The effects of atmospheric nitrogen deposition on the soil chemistry of coniferous forests in The Netherlands

Nitrogen fluxes, particularly those of ammonium, are extremely high in Dutch forests. In soils exposed to high ammonium deposition, acidification, eutrophication or a combination of both processes may occur. In addition to the amounts of ammonium deposited, the rate of soil nitrification determines which process takes place. A nation-wide investigation, in which three coniferous tree species were involved, was carried out to study the relation between deposition fluxes, measured by means of throughfall and bulk samplers, and the chemical composition of the soil. The ammonium deposition accounted directly for the high ammonium content and the high ammonium/cation ratios in the soil. In the top layer of most of the forest soils which were investigated nitrification rates were low. In these stands ammonium/cation ratios in the soil often reflected ammonium/cation ratios in throughfall water. Even in soils with relatively high nitrification rates, ammonium concentrations exceeded those of nitrate in the top layer of the mineral soil, indicating that ammonium deposition was more important than nitrification rate in determining the predominant form of nitrogen.     

Impact of artificial, ammonium-enriched rainwater on soils and young coniferous trees in a greenhouse. Part II--Effects on the trees

To obtain an insight into the effects of a high ammonium deposition on trees young, coniferous trees were planted in pots in a greenhouse and treated with artificial ammonium-enriched rainwater. Application of 480 kg ammonium N ha(-1) year(-1) resulted in an increase of the shoot/root ratio. The biomass of fine roots strongly declined, as did the numbers of mycorrhizae. The fructification of the mycorrhizal fungi was totally inhibited. The nitrogen content of the needles was enhanced, but the levels of potassium, magnesium and calcium decreased sharply. The phosphorus content remained almost unaffected. Much of the nitrogen was stored as arginine. The levels of leaf pigments also increased. Within one year of treatment many of the trees died. The trees that were treated with 48 kg ammonium-N ha(-1) year(-1) did not show any signs of deterioration.     

Response of magnesium-deficient saplings in a young, open stand of Picea abies (L.) Karst. to elevated soil magnesium, nitrogen and carbon

A decline in a Picea abies L. (Karst.) stand in the Fichtelgebirge, NE-Bavaria, FRG has been attributed to a nutritional disharmony-a seasonal imbalance between a high supply of nitrogen, caused by high nitrogen deposition, and a low supply of soil magnesium, caused by soil acidification (Oren et al., 1988a). The nutritional disharmony hypothesis was tested on ten-year-old P. abies saplings in an adjacent stand growing on identical soil. The supply rate of magnesium relative to nitrogen was continuously increased or decreased during three successive growing seasons. Increasing the nitrogen or carbon supply resulted in a small increase in foliar nitrogen concentrations. Magnesium or carbon addition slightly raised the concentration of magnesium in the foliage. Reduction in crown leaf-area did not result in any appreciable changes in nutrient concentrations. Increased N supply decreased foliar Mg concentrations. In spite of the changes in the nutritional status of the needles, gas-exchange rates, pigment concentrations, needle characteristics and growth of twigs and stems did not differ among treatments. It appears that the growth of saplings was unimpaired at the foliar magnesium concentration at which the growth of adjacent mature trees was reduced. Moreover, it was not possible to promote nutritional disharmony in the saplings, including those receiving three times the annual nitrogen input. The study demonstrates that in young, relatively open stands of P. abies, much of the deposited nitrogen is not absorbed by the roots of saplings. Thus, the conceptual model of nutritional disharmony cannot explain forest decline if nitrogen uptake does not increase with deposition. Identifying the processes which control the uptake relative to the supply of all nutrients, and quantifying the rates of nutrient uptake are essential steps in using the conceptual model to explain specific decline symptoms.     

