Regional patterns in foliar (15)N across a gradient of nitrogen deposition in the northeastern US

Recent studies have demonstrated that natural abundance (15)N can be a useful tool for assessing nitrogen saturation, because as nitrification and nitrate loss increase, delta(15)N of foliage and soil also increases. We measured foliar delta(15)N at 11 high-elevation spruce-fir stands along an N deposition gradient in 1987-1988 and at seven paired northern hardwood and spruce-fir stands in 1999. In 1999, foliar delta(15)N increased from -5.2 to -0.7 per thousand with increasing N deposition from Maine to NY. Foliar delta(15)N decreased between 1987-1988 and 1999, while foliar %N increased and foliar C:N decreased at most sites. Foliar delta(15)N was strongly correlated with N deposition, and was also positively correlated with net nitrification potential and negatively correlated with soil C:N ratio. Although the increase in foliar %N is consistent with a progression towards N saturation, other results of this study suggest that, in 1999, these stands were further from N saturation than in 1987-1988.     

Soil acidification and foliar nutrient status of Ontario's deciduous forest in 1986 and 2005

To assess the impacts of the decline in sulphur (S) deposition over the past 20 years in Ontario, soil chemistry and sugar maple (Acer saccharum Marsh) foliar chemistry were measured at 17 sites in south and central Ontario in 2005 and compared with archived samples collected in 1986. Foliar S concentrations were lower in 2005, reflecting the decline in S deposition whereas foliar N remained unchanged, reflecting the lack of change in N deposition in Ontario. Mineral soil pH, exchangeable base cations were lower in 2005 whereas total S, N and cation exchange capacity (CEC) were unchanged. Foliar concentrations of Ca were positively related to soil Ca levels in the A-horizon and were lower in 2005. Despite evidence of increasing soil acidity and losses of calcium, foliar base cation concentrations at most sites were adequate for sugar maple and forest health in terms of canopy appearance (Decline Index) improved.     

Inferred effects of cloud deposition on forest floor nutrient cycling and microbial properties along a short elevation gradient

Cloud water deposition often increases with elevation, and it is widely accepted that this cloud water increases acid loading to upland forest ecosystems. A study was undertaken in south-eastern Quebec to determine if a 250 m elevation gradient (i.e. 420-665 m), along a uniform sugar-maple stand on the slope of Mount Orford, corresponded to a pH gradient in the forest floor and to predictable changes in soil nutrient availability and microbial properties. Precipitation data from a nearby study, and a photographic survey, provided presumptive evidence that this elevation gradient corresponded to a strong gradient in cloud water deposition. Forest floor temperature did not differ significantly across elevations. Forest floor moisture content was significantly higher, whereas pH and exchangeable Ca and Mg were significantly lower, at the higher elevations. Average seasonal net nitrification rates, determined by long-term laboratory incubations, did not differ significantly across elevations, whereas average seasonal net ammonification rates were significantly higher at higher elevations. Basal respiration rates and microbial biomass did not differ significantly across elevations, but metabolic quotient was significantly higher at higher elevations indicating possible environmental stress on forest floor microbial communities due to cloud water deposition. Anaerobic N mineralisation rates were significantly higher at higher elevations suggesting that N-limited microbial communities frequently exposed to cloud cover can be important short-term sinks for atmospheric N, thereby contributing to increase the active-N fraction of forest floors. We conclude that, where no significant changes in vegetation or temperature occur, elevation gradients can still be used to understand the spatial variability of nutrient cycles and microbial properties.     

Spatial and temporal small-scale variability of nitrogen mobilization in a forest ecosystem with high N deposition in NW-Germany

For conifer stands in NW-Germany with high DIN load (23-35 kg N ha⁻¹ a⁻¹) and a long history of nitrogen export the risk of N mobilization were investigated. Ammonium is the most mobilized N species, pointing towards either conditions not favoring nitrification or, more likely - under the dominant aerobic conditions - a very high amount of ammonium in the forest floor. Independence of net nitrification and net ammonification from each other indicates the existence of two separate systems. The nitrifying system depends very much on biotic conditions - as a function of energy and moisture - and seems not to be directly related to N deposition. In contrast, for the ammonification system (Oe horizon) a correlation with the sum of ammonium deposition three months prior to sampling was found. However, the role of disturbance, i.e. nitrogen export, during the last centuries and the role of recovery of the N balance during the last 150 years is still not clear.     

Acidification-induced chemical changes in coniferous forest soils in southern Sweden 1988-1999

Thirty-two Norway spruce [Picea abies (L.) Karst.] and Scots pine (Pinus sylvestris L.) stands in southern Sweden were studied for a period of 12 years to evaluate acidification-induced chemical changes in the soil. Soil, at 20-30 cm depth in the mineral layer, was sampled three times during this period (1988, 1993 and 1999). The results show that pH(BaCl2) in mineral soil decreased by, on average, 0.17 units between 1988 and 1999, accompanied by an increase in aluminium (Al) concentration and a decrease in base saturation in the soil. In 1999, the base saturation was below 5% in 58% of the 32 sites compared with 16% in 1988 and 7% in 1993. Concentrations of calcium (Ca), potassium (K) and magnesium (Mg) are low and decreasing. Based on C/N ratios in humus, 45% of the sites may be subjected to leaching of considerable amounts of nitrate. The results show that the acidification of coniferous forest soils in southern Sweden is continuing, and that the negative effects on the nutrient status in soil are extensive. The results are compared with reference values for productive, long-term sustainably managed boreal coniferous or mixed forest soils and implications for long-term sustainability are discussed.     

Physiological and foliar symptom response in the crowns of Prunus serotina, Fraxinus americana and Acer rubrum canopy trees to ambient ozone under forest conditions

The crowns of five canopy dominant black cherry (Prunus serotina Ehrh.), five white ash (Fraxinus americana L.), and six red maple (Acer rubrum L.) trees on naturally differing environmental conditions were accessed with scaffold towers within a mixed hardwood forest stand in central Pennsylvania. Ambient ozone concentrations, meteorological parameters, leaf gas exchange and leaf water potential were measured at the sites during the growing seasons of 1998 and 1999. Visible ozone-induced foliar injury was assessed on leaves within the upper and lower crown branches of each tree. Ambient ozone exposures were sufficient to induce typical symptoms on cherry (0-5% total affected leaf area, LAA), whereas foliar injury was not observed on ash or maple. There was a positive correlation between increasing cumulative ozone uptake (U) and increasing percent of LAA for cherry grown under drier site conditions. The lower crown leaves of cherry showed more severe foliar injury than the upper crown leaves. No significant differences in predawn leaf water potential (psi(L)) were detected for all three species indicating no differing soil moisture conditions across the sites. Significant variation in stomatal conductance for water vapor (g(wv)) was found among species, soil moisture, time of day and sample date. When comparing cumulative ozone uptake and decreased photosynthetic activity (P(n)), red maple was the only species to show higher gas exchange under mesic vs. drier soil conditions (P < 0.05). The inconsistent differences in gas exchange response within the same crowns of ash and the uncoupling relationship between g(wv) and P(n) demonstrate the strong influence of heterogeneous environmental conditions within forest canopies.     

Evidence from nitrogen fertilisation in the forests of Germany

In the first part, this contribution presents German results of nitrogen fertilisation experiments on stand growth, published 1958-1987. Over this period Norway spruce and Scots pine have generally responded positively to nitrogen fertilisation, without showing signs of damage, at levels up to at least 1000 kg N ha(-1). In the second part, growth patterns of forest trees and stands in Southern Germany are presented. Norway spruce in particular is now growing better than in earlier years of this century, starting around 1960. It is plausible that increasing nitrogen deposition, which coincides with the growth increase, is the cause, though this cannot be shown unequivocally. At a few sites with poor soil and management and high acid deposition a deterioration is taking place.     

Rain and throughfall chemistry in a Norway spruce forest in the Western Prealps

From January 1987 to December 1989 precipitation was collected in an open field in a forested area of the Western Prealps. The site faces the intensely industrialized area of the Po Valley. In a Norway spruce plantation, next to the open field sampler, throughfall was collected from January 1987 to August 1989 for a period of 32 months. In the common sampling period median pH values of the open field precipitation and throughfall were 4.28 and 4.19 respectively. Open field precipitation has yearly deposition rates of acidity, Ca, S, and N of 0.81, 9.82, 24.83 and 32.81 kg/ha(-1) respectively. The throughfall showed significant increases in the deposition rates for Ca, Mg, K, N and S, due to abundant dry deposition and, probably, to nutrient leaching contributed by the canopies of the spruce. On the basis of available data, an initial approach to atmosphere-canopy interaction was made.     

Foliar and ground deposits of fenitrothion following aerial application over a plantation forest

Abstract

Spray deposits were measured on spruce foliage at tree canopy level and on glass plates at ground level, after aerial application of an emulsion formulation of fenitrothion at the rate of 0.21 kg AI in 1.46 L per ha over a 16 ha plot in a plantation forest. Fenitrothion deposits were quantified by gas‐liquid chromatography. A wide variation was observed in deposits on foliage and at the forest floor. Deposits were relatively higher on foliar samples collected from the upwind side of a tree canopy than those on the downwind side. Similarly, the glass plate placed under a tree on the upwind side received relatively higher deposit than the one on the downwind side. However, the glass plates placed in the adjacent forest openings collected markedly higher deposits. Results clearly indicate filtration of the spray droplets by canopy foliage. Assessment of the average deposit of fenitrothion at ground level (mean deposit from all sampling locations) indicated that ca. 19.4% of the applied material reached the forest floor. Within a sampling station, correlation was poor between foliar depsits and those on glass plates under the same trees or in nearby clearings. Analysis of fenitrothion deposits on foliage collected at 1 and 2 h after application indicated that the droplets took, more than 1 h for deposition on the tree canopy. On the other hand, deposition on the glass plates at ground level appeared to be practically complete within 1 h post‐treatment. This was attributed to the higher sedimentation velocities of the larger droplets which tend to travel faster to the floor level than the smaller droplets which float for a longer period near the tree canopy.     

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.     

Detrital control on the release of dissolved organic nitrogen (DON) and dissolved inorganic nitrogen (DIN) from the forest floor under chronic N deposition

The role of detrital quantity and quality in forest floor N leaching was investigated in a litter manipulation experiment at a deciduous forest under chronic N deposition. Dissolved inorganic nitrogen (DIN) comprised the bulk of nitrogen leaching from the control except a short period following autumn litterfall. The dominance of DIN was strengthened by litter exclusion, whereas the addition of glucose or fresh litter led to a small increase in dissolved organic nitrogen (DON) and either a temporary or gradual reduction in NO(3)(-) release, respectively. Changes in soluble organic C and microbial C in the forest floor implied that increased availability of C sources might have enhanced microbial immobilization of DIN, either temporarily following glucose application or over the longer term following litter addition. The results suggest that detrital quantity and quality can play a crucial role in determining the balance between DIN and DON in N-enriched forest soils.     

Regional estimation of current and future forest biomass

The 90,674 wildland fires that burned 2.9 million ha at an estimated suppression cost of $1.6 billion in the United States during the 2000 fire season demonstrated that forest fuel loading has become a hazard to life, property, and ecosystem health as a result of past fire exclusion policies and practices. The fire regime at any given location in these regions is a result of complex interactions between forest biomass, topography, ignitions, and weather. Forest structure and biomass are important aspects in determining current and future fire regimes. Efforts to quantify live and dead forest biomass at the local to regional scale has been hindered by the uncertainty surrounding the measurement and modeling of forest ecosystem processes and fluxes. The interaction of elevated CO2 with climate, soil nutrients, and other forest management factors that affect forest growth and fuel loading will play a major role in determining future forest stand growth and the distribution of species across the southern United States. The use of satellite image analysis has been tested for timely and accurate measurement of spatially explicit land use change and is well suited for use in inventory and monitoring of forest carbon. The incorporation of Landsat Thematic Mapper data coupled with a physiologically based productivity model (PnET), soil water holding capacity, and historic and projected climatic data provides an opportunity to enhance field plot based forest inventory and monitoring methodologies. We use periodic forest inventory data from the USDA Forest Service's Forest Inventory and Analysis (FIA) project to obtain estimates of forest area and type to generate estimates of carbon storage for evergreen, deciduous, and mixed forest classes for use in an assessment of remotely sensed forest cover at the regional scale for the southern United States. The displays of net primary productivity (NPP) generated from the PnET model show areas of high and low forest carbon storage potential and their spatial relationship to other landscape features for the southern United States. At the regional scale, predicted annual NPP in 1992 ranged from 836 to 2181 g/m2/year for evergreen forests and 769-2634 g/m2/year for deciduous forests with a regional mean for all forest land of 1448 g/m2/year. Prediction of annual NPP in 2050 ranged from 913 to 2076 g/m2/year for evergreen forest types to 1214-2376 g/m2/year for deciduous forest types with a regional mean for all forest land of 1659 g/m2/year. The changes in forest productivity from 1992 to 2050 are shown to display potential areas of increased or decreased forest biomass. This methodology addresses the need for spatially quantifying forest carbon in the terrestrial biosphere to assess forest productivity and wildland fire fuels.     

A conceptual framework: Redefining forest soil's critical acid loads under a changing climate

Federal agencies of several nations have or are currently developing guidelines for critical forest soil acid loads. These guidelines are used to establish regulations designed to maintain atmospheric acid inputs below levels shown to damage forests and streams. Traditionally, when the critical soil acid load exceeds the amount of acid that the ecosystem can absorb, it is believed to potentially impair forest health. The excess over the critical soil acid load is termed the exceedance, and the larger the exceedance, the greater the risk of ecosystem damage. This definition of critical soil acid load applies to exposure of the soil to a single, long-term pollutant (i.e., acidic deposition). However, ecosystems can be simultaneously under multiple ecosystem stresses and a single critical soil acid load level may not accurately reflect ecosystem health risk when subjected to multiple, episodic environmental stress. For example, the Appalachian Mountains of western North Carolina receive some of the highest rates of acidic deposition in the eastern United States, but these levels are considered to be below the critical acid load (CAL) that would cause forest damage. However, the area experienced a moderate three-year drought from 1999 to 2002, and in 2001 red spruce (Picea rubens Sarg.) trees in the area began to die in large numbers. The initial survey indicated that the affected trees were killed by the southern pine beetle (Dendroctonus frontalis Zimm.). This insect is not normally successful at colonizing these tree species because the trees produce large amounts of oleoresin that exclude the boring beetles. Subsequent investigations revealed that long-term acid deposition may have altered red spruce forest structure and function. There is some evidence that elevated acid deposition (particularly nitrogen) reduced tree water uptake potential, oleoresin production, and caused the trees to become more susceptible to insect colonization during the drought period. While the ecosystem was not in exceedance of the CAL, long-term nitrogen deposition pre-disposed the forest to other ecological stress. In combination, insects, drought, and nitrogen ultimately combined to cause the observed forest mortality. If any one of these factors were not present, the trees would likely not have died. This paper presents a conceptual framework of the ecosystem consequences of these interactions as well as limited plot level data to support this concept. Future assessments of the use of CAL studies need to account for multiple stress impacts to better understand ecosystem response.     

Influence of foliar morphology and crown geometry on insecticide deposition and dissipation following an aerial spray over a mixed forest

Abstract

Deposition and dissipation of aerially sprayed phosphamidon insecticide over a mixed forest consisting of different species of coniferous and deciduous trees were studied. The data showed that the chemical deposited more and dissipated slower from deciduous foliage than from conifer needles. Differences in deposits and persistence were also found within each of the two foliar types. Leaf shape and orientation, crown geometry and foliar morphology appear to have played a role. No correlation was found between initial deposit levels and cuticular wax content of different foliar species. A controlled laboratory study showed that surface area/mass ratio played a more important role on initial deposits than pubescence/serrations present on foliar surfaces.     

Modelling changes in forest soil chemistry at 16 Swedish coniferous forest sites following deposition reduction

The dynamic forest ecosystem model ForSAFE was applied at 16 coniferous forest sites in Sweden to investigate past and future changes in soil chemistry following changes in atmospheric deposition. The simulation shows a considerable historical soil acidification. Acidification in the southwest, where deposition has been greatest, was more expressed in the deepest soil layers, while it was more evenly distributed through the soil profile in central Sweden, and was greater in the upper soil layers in the north. The simulation also shows that a slight recovery took place after the reduction in emissions, but was counteracted by the effect of harvesting. The simulation predicts an increase in the number of acidified sites in the future. The results also suggest that future acidification will be mainly due to the enhanced tree growth resulting from the chronic high deposition of nitrogen and the removal of soil base cations through harvesting.     

Dry deposition and internal circulation of nitrogen,sulphur and base cations to a coniferous forest

The dry deposition of sulphur, nitrogen and base cations to a spruce stand was estimated during a five year period using a surrogate surface resembling needles, throughfall and bulk deposition measurements. The deposition was calculated from the ratio between the deposition of an ion and sodium on the surrogate surface and the net throughfall of sodium to the forest. The dry deposition represented a large fraction of the total atmospheric input of base cations. For Na⁺, Mg²⁺, Ca²⁺, and K⁺ they were 66, 67, 53 and 42%, respectively. The internal circulation was 95% of non-marine net throughfall fro K⁺ and 76% for Ca²⁺. The dry deposition of SO₂ to the canopies regulates the internal circulation of Ca²⁺. The dry deposition of SO₂ to the canopies regulates the internal circulation of Ca²⁺. The dry depositions of ammonium and nitrate are close to the net throughfall of Kjeldahl-N and nitrate, respectively. The obtained deposition velocities are comparable to other studies. The calculated dry deposition of ammonium was compared to the net throughfall of ammonium at three nearby forest stands receiving different ammonium amounts on the soils. No correlation to nitrogen level was found, but most ammonium was lost and converted to organic nitrogen in the canopies of the wettest forest stand.     

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.     

Atmospheric deposition to the edge of a spruce forest in Denmark

Atmospheric deposition was measured during 1 year at the forest edge of a Norway spruce stand in Denmark. Inside the forest the deposition of H(+), Ca(2+), Mg(2+), Na(+), K(+), Cl(-), NO(3)(-), NH(4)(2) and SO(4)(2-) with canopy throughfall varies with the distance from the forest edge. The deposition at the edge is found to be 10-20 times as high as deposition to an open field and 2-8 times as high as deposition inside the stand. An exponential decrease in deposition as a function of the distance from the forest edge is found. Increased deposition of K(+) and non-sea salt Mg(2+), which mainly originates as a result of leaching from the needles may be explained by a larger leaf area index (LAI) at the forest edge. Deposition of particulate substances, especially Na(+), Cl(-), Mg(2+) and to some extent SO(4)(2-), NH(4)(+) and NO(3)(-) is increased much more than the LAI, which we believe to be caused by changes in wind movements at the forest edge.     

Spatial variability of throughfall fluxes in a spruce forest

The spatial variability of throughfall deposition of H(+), Ca(2+), Mg(2+), Na(+), K(+), Cl(-), NO(3)(-), NH(4)(+), O(4)(2-) to a Norway spruce (Picea abies (L.) Karst.) forest was intensively examined during the period October 1986 to October 1987. Large systematic spatial variability of the atmospheric deposition within the forest was observed. The flux of throughfall water was higher away from the trunk compared to the flux close to the trunk. In contrast to this, the deposition of all substances was considerably higher close to the trunk compared to the deposition at the periphery of the canopy. A linear decrease in deposition as a function of the distance from the nearest tree trunk was found. Further, the deposition varied quite dramatically between trees according to their size. The observed spatial variability in throughfall may be due to variabilities in the processes taking part in altering the distribution and composition of the precipitated water as it moves through the canopy. The influence of these processes of precipitation, wash-off, dry deposition and canopy exchange is discussed, and it is found that both increased dry deposition and canopy exchange in the tree tops contribute to the higher solute fluxes found close to the tree trunk.     

Can the foliar nitrogen concentration of upland vegetation be used for predicting atmospheric nitrogen deposition? Evidence from field surveys

The deposition of atmospheric nitrogen can be enhanced at high altitude sites as a consequence of cloud droplet deposition and orographic enhancement of wet deposition on hills. The degree to which the increased deposition of nitrogen influences foliar nitrogen concentration in a range of upland plant species was studied in a series of field surveys in northern Britain. A range of upland plant species sampled along altitudinal transects at sites of known atmospheric nitrogen deposition showed marked increases in foliar nitrogen concentration with increasing nitrogen deposition and altitude (and hence with decreasing temperature). For Nardus stricta L., Deschampsia flexuosa (L.) Trin., Calluna vulgaris (L.) Hull, Erica cinerea L. and Hylocomium splendens (Hedw.) Br. Eur. on an unpolluted hill, foliar nitrogen increased by 0.07, 0.12, 0.15, 0.08 and 0.04% dry weight respectively for each 1 kg ha(-1) year(-1) increase in nitrogen deposition. Most species showed an approximately linear relationship between foliar nitrogen concentration and altitude but no trend with altitude for foliar phosphorus concentration. This provided evidence that the tissue nutrient status of upland plants reflects nutrient availability rather than the direct effects of climate on growth. However, differences in the relationship between foliar nitrogen concentration and atmospheric nitrogen deposition for N. stricta sampled on hills in contrasting pollution climates show that the possibility of temperature-mediated growth effects on foliar nitrogen concentration should not be ignored. Thus, there is potential to use upland plant species as biomonitors of nitrogen deposition, but the response of different species to nitrogen addition, in combination with climatic effects on growth, must be well characterised.