Deposition of fixed atmospheric nitrogen and foliar nitrogen content of bryophytes and Calluna vulgaris (L.) Hull

Atmospheric deposition of fixed nitrogen as nitrate and ammonium in rain and by dry deposition of nitrogen dioxide, nitric acid and ammonia has increased throughout Europe during the last two decades, from 2-6 kg N ha(-1) year(-1) to 15-60 kg N ha(-1) year(-1). The nitrogen contents of bryophytes and the ericaceous shrub Calluna vulgaris have been measured at a range of sites, with the objective of showing the degree to which nitrogen deposition is reflected in foliar plant nitrogen. Tissue nitrogen concentrations of herbarium bryophyte samples and current samples of the same species collected from the same sites were compared. No significant change in tissue nitrogen was recorded at a remote site in north-west Scotland where nitrogen inputs are small (< 6 kg N ha(-1) year(-1)). Significant increases in tissue N occurred at four sites ranging from 38% in central Scotland to 63% in Cumbria where nitrogen inputs range from 15 to 30 kg N ha(-1) year(-1). The relationships found between the estimated input of atmospheric nitrogen and the tissue nitrogen content of the selected bryophytes and Calluna at the sites investigated were found to be generally linear and fitted the form N(tissue) = 0.62 + 0.022 N(dep) for bryophytes and N(tissue) = 0.83 + 0.045 N(dep) for Calluna. There was thus an increase in total tissue nitrogen of 0.02 mg g(-1) dry weight for bryophytes and 0.045 mg g(-1) dry weight for Calluna for an increase in atmospheric nitrogen deposition of 1 kg ha(-1) year(-1). The lowest concentrations were found in north-west Scotland and the highest in Cumbria and the Breckland heaths of East Anglia, both areas of high atmospheric nitrogen deposition (30-40 kg N ha(-1) year(-1)). The implications of increased tissue nitrogen content in terms of vegetation change are discussed. Changes in atmospheric nitrogen deposition with time were also examined using measured values and values inferred from tissue nitrogen content of mosses. The rate of increase in nitrogen deposition is not linear over the 90-year period, and the increases were negligible over the period 1880-1915. However, during the period 1950 to 1990 the data suggest an increase in nitrogen deposition of 2 kg N ha(-1) every 10 years.     

The influence of nitrogen in stemflow and precipitation on epiphytic bryophytes, Isothecium myosuroides Brid., Dicranum scoparium Hewd. and Thuidium tamariscinum (Hewd.) Schimp of Atlantic oakwoods

The spatial relationship between the concentration and deposition of the major ions in precipitation and stemflow and their influence on the tissue nitrogen concentration of three epiphytic bryophytes on Quercus petraea (Matt) Liebl. and Q. robur L. was investigated at seven UK Atlantic oak woodland sites with a range of total N deposition of 55-250mmolm(-2). The main driver of change in tissue N concentrations of three epiphytic bryophytes (Isothecium myosuroides Brid. (Eurhynchium myosuroides (Brid.) Schp.), Dicranum scoparium Hewd. and Thuidium tamariscinum (Hewd.) Schimp.) was total N deposition in stemflow, dominated by ammonium deposition. The three epiphytic species also showed strong relationships between tissue N concentration and total N deposition in rainfall but a poor correlation with total N ion concentration in rainfall. This study shows that epiphytic bryophytes utilise stemflow N and thus increase their risk from inputs of total N deposition compared to terricolous species at the same site.     