Flow rates of ions in waters percolating through a model ecosystem with forest trees

From 1983-88 the long-term effects of low level exposure with O(3), SO(2) and simulated acid rain on mineral cycling in model ecosystems with spruce, fir and beech seedlings were investigated. Systems consisting of open-top chambers built above lysimeters were protected against the intrusion of ambient rain and dust. As part of the investigations on mineral cycling the fluxes of elements with water input and output of the canopy and soil compartments are presented. During the 5 year duration of the experiment, pronounced effects on canopy deposition and cation leaching were observed. Most noticeable were throughfall enrichment with sulfate through dry deposition of SO(2) as influenced by duration of needlewetting and factors promoting SO(2) oxidation. Depending on sulfur deposition, leaching of calcium, magnesium, manganese, zinc and ammonium from canopies was elevated, in total leading to enhanced soil input of acid. After 15 months, the water percolating the soils in the lysimeters of these treatments was acidified, with elevated flowrates of sulfate, manganese, calcium and magnesium. The results on canopy/soil leaching are compared to those from old conifer stands in the field.     

Trends in needle and soil chemistry of Norway spruce and Scots pine stands in South Sweden 1985-1994

The nutrient status of the trees and soil in 42 stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in Scania, South Sweden was followed from 1985 to 1994. Samples from needles taken in winter 1985, 1987, 1990, and 1994, and soils in 1988 and 1993 were analyzed. Concentrations, as well as ratios to N, of K and Cu in needles of both species decreased by approximately 40% from 1985 to 1994. Soil analyses indicate ongoing soil acidification and leaching of mineral nutrients from the soil profile. Together with deposition data and corroboration from modeled scenarios, these data support the recent contention that one consequence of enhanced deposition of N and S will be the development of nutrient imbalances in trees growing in southern Sweden.     

Anthropogenic impacts on natural nitrogen isotope variations in Pinus sylvestris stands in an industrially polluted area

Natural variations of the nitrogen isotopes 15N/14N (delta15N values) and the N concentrations of one-year-old needles from 7-12-year-old pine trees (Pinus sylvestris L.) were determined on 27 sites in the heavily polluted Leipzig-Halle region (former GDR). At three selected sites measurements were repeated over a period of 2 years. N concentrations and delta15N values in different needle age classes were compared at the three sites. The delta15N values of the N in the humus layer and the potential plant available N in the A(h) horizon of the local soil were determined. The 15N/14N isotope ratios (delta15N values) of one-year-old pine needles in the region of Leipzig-Halle were found to vary depending on their specific location by a factor of up to one order of magnitude (-9.6 per thousand to + 0.4 per thousand ). N concentrations in one-year-old pine needles varied between 0.71 and 1.38 mmol eq N g dw(-1). Pine stands with positive or slightly negative delta15N values and high N concentrations in one-year-old needles were concentrated around the cities of Leipzig and Halle and in the industrial areas. More negative delta15N values and lower N concentrations in one-year-old pine needles were found on sites at greater distances from the industrial agglomerations, mainly in the NE forested part. Site specific differences in the delta15N values of the N in the humus layer from three selected sites were similar to those found for the needles. No site specific differences, however, were found for the delta15N values of the water soluble nitrogen fraction from the mineral soil horizons of the same sites.     

Effects of ammonium and aluminium on the development and nutrition of Pinus nigra in hydroculture

Application of ammonium and aluminium to young Pinus nigra var. maritima (Ait.) Melville trees resulted in a variety of negative effects. Excess ammonium led to an increase in shoot/root ratio. The biomass of the fine roots declined, resulting in an increase of the coarse/fine root ratio. The degree of mycorrhizal infection of the roots decreased. The nitrogen content of the trees increased considerably, whereas particularly the levels of calcium magnesium, manganese and zinc decreased sharply. Excess aluminium resulted in a simultaneous reduction of root and shoot biomass, a decline of the fine root system, an increase in the coarse/fine root ratio and a decrease in the degree of mycorrhizal infection. Uptake of the divalent cations calcium, magnesium, iron, manganese and zinc was restricted substantially, The nitrogen and phosphorus contents of the trees were hardly affected, whilst the potassium content of the shoot increased and of the roots decreased. This implicates that a deteriorating fine root system has to supply water and nutrients to a more demanding shoot. In the long term, high ammonium inputs and aluminium dissolution in forest ecosystems will lead to substantial nutrient deficiencies, just as has been found in the field.     

Influence of air pollution on the foliar nutrition of conifers in Great Britain

The nutritional status of needles from Sitka spruce, Norway spruce and Scots pine in a total of 108 stands was assessed. There was little evidence of nutritional deficiency, although potassium levels were frequently quite low. Analysis of some heavy metals (lead and copper) failed to reveal any likely toxicity problems. Sulphur, nitrogen and iron levels in/on the foliage were all related to various measures of sulphur and nitrogen pollution, determined using improved deposition models that take into account cloud deposition and the seeder-feeder mechanism. The analysis strongly suggested that direct air pollution has a greater effect on sulphur, nitrogen and iron foliar analyses than indirect pollution (wet deposition). The relationships were identified for levels of pollution that were generally lower than those seen in traditional gradient studies.     

Acid deposition and the element composition of pine tree rings

The element contents of pine tree rings (Pinus sylvestris L.) were analyzed in five areas differing in the level of sulphur deposition. The elements studied and their average content in xylem were as follows:

aluminium 4.2, manganese 65, zinc 8.4, potassium 390, rubidium 2.2, calcium 930, magnesium 131, phosphorus 38, sulphur 56, nitrogen 0.65, boron 4.3 and molybdenum 0.03 mg/kg.

The contents of aluminium, potassium and rubidium were higher in tree rings from the 1950's on, particularly in southern Finland. The increase is probably due to acid deposition in the soil.     

Does (K+Mg+Ca+P) fertilization lead to recovery of tree health in a nitrogen stressed Quercus rubra L. stand?

In The Netherlands atmospheric deposition of nitrogen compounds to forest ecosystems has been very high for some decades and has led to severe nutritional imbalances in soils as well as in trees. At this moment legislation is not fully in effect with respect to lowering emission/deposition fluxes, particularly of nitrogen. The trees suffer mainly from severe magnesium, potassium and calcium and sometimes phosphorus deficiencies. In this study it was investigated whether fertilization with potassium, magnesium, calcium and phosphorus could prevent a Quercus rubra stand from further decline by restoring the nutritional balance, though the detrimental input of nitrogen compounds still continues. Three fertilization treatments were applied: (i) a standard dose of 1250 kg.ha(-1) containing 60 kg P.ha(-1) (as P2O5), 100 kg K.ha(-1) (as K2O), 80 kg Mg.ha(-1) (as MgO) and 340 kg Ca.ha(-1) (as CaCO3); (ii) two times the standard dose; and (iii) three times the standard dose. Soil solution chemistry showed that the highest dose led to the most significant results: an improved nutrient balance and an increased availability of nutrients. After one growing season following fertilization, the trees looked much healthier and crown density had increased. This revitalization lasted for at least four years. For tree health the mid dose seemed appropriate. After the first growing season potassium is the most mobile nutrient, in the soil and also in the trees, but tended to decrease in the fourth year after fertilization. Magnesium and calcium reached normal values in the leaves after four growing seasons. No obvious effects of phosphorus were found.     

Comparison of throughfall and soil solution chemistry between a high-density Corsican pine stand and a naturally regenerated silver birch stand

In Flanders, critical loads for acidification and eutrophication are exceeded in the majority of the forest stands, and many previously nitrogen limited forest ecosystems have become nitrogen saturated. The present study investigates whether a naturally regenerated stand of silver birch (Betula pendula Roth) contributes less to the acidification and eutrophication of the forest soil than a high-density plantation of Corsican pine (Pinus nigra ssp. laricio Maire). Throughfall deposition of inorganic nitrogen was about 3.5 times higher in the Corsican pine stand than in the birch stand. Potassium throughfall deposition was significantly higher under birch due to higher canopy leaching. Magnesium throughfall deposition was significantly higher under the pine canopy due to higher dry deposition. The lower nitrogen throughfall deposition in the birch stand was reflected in a 60% lower nitrate percolation at 1m depth compared with pine. Nitrate soil percolation is linked to losses of aluminium and base cations.     

Nitrate reductase activity as an indicator of ponderosa pine response to atmospheric nitrogen deposition in the San Bernardino Mountains

Determinations of nitrate reductase (NR) activity in ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) needles were performed during summer 1994 in two areas (consisting of six different sites) with different nitrogen (N) deposition levels in the San Bernardino Mountains, southern California. Nitrate reductase activity was used as an integrative indicator of atmospheric nitrogen deposition to pine trees (direct uptake of N species from the atmosphere and N transported from the soil). Deposition of nitrate (NO3-) to pine branches was measured in order to determine dry atmospheric inputs of the oxidized N species to tree foliage. High NR activity was detected in all of the experimental sites. Activity of the enzyme was significantly higher at the locations characterized by higher NO3- deposition to branches--slight positive correlation between branch deposited NO3- and NR activity was found. However, high variability of NR in time and between the experimental sites discredit the NR assay as a reliable indicator of N deposition for ponderosa pine in the field conditions. This could be caused by substantial interference from other abiotic and biotic factors with tropospheric ozone as probably the most important one.     

Effects of acidic deposition on forest and aquatic ecosystems in New York State

Acidic deposition is comprised of sulfuric and nitric acids and ammonium derived from atmospheric emissions of sulfur dioxide, nitrogen oxides, and ammonia, respectively. Acidic deposition has altered soil through depletion of labile pools of nutrient cations (i.e. calcium, magnesium), accumulation of sulfur and nitrogen, and the mobilization of elevated concentrations of inorganic monomeric aluminum to soil solutions in acid-sensitive areas. Acidic deposition leaches essential calcium from needles of red spruce, making this species more susceptible to freezing injury. Mortality among sugar maples appears to result from deficiencies of nutrient cations, coupled with other stresses such as insect defoliation or drought. Acidic deposition has impaired surface water quality in the Adirondack and Catskill regions of New York by lowering pH levels, decreasing acid-neutralizing capacity, and increasing aluminum concentrations. Acidification has reduced the diversity and abundance of aquatic species in lakes and streams. There are also linkages between acidic deposition and fish mercury contamination and eutrophication of estuaries.     

Crown condition of Norway spruce in relation to sulphur and nitrogen deposition and soil properties in southeast Norway

Atmospheric deposition of sulphur and nitrogen compounds may lead to enhanced leaching of base cations, accumulation of nitrogen in organic matter, lowered pH and increased concentration of toxic aluminium in soil, which in turn may affect the vitality of forest trees. A general monitoring of forest condition has been initiated in many European countries, partly in order to reveal stresses caused by acidification. However, forest condition is also affected by many other factors. This paper examines a seven-year series of crown-condition data from Local County Monitoring Plots in Norway spruce stands in Norway. Average, time trend and lability variables were calculated for crown density and crown colour for each plot. Wet deposition of sulphate, ammonium and nitrate for each plot were estimated using data from the national air and precipitation monitoring programmes. Soil data are based on soil sampling within the plots. The analysis gave no evident support for the hypothesized negative effect on crown condition from sulphur and nitrogen deposition and related alterations in soil.     

Impact of artificial ammonium-enriched rainwater on soils and young coniferous trees in a greenhouse. Part I-effects on the soils

A pot experiment was used to investigate whether ammonium sulphate, when applied in amounts comparable to those deposited in the Dutch forests, is detrimental to trees. Young coniferous trees were planted in pots and treated with artificial ammonium-enriched rainwater in a greenhouse. The deposition of ammonium resulted in a strong acidification of the soil solution, leaching of base cations and an increased solubility of aluminium, manganese and zinc. Ammonium accumulated in the soil solution, causing severe imbalances between plant nutrients. The effects of these chemical changes in the soil on tree performance are described in a second paper.     

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.     

The effects of sulphur gas and elemental sulphur dust deposition on Pinus contorta x Pinus banksiana: cell walls and water relations

The bulk modulus of elasticity (E) for Pinus contorta (lodgepole pine) x Pinus banksiana (jack pine) hybrids was compared between a site (AI) close to a sour gas processing plant and a control site (AV). The mean bulk modulus of elasticity for branches from AI was 47.5 MPa vs 18.5 MPa for the control site (AV). Site AI had been exposed to S-gas emissions and large amounts of elemental S deposition and had an acidic soil (pH 4.0 at 10 cm depth). During 1981 the needles at AI had more aluminum and iron compared to those at AV (900 ppm vs 390 ppm AI in the 3-year-old needles). Mean leader growth was measured over a 3-year period and was observed to be greater at AI than AV (46+/-7 cm vs 29+/-9 cm for 1988). Histochemically, the needles at AI had higher phenol and lignin content than AV. These results suggest that the S-gas fumigation, S-dust deposition, plus increased concentrations of soluble aluminum and iron had altered the cell wall elastic properties resulting in altered water relations. The implications of this on leaf diffusive resistance and photosynthesis are discussed.     

Effect of fertilisation on the potassium and radiocaesium distribution in tree stands (Pinus sylvestris L.) and peat on a pine mire

This paper compares the effects of single and repeated fertilisation on the contents of potassium, 134Cs and 137Cs in different Scots pine compartments at different levels above ground and in the peat profile 9 years after the Chernobyl disaster. The material was collected from a ditch spacing and fertilisation experiment in Finland. Above a needle potassium concentration of 3.0 mg g(-1) in composed crown samples, 137Cs and 134Cs concentrations remained at about the same level but below that the values were higher on average. This potassium value corresponded to the potassium concentrations of 3.5-3.6 mg g(-1) in the current-year needles of two topmost whorls. The result indicates an enhanced radiocaesium uptake by pine trees under severe potassium deficiency. Fertilisation with potassium-containing fertilisers decreased the caesium uptake considerably. The inhibiting effect of fertilisation on caesium uptake by trees seemed to be fairly long lasting. Fertilisation had sped up the penetration of caesium downwards in the peat profile and its moving out of the active circulation of elements between soil and plants.     

Effects of acidic fog on seedlings of Pinus ponderosa and Abies concolor: foliar injury, physiological and biochemical responses

Seedlings of Pinus ponderosa (ponderosa pine) and Abies concolor (white fir) were exposed to acidic fog (pH 2.0, 3.0 or 4.0) in open-field plots for six weeks. The two species exhibited dissimilar injury responses; neither current year nor previous year needles of ponderosa pine were injured by pH 2.0 fog, but current year needles exhibited higher membrane permeability responses (i.e. needle extract conductivity, K+ concentration). In comparison, both needle age classes in white fir were significantly injured by pH 2.0 fog, but no significant effects on membrane permeability were observed. For both species, whole-study average rates of net photosynthesis in previous year needles were lower in plants exposed to pH 2.0 fog than in plants treated with pH 4.0 fog. While decreased process rates coincided with leaf necrosis in white fir, stomatal closure appeared to be the mechanism of inhibition in ponderosa pine with pH 2.0 fog (i.e. no visible injury). The findings of the present study provide evidence that frequent applications of highly acidic fog (i.e. pH 2.0-3.0) can cause temporal alterations in membrane permeability and gas exchange rates in western conifer seedlings, in the presence or absence of visible injury. However, because incipient effects on other measures of foliage health were species-specific (i.e. concentrations of starch, photosynthetic pigments, inorganic nutrients), a general mechanism of phