second pond the remaining ammonium and produced nitrate can be converted by a combination of nitrate reduction to nitrite and anammox. Such optimization of microbial nitrogen transformations can contribute to alleviating the ammonium discharge to surface water draining the landfill.     

The Fernow Watershed Acidification Study: Ecosystem Acidification, Nitrogen Saturation and Base Cation Leaching

In 1989, a watershed acidification experiment was begun on the Fernow Experimental Forest in West Virginia, USA. Ammonium sulfate fertilizer (35.5 kg N ha⁻¹ yr⁻¹and 40.5 kg S ha⁻¹ yr⁻¹) was applied to a forested watershed (WS3) that supported a 20-year-old stand of eastern deciduous hardwoods. Additions of N and S are approximately twice the ambient deposition of nitrogen and sulfur in the adjacent mature forested watershed (WS4), that serves as the reference watershed for this study. Acidification of stream water and soil solution was documented, although the response was delayed, and acidification processes appeared to be driven by nitrate rather than sulfate. As a result of the acidification treatment, nitrate solution concentrations increased below all soil layers, whereas sulfate was retained by all soil layers after only a few years of the fertilization treatments, perhaps due to adsorption induced from decreasing sulfate deposition. Based on soil solution monitoring, depletion of calcium and magnesium was observed, first from the upper soil horizons and later from the lower soil horizons. Increased base cation concentrations in stream water also were documented and linked closely with high solution levels of nitrate. Significant changes in soil chemical properties were not detected after 12 years of treatment, however.     

A chronology of nitrogen deposition in the UK between 1900 and 2000

Measurements of the concentrations of nitrogen compounds in air and precipitation in the UK have been made since the mid-19th century, but no networks operating to common protocols and having traceable analytical procedures were established until the 1950s. From 1986 onwards, a high-quality network of sampling stations for precipitation chemistry was established across the UK. In the following decade, monitoring networks provided measurement of NO₂, NH₃, HNO₃ and a satisfactory understanding of the dry deposition process for these gases allowed dry deposition to be quantified. Maps of N deposition for oxidized and reduced compounds at a spatial scale of 5 km × 5 km are available from 1986 to 2000. Between 1950 and 1985, the more limited measurements, beginning with those of the European Air Chemistry Network (EACN) provide a reasonable basis to estimate wet deposition of NO^? ₃?N and NH⁺ ₄?N. For the first half of the century, estimates of deposition were scaled with emissions assuming a constant relationship between emission and deposition for each of the components of the wet and dry deposition budget at the country scale. Emissions of oxidized N, which dominated total nitrogen emissions throughout the century, increased from 312 kt N annually in 1900 to a peak of 787 kt for the decade 1980–1990 and then declined to 460 kt in 2000. Emissions of reduced N, largely from coal combustion were about 168 kt N in 1900, increasing to a peak of 263 kt N in 2000 and by now dominated by agricultural sources. Reduced N dominated the deposition budget at the country scale, increasing from 163 kt N in 1900 to 211 kt N in 2000, while deposition of oxidized N was 66 kt N in 1900 and 191 kt N in 2000. Over the century, 68 Mt (Tg) of fixed N was emitted within the UK, 78% as NO _x , while 29 Mt of nitrogen was deposited (43% of emissions), equivalent to 1.2 t N ha^?1, on average, with 60% in the reduced form. Deposition to semi-natural ecosystems is approximately 15 Tg N, equivalent to between 1 and 5 t N ha^?1, over the century and appears to be accumulating in soil. The N deposition over the century is similar in magnitude to the total soil N inventory in surface horizons.     

Nitrate Leaching From a Mountain Forest Ecosystem with Gleysols Subjected to Experimentally Increased N Deposition

Nitrate leaching was measured over seven years of nitrogen (N) addition in a paired-catchment experiment in Alptal, central Switzerland (altitude: 1200 m, bulk N deposition: 12 kg ha^-1 a^-1). Two forested catchments (1500 m² each) dominated by Picea abies) were delimited by trenches in the Gleysols. NH₄NO₃ was added to one of the catchments using sprinklers. During the first year, the N addition was labelled with ¹⁵N. Additionally, soil N transformationswere studied in replicated plots. Pre-treatment NO₃ ^--N leaching was 4 kg ha^-1 a^-1 from both catchments, and remained between 2.5 and 4.8 kg ha^-1 a^-1 in the control catchment. The first year of treatment induced an additional leaching of 3.1 kg ha^-1, almost 90% of which was labelled with ¹⁵N, indicating that it did not cycle through the large N pools of the ecosystem (soil organic matter and plants). These losses partly correspond to NO₃ ^- from precipitation bypassing the soil due to preferential flow. During rain or snowmelt events, NO₃ ^- concentration peaks as the water table is rising, indicating flushing from the soil. Nitrification occurs temporarily along the water flow paths in the soil and can be the source of NO₃ ^- flushing. Its isotopic signature however, shows that this release mainly affects recently applied N, stored only between runoff events or up to a few weeks. At first, the ecosystem retained 90% of the added N (2/3 in the soil), but NO₃ ^- losses increased from 10 to 30% within 7 yr, indicating that the ecosystem became progressively N saturated.     

Controls on leaching of N species in upland moorland catchments

Spatial and temporal changes in mobility of N species have been studied for three UK upland river networks, the Etherow in the South Pennines, the Nether Beck in the Lake District and the Dee in NE Scotland. The catchments are subject to N deposition at 35.1, 22.0 and 10.8–15.6 kg N ha^?1 yr^?1, respectively. The NH⁺ ₄ leaching appears to be predominantly regulated by flow path in more polluted upland catchments. It is greatest where water draining acidified peaty soils contributes more to total discharge. Soluble organic matter may provide the dominant counter anion. In the Etherow and Dee catchments, which are dominated by acid mineral and organic soils, at high discharge NO^? ₃ also appears to be associated with greater input of water from acidified soils. In contrast, for the Nether Beck, higher NO^? ₃ concentrations are associated with tributaries draining soils contributing water with higher alkalinity, suggesting nitrification is important. For the Etherow and Dee, dissolved organic N (DON) appears to originate predominantly from acidified, peaty soils. Spiking experiments with peat soil from the Etherow catchment confirmed the limited capacity of these soils to utilize inorganic N inputs, favouring equilibration with NH⁺ ₄ inputs and leaching losses of inorganic N throughout the year.     

Ammonia volatilization,N2O and CO2 emissions from landfill leachate-irrigated soils

Effects of leachate addition on ammonia volatilization and N₂O and CO₂ emissions from two different soils were investigated using the 10-day laboratory incubation method at two levels of moisture content. Ammonia volatilization was dominated by soil pH and only occurred in alkaline clay soil, where 0.26–0.32% of soil ammonia could be lost. The N₂O emission from the alkaline clay soil was one order of magnitude greater than that from the acidic sandy soil, when either water or leachate was irrigated. Increasing the moisture content from 46% water-filled pore space (WFPS) to 70% WFPS in the alkaline clay soil or the acidic sandy soil by either water or leachate irrigation increased the N₂O emission by over twofold. The CO₂ emission from each soil sample at the two WFPSs was almost the same. The CO₂ emission from the alkaline clay soil with leachate addition was 72% lower than that from the acidic sandy soil with leachate addition, and 6.7 times higher than that from the alkaline clay soil with distilled water addition. Ammonia volatilization and N₂O emission under leachate irrigation could be minimized by avoiding the excessively wet condition and by selecting the acidic sandy soil with low organic carbon and total nitrogen content.     

An Automated Wet Deposition System to Compare the Effects of Reduced and Oxidised N on Ombrotrophic Bog Species: Practical Considerations

Critical N loads for ombrotrophic bogs, which often contain rare and N-sensitive plants (especially those in lower plant groups: lichens, mosses and liverworts), are based on very few experimental data from measured, low background N deposition areas. Additionally the relative effects of reduced versus oxidised N are largely unknown. This paper describes an automated field exposure system (30 km S. of Edinburgh, Scotland) for treating ombrotrophic bog vegetation with fine droplets of oxidised N (NaNO₃) and reduced N (NH₄Cl). Whim Moss exists in an area of low ambient N deposition (ca. 8 kg N ha^–1 y^–1), the sources and quantification of which are described. The wet N treatment system is run continuously, and is controlled/activated by wind speed and rainfall to provide a unique simulation of real world treatment patterns (no rain=no treatment). Simulated precipitation is supplied at ionic concentrations below4mMin rainwater collected on site. Treatments provide a replicated dose response to 16, 32 and 64 kg N ha^–1 y^–1 adjusted for ambient deposition (8 kg N ha^–1 y^–1). The 16 and 64 kg N ha^–1 y^–1 are duplicated with a P+K supplement. Baseline soil chemistry and foliar nutrient status was established for all 44 plots for Calluna vulgaris, Sphagnum capillifolium, Hypnum jutlandicum and Cladonia portentosa.     

Fate and distribution of nitrogen in soil and plants irrigated with landfill leachate

Landfill leachate contains a high concentration of ammoniacal substances which can be a potential supply of N for plants. A bioassay was conducted using seeds of Brassica chinensis and Lolium perenne to evaluate the phytotoxicity of the leachate sample. A soil column experiment was then carried out in a greenhouse to study the effect of leachate on plant growth. Two grasses (Paspalum notatum and Vetiver zizanioides) and two trees (Hibiscus tiliaceus and Litsea glutinosa) were irrigated with leachate at the EC50 levels for 12 weeks. Their growth performance and the distribution of N were examined and compared with columns applied with chemical fertilizer. With the exception of P. notatum, plants receiving leachate and fertilizer grew better than those receiving water alone. The growth of L. glutinosa and V. zizanioides with leachate irrigation did not differ significantly from plants treated with fertilizer. Leachate irrigation significantly increased the levels of NH_x-N in soil. Although NO_x-N was below 1 mg N L⁻¹ in the leachate sample, the soil NO_x-N content increased by 9-fold after leachate irrigation, possibly as a result of nitrification. Leachate irrigation at EC50 provided an N input of 1920 kg N ha⁻¹ over the experimental period, during which up to 1050 kg N ha⁻¹ was retained in the soil and biomass, depending on the type of vegetation. The amount of nutrient added seems to exceed beyond the assimilative capability. Practitioners should be aware of the possible consequence of N saturation when deciding the application rate if leachate irrigation is aimed for water reuse.     

Applying metabolic fingerprinting to ecology: The use of Fourier-transform infrared spectroscopy for the rapid screening of plant responses to N deposition

The potential for metabolic fingerprinting via Fourier-transform infrared (FT-IR) spectroscopy to provide a novel approach for the detection of plant biochemical responses to N deposition is examined. An example of spectral analysis using shoot samples taken from an open top chamber (OTC) experiment simulating wet ammonium deposition is given. Sample preparation involved oven drying and homogenisation via mill grinding. Slurries of a consistent dilution were then prepared prior to FT-IR analysis. Spectra from control, 8 and 16 kg N ha^–1 yr^–1 treatments were then subjected to cross-validated discriminant function analysis. Ordination diagrams showed clear separation between the three N treatments examined. The potential for using Calluna vulgaris (L.) Hull as a bioindicator of N deposition is further evident from these results. The results also clearly demonstrate the power of FT-IR in discriminating between subtle phenotypic alterations in overall plant biochemistry as affected by ammonium pollution.     

Stem Growth of Picea Abies in South Western Sweden in the 10 Years Following Liming and Addition of PK and N

Liming and/or application of specific nutrients have been proposed as countermeasures to the acidification of forest soils in southern Sweden. In this study the stem growth of Picea abies (L.) Karst. growing on acidic mineral soils in SW Sweden was investigated 10 years after additions of lime (Ca; 3000 kg lime ha⁻¹), lime plus P (25 kg ha⁻¹) and K (80 kg ha⁻¹), or N in low doses (2 × 10 kg ha⁻¹ yr⁻¹) (treatments: CaPK, Ca, N, CaPKN, and 2Ca2P2K, respectively). Compared with the control, stem growth was increased following all treatments involving lime additions, including liming alone. The PK addition did not seem to affect growth. The most plausible cause of the observed growth increases was that the lime additions indirectly increased the supply of plant-available N. The annual low-dose N addition did not significantly affect growth. This suggests that air-borne deposition of N, which supplies very small doses of N throughout the year, has a minor or even negligible influence on P. abies growth.     

Early effects of atmospheric ammonia deposition on Calluna vulgaris (L.) hull growing on an ombrotrophic peat bog

This paper reports data from a field study investigating the impacts of elevated ammonia (NH₃) deposition on Calluna vulgaris growing on an ombrotrophic peat bog in S.E. Scotland. Shoot extension, foliar N concentrations, chlorophyll concentration and chlorophyll fluorescence were measured during the second growing season of exposure to a gradient of ammonia concentrations. Results indicate that NH₃ increases growth between 150–200 kg N ha^?1y^?1 cumulative deposition. Foliar N content increased significantly in response to NH₃ cumulative deposition up to 400 kg N ha^?1 y^?1 whereas chlorophyll a content significantly decreased. Measurements of Fv/Fm suggest that although NH₃ exposure altered the growth and reduced chlorophyll a, the efficiency of photosystem II was insensitive to NH₃–N deposition at this stage.     

Forest Ecosystem Responses to Atmospheric Pollution: Linking Comparative With Experimental Studies

The impact on an ecosystem of an environmental stress, such as climate change or air pollution, can be studied through experimentation, through comparisons of sites across a gradient of the stress, through long-term studies at a single site, or through theoretical or modelling approaches. Although the former three techniques often are used to develop and test models, it is much rarer to explicitly link experimental, comparative or long-term studies together. Here we present a concept for combining experimental and comparative research to assess the direction and rate of change, the expected long-term state, and the rate at which the long-term state is achieved after an ecosystem is exposed to an environmental stress. We do this by comparing the response of a forest in Denmark to experimentally increased N deposition with the expected long-term response based on a European database of forests exposed to different levels of N deposition over long time periods. The analysis suggests that if N deposition were to increase by 3-fold to about 50 kg N ha^-1 a^-1 at the Danish site, and remain at this level, the N concentration in needles would respond within 2–4 yr after the onset of the enhanced N deposition, and would rapidly plateau to an expected mean value of 18.0 mg N g^-1 dry mass (95% confidence interval ± 2.5 mg g^-1). The N concentration of new litter also would respond rapidly (1–2 yr) to reach an expected value of 16.6 mg N kg^-1 dry mass (± 3). The N concentration of the organic layer in the soil would increase much more slowly, but a significant increase would be expected within 5–10 yr. Mineral soil pH would take more than 7 yr to change. Finally, the flux of dissolved inorganic N in leachate wouldbegin to increase immediately, but would take many years to reach the expected level of 22.4 kg N ha^-1 a^-1(± 4).     

Fluxes of NO3-, NH4+, NO,NO2, and N2O in an Old Danish Beech Forest

The fluxes of the major nitrogen compounds havebeen investigated in many ecosystem studies over the world.However, only in few studies has attention been drawn to theimportance of the fluxes of minor gaseous nitrogen compoundsto complete the nitrogen cycle. In Denmark a detailed study onthe nitrogen cycle in an old beech forest has been implementedin 1997 at Gyrstinge near Sorø, Zealand. The study includesthe fluxes of the gases NO, N₂O and water mediatedtransport of NO₃ ^- and NH₄ ⁺. Measurementsof the fluxes of the gaseous compounds are performed withmicro-meteorological methods (eddy-correlation and gradient)and with chambers. Water mediated fluxes encompass rain,throughfall, stem-flow and leaching from the root zone. Thehydrological model is verified by TDR measurements. The findings show that the total water mediated N input tothe forest floor with throughfall and stemflow was 25.6 kg Nha^-1 yr ^-1, and open field wet deposition withprecipitation was 19.0 kg N ha^-1 yr ^-1. The internalcycling of N in the ecosystem measured as turnover oflitterfall and plant uptake was 100 kg N ha^-1 yr ^-1and 14 kg N ha^-1 yr ^-1, respectively. The fluxes ofthe gaseous N compounds NO and N₂O were of minorimportance for the total N turnover in the forest, NO_xemission being <1 kg N ha^-1 yr ^-1 and N₂Oemission from the soil being 0.5 kg N ha^-1 yr ^-1 withno significant difference between wet and dry soils.Concentrations of NO₃ ^- and NH₄ ⁺ in thesoil solution beneath the rooting zone are very small andconsequently the N leaching is almost negligible. It isconcluded that the nitrogen mass balance of this old beechforest ecosystem mainly is controlled by the input by dry andwet deposition and a large internal N cycle with a fast litterturnover. The nitrogen input tothe forest ecosystem which currently exceeds the critical loadby 5 kg N ha^-1 yr ^-1is mainly accumulated in the soil and no significant nitrateleaching is occurring.