Atmospheric deposition and canopy exchange processes in heathland ecosystems

The aims of the present study were to determine canopy exchange processes and to quantify total atmospheric deposition of sulphur and nitrogen in heathland. The study was carried out in dry inland heath vegetation, dominated by Calluna vulgaris, in two nature reserves in the eastern part of the Netherlands. Atmospheric deposition was determined with throughfall-stemflow measurements, adapted for low vegetation. Throughflow measurements (sum of throughfall and stemflow) in artificial Calluna canopies showed co-deposition of SOx and NHy upon heathland vegetation. In the real Calluna canopy, a significant part of the deposited ammonia/ammonium was directly assimilated by the Calluna shoots, especially in wet periods. The concentrations of potassium, calcium and magnesium in throughflow, after passage through the Calluna canopy, increased significantly compared with bulk precipitation. The amount of cations lost from the canopy were in good agreement with the observed ammonium uptake by the Calluna. A field experiment demonstrated that losses of the above-mentioned cations can be doubled by application of ammonium sulphate. It was shown that interception deposition is an important component of the atmospheric deposition of sulphur and nitrogen upon Calluna heathland; bulk precipitation amounted to only c. 35-40% of total atmospheric input. Total atmospheric deposition of sulphur and nitrogen in the investigated heathlands was 1.5-2.1 (27-33 kg S ha(-1) yr(-1)) and 2.1-3.1 kmolc ha(-1) yr(-1) (30-45 kg N ha(-1) yr(-1)), respectively. It is concluded that the present atmospheric nitrogen deposition is a continuous threat for the existence of heathlands in Western Europe.     

Nitrogen deposition and the biodiversity of boreal forests: implications for the nitrogen critical load

The critical load concept is used to establish the deposition levels which ecosystems can tolerate without significant harmful effects. Here we summarize work within the Swedish research program Abatement Strategies for Transboundary Air Pollution (ASTA) assessing the critical load of N for boreal forests. Results from both field experiments in an area with low background N deposition in northern Sweden, and from a large-scale monitoring study, show that important vegetational changes start to take place when adding low N doses and that recovery of the vegetation after ceasing N input is a very slow process. The data presented indicate that changes in key ecosystem components occur even at a lower rate of N input than the present recommended empirical critical load for boreal forest understorey vegetation of 10-15 kg N ha(-1) yr(-1). Based on the data presented, we suggest that the critical load should be lowered to 6 kg N ha(-1) yr(-1).     

Nitrogen content, 15N natural abundance and biomass of the two pleurocarpous mosses Pleurozium schreberi (Brid.) Mitt. and Scleropodium purum (Hedw.) Limpr. in relation to atmospheric nitrogen deposition

The suitability of the two pleurocarpous mosses Pleurozium schreberi and Scleropodium purum for assessing spatial variation in nitrogen deposition was investigated. Sampling was carried out at eight sites in the western part of Germany with bulk deposition rates ranging between 6.5 and 18.5 kg N ha(-1) yr(-1). In addition to the effect of deposition on the nitrogen content of the two species, its influence on 15N natural abundance (delta15N values) and on productivity was examined. Annual increases of the mosses were used for all analyses. Significant relationships between bulk N deposition and nitrogen content were obtained for both species; delta15N-values reflected the ratio of NH4-N to NO3-N in deposition. A negative effect of nitrogen input on productivity, i.e. decreasing biomass per area with increasing N deposition due to a reduction of stem density, was particularly evident with P. schreberi. Monitoring of N deposition by means of mosses is considered an important supplement to existing monitoring programs. It makes possible an improved spatial resolution, and thus those areas that receive high loads of nitrogen are more easily discernible.     

Current and historical relationships between the tissue nitrogen content of a snowbed bryophyte and nitrogenous air pollution

Snowbed vegetation is under threat from atmospheric pollution. Most of the late lying snowbeds in Britain are in the central highlands of Scotland, coinciding with an area of very high deposition of nitrogenous air pollutants. Snow is a very efficient scavenger of atmospheric pollution and, due to the dynamics of snowmelt, much of the pollution load of a snow pack is released at very high concentrations in episodes known as 'acid flushes'. This study demonstrates the existence of a positive relationship between duration of snow-lie and tissue nitrogen content of Kiaeria starkei, a bryophyte characteristic of late snowbeds. An increase in the tissue nitrogen content of this bryophyte over this century is also shown, reflecting increasing air pollution. Maximum tissue nitrogen concentration in K. starkei is up to 50% greater than that recorded in other upland bryophyte species, demonstrating the exceptional threat of pollution to snowbed bryophytes. This has implications for the critical loads approach to pollution emission controls, as it indicates that some mountain communities are receiving higher pollution loadings than previously realised and therefore current exceedence of critical loads is probably greater than recognised at present.     

Short- and medium-term effects of experimental nitrogen fertilization on arthropods associated with Calluna vulgaris heathlands in north-west Spain

We studied the short- and medium-term effects of experimental nitrogen fertilization (3 and 15 months after the treatment) on the arthropods of Calluna vulgaris heathlands in NW Spain. Three heathland sites were selected with two permanent plots per site: control and fertilized. Ammonium nitrate fertilizer (56 kg N ha(-1) yr(-1)) was applied monthly and insects were caught using pitfall traps. We found mainly species-level responses to nitrogen addition. Seven species (e.g. Lochmaea suturalis) showed a consistent trend (benefited or harmed) in both periods and were proposed as possible reliable indicators of the effects of nitrogen deposition in these ecosystems. We also found variable arthropod trophic-group responses: (a) herbivores (leaf beetles, true bugs) increased in abundance on a short-term scale; (b) predators (carabid beetles, true bugs) showed opposite and less clear responses in both periods. Further long-term studies are needed to determine the mechanisms underlying the observed arthropod responses.     

Fluxes of oxidised and reduced nitrogen above a mixed coniferous forest exposed to various nitrogen emission sources

Concentrations of nitrogen gases (NH(3), NO(2), NO, HONO and HNO(3)) and particles (pNH(4) and pNO(3)) were measured over a mixed coniferous forest impacted by high nitrogen loads. Nitrogen dioxide (NO(2)) represented the main nitrogen form, followed by nitric oxide (NO) and ammonia (NH(3)). A combination of gradient method (NH(3) and NO(x)) and resistance modelling techniques (HNO(3), HONO, pNH(4) and pNO(3)) was used to calculate dry deposition of nitrogen compounds. Net flux of NH(3) amounted to -64 ng N m(-2) s(-1) over the measuring period. Net fluxes of NO(x) were upward (8.5 ng N m(-2) s(-1)) with highest emission in the morning. Fluxes of other gases or aerosols substantially contributed to dry deposition. Total nitrogen deposition was estimated at -48 kg N ha(-1) yr(-1) and consisted for almost 80% of NH(x). Comparison of throughfall nitrogen with total deposition suggested substantial uptake of reduced N (+/-15 kg N ha(-1) yr(-1)) within the canopy.     

Sulphur and seasalt deposition as reflected by throughfall and runoff chemistry in forested catchments

At forested catchments at Lake G?rdsj?n on the Swedish west coast the deposition and runoff chemistry has been followed during the period 1979-1990 by throughfall and runoff measurements as well as by measurements of atmospheric concentrations. The 10-year means in throughfall and runoff are very similar for sulphur and the main seasalt ions sodium and chloride; for sulphur 26.1 and 27.6 kg ha(-1) yr(-1), for sodium 49 and 52 kg ha(-1) yr(-1) and for chloride 96 kg ha(-1) yr(-1) and 93 kg ha(-1) yr(-1), respectively. The actual flows are 100-200% higher than the wet deposition as collected in open bulk precipitation collectors indicating a very large input by dry deposition. One important question is to what extent the throughfall and runoff values can be used as measures of total deposition. We present results from studies at different experimental catchments illustrating the possibilities of using throughfall and runoff data as measures of atmospheric deposition of sulphur and seasalt.     

Lichen and bryophyte distribution on oak in London in relation to air pollution and bark acidity

Epiphytic lichen and bryophyte distribution and frequency were investigated on the trunks of 145 young oak trees throughout London and surrounding counties, and compared with pollution levels and bark pH. Sixty-four lichen and four bryophyte species were recorded. Three major zones were identified: (i) two central regions with a few lichens, bryophytes absent; (ii) a surrounding region with a more diverse flora including a high cover of nitrophyte lichens; and (iii) an outer region, characterised by species absent from central London, including acidophytes. Nineteen species were correlated with nitrogen oxides and 16 with bark pH, suggesting that transport-related pollution and bark acidity influence lichen and bryophyte distribution in London today. Lichens and bryophytes are responding to factors that influence human and environmental health in London. Biomonitoring therefore has a practical role to assess the effects of measures to improve London's air quality.     

Empirical and simulated critical loads for nitrogen deposition in California mixed conifer forests

Empirical critical loads (CL) for N deposition were determined from changes in epiphytic lichen communities, elevated NO(3)(-) leaching in streamwater, and reduced fine root biomass in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) at sites with varying N deposition. The CL for lichen community impacts of 3.1kg ha(-1) year(-1) is expected to protect all components of the forest ecosystem from the adverse effects of N deposition. Much of the western Sierra Nevada is above the lichen-based CL, showing significant changes in lichen indicator groups. The empirical N deposition threshold and that simulated by the DayCent model for enhanced NO(3)(-)leaching were 17kg N ha(-1) year(-1). DayCent estimated that elevated NO(3)(-) leaching in the San Bernardino Mountains began in the late 1950s. Critical values for litter C:N (34.1), ponderosa pine foliar N (1.1%), and N concentrations (1.0%) in the lichen Letharia vulpina ((L.) Hue) are indicative of CL exceedance.     

The interactions between plant growth,vegetation structure and soil processes in semi-natural acidic and calcareous grasslands receiving long-term inputs of simulated pollutant nitrogen deposition

Regular applications of ammonium nitrate (35-140 kg N ha(-1) year(-1)) and ammonium sulphate (140 kg N ha(-1) year(-1)) to areas of acidic and calcareous grassland in the Derbyshire Peak District over a period of 6 years, have resulted in significant losses in both overall plant cover, and the abundance of individual species, associated with clear and dose-related increases in shoot nitrogen content. No overall growth response to nitrogen treatment was seen at any stage in the experiment. Phosphorus additions to the calcareous plots did however lead to significant increases in plant cover and total biomass, indicative of phosphorus limitation in this system. Clear and dose-related increases in soil nitrogen mineralization rates were also obtained, consistent with marked effects of the nitrogen additions on soil processes. High nitrification rates were seen on the calcareous plots, and this process was associated with significant acidification of the 140 kg N ha(-1) year(-1) treatments.     

Deposition of nitrogen-containing compounds to an extensively managed grassland in central Switzerland

During four intensive observation periods in 1992 and 1993, dry deposition of nitrogen dioxide (NO(2)) and ammonia (NH(3)), and wet deposition of nitrogen (N) were determined. The measurements were carried out in a small, extensively managed litter meadow surrounded by intensively managed agricultural land. Dry deposition of NH(3) was estimated by the gradient method, whereas eddy correlation was used for NO(2). Rates of dry deposition of total nitrate (= nitric acid (HNO(3)) + nitrate (NO(3)(-))), total nitrite (= nitrous acid (HONO) + nitrite (NO(2)(-))) and aerosol-bound ammonium (NH(4)(+)) were estimated using deposition velocities from the literature and measured concentrations. Both wet N deposition and the vertical NH(3) gradient were measured on a weekly basis during one year. Dry deposition was between 15 and 25 kg N ha(-1) y(-1), and net wet deposition was about 9.0 kg N ha(-1) y(-1). Daily average NO(2) deposition velocity varied from 0.11 to 0.24 cm s(-1). Deposition velocity of NH(3), was between 0.13 and 1.4 cm s(-1), and a compensation point between 3 and 6 ppbV NH(3) (ppb = 10(-9)) was found. Between 60 and 70% of dry deposition originated from NH(3) emitted by farms in the neighbourhood. It is concluded that total N deposition is exceeding the critical load for litter meadows, is highly correlated to local NH(3) emissions, and that NH(3) is of utmost importance with respect to possible strategies to reduce N deposition in rural regions.     

The effects of atmospheric nitrogen deposition in the Rocky Mountains of Colorado and southern Wyoming, USA-a critical review

The Rocky Mountains of Colorado and southern Wyoming receive atmospheric nitrogen (N) deposition that ranges from 2 to 7 kg ha(-1) yr(-1), and some previous research indicates pronounced ecosystem effects at the highest rates of deposition. This paper provides a critical review of previously published studies on the effects of atmospheric N deposition in the region. Plant community changes have been demonstrated through N fertilization studies, however, N limitation is still widely reported in alpine tundra and subalpine forests of the Front Range, and sensitivity to changes in snow cover alone indicate the importance of climate sensitivity in these ecosystems. Retention of N in atmospheric wet deposition is <50% in some watersheds east of the Continental Divide, which reflects low biomass and a short growing season relative to the timing and N load in deposition. Regional upward temporal trends in surface water NO(3)(-) concentrations have not been demonstrated, and future trend analyses must consider the role of climate as well as N deposition. Relatively high rates of atmospheric N deposition east of the Divide may have altered nutrient limitation of phytoplankton, species composition of diatoms, and amphibian populations, but most of these effects have been inconclusive to date, and additional studies are needed to confirm hypothesized cause and effect relations. Projected future population growth and energy use in Colorado and the west increase the likelihood that the subtle effects of atmospheric N deposition now evident in the Front Range will become more pronounced and widespread in the future.     

Effects of nitrogen enrichment on coastal dune grassland: a mesocosm study

Mesocosms filled with dune sand were planted with graminoid (Calamagrostis epigejos, Carex arenaria) and herbaceous species (Carlina vulgaris, Galium verum). Strong effects of nitrogen addition on the vegetation were found within two to three years. The above-ground biomass of C. epigejos and C. arenaria increased at deposition rates between 10 and 80 kg N ha(-1) yr(-1). Both grasses were limited by N. In latter stages P limitation was suggested for C. arenaria. At high N-levels, C. epigejos dominated the vegetation within two years. C. vulgaris and G. verum declined drastically as a result of increased competition for light by the highly competitive grass C. epigejos. It is concluded that increased (ambient) N inputs are of major importance for the increased dominance of tall grasses in stable dune grasslands.     

Effects of increased deposition of atmospheric nitrogen on an upland moor: leaching of N species and soil solution chemistry

This study was designed to investigate the leaching response of an upland moorland to long-term (10 yr) ammonium nitrate additions of 40, 80 and 120 kg N ha(-1) yr(-1) and to relate this response to other indications of potential system damage, such as acidification and cation displacement. Results showed increases in nitrate leaching only in response to high rates of N input, in excess of 96 and 136 kg total N input ha(-1) yr(-1) for the organic Oh horizon and mineral Eag horizon, respectively. Individual N additions did not alter ammonium leaching from either horizon and ammonium was completely retained by the mineral horizon. Leaching of dissolved organic nitrogen (DON) from the Oh horizon was increased by the addition of 40 kg N ha(-1) yr(-1), but in spite of increases, retention of total dissolved nitrogen reached a maximum of 92% and 95% of 80 kg added N ha(-1) yr(-1) in the Oh and Eag horizons, respectively. Calcium concentrations and calcium/aluminium ratios were decreased in the Eag horizon solution with significant acidification mainly in the Oh horizon leachate. Nitrate leaching is currently regarded as an early indication of N saturation in forest systems. Litter C:N ratios were significantly lowered but values remained above a threshold predicted to increase leaching of N in forests.     

Effects of increased deposition of atmospheric nitrogen on an upland Calluna moor: N and P transformations

This study determined the effects of increased N deposition on rates of N and P transformations in an upland moor. The litter layer and the surface of the organic Oh horizon were taken from plots that had received long-term additions of ammonium nitrate at rates of 40, 80 and 120 kg N ha(-1) yr(-1). Net mineralisation processes were measured in both field and laboratory incubations. Soil phosphomonoesterase (PME) activity and rates of N(2)O release were measured in laboratory incubations and root-surface PME activity measured in laboratory microcosms using Calluna vulgaris bioassay seedlings. Net mineralisation rates were relatively slow, with net ammonification consistently stimulated by N addition. Net nitrification was marginally stimulated by N addition in the laboratory incubation. N additions also increased soil and root-surface (PME) activity and rates of N(2)O release. Linear correlations were found between litter C:N ratio and all the above processes except net nitrification in field incubations. When compared with data from a survey of European forest sites, values of litter C:N ratio were greater than a threshold below which substantial, N input-related increases in net nitrification rates occurred. The maintenance of high C:N ratios with negligible rates of net nitrification was associated with the common presence of ericaceous litter and a mor humus layer in both this moorland as well as the forest sites.     

Cumulative impact of 40 years of industrial sulfur emissions on a forest soil in west-central Alberta (Canada)

The impact of 40 years of sulfur (S) emissions from a sour gas processing plant in Alberta (Canada) on soil development, soil S pools, soil acidification, and stand nutrition at a pine (Pinus contorta x Pinus banksiana) ecosystem was assessed by comparing ecologically analogous areas subjected to different S deposition levels. Sulfur isotope ratios showed that most deposited S was derived from the sour gas processing plant. The soil subjected to the highest S deposition contained 25.9 kmol S ha(-1) (uppermost 60 cm) compared to 12.5 kmol S ha(-1) or less at the analogues receiving low S deposition. The increase in soil S pools was caused by accumulation of organic S in the forest floor and accumulation of inorganic sulfate in the mineral soil. High S inputs resulted in topsoil acidification, depletion of exchangeable soil Ca2+ and Mg2+ pools by 50%, podzolization, and deterioration of N nutrition of the pine trees.     

Are N and S deposition altering the mineral composition of Norway spruce and Scots pine needles in Finland?

Data from a large-scale foliar survey were used to calculate the extent to which N and S deposition determined the mineral composition of Scots pine and Norway spruce needles in Finland. Foliar data were available from 367 needle samples collected on 36 plots sampled almost annually between 1987 and 2000. A literature study of controlled experiments revealed that acidifying deposition mediates increasing N and S concentrations, and decreasing Mg:N and Ca:Al ratios in the needles. When this fingerprint for N and S elevated deposition on tree foliage was observed simultaneously with increased N and S inputs, it was considered sufficient evidence for assuming that acidifying deposition had altered the mineral composition of tree needles on that plot in the given year. Evidence for deposition-induced changes in the mineral composition of tree foliage was calculated on the basis of a simple frequency model. In the late eighties the evidence was found on 43% of the Norway spruce and 27% of Scots pine plots. The proportion of changed needle mineral composition decreased to below 8% for both species in the late nineties.     

The impact of nitrogen deposition on acid grasslands in the Atlantic region of Europe

A survey of 153 acid grasslands from the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is changing plant species composition and soil and plant-tissue chemistry. Across the deposition gradient (2-44 kg N ha⁻¹ yr⁻¹) grass richness as a proportion of total species richness increased whereas forb richness decreased. Soil C:N ratio increased, but soil extractable nitrate and ammonium concentrations did not show any relationship with nitrogen deposition. The above-ground tissue nitrogen contents of three plant species were examined: Agrostis capillaris (grass), Galium saxatile (forb) and Rhytidiadelphus squarrosus (bryophyte). The tissue nitrogen content of neither vascular plant species showed any relationship with nitrogen deposition, but there was a weak positive relationship between R. squarrosus nitrogen content and nitrogen deposition. None of the species showed strong relationships between above-ground tissue N:P or C:N and nitrogen deposition, indicating that they are not good indicators of deposition rate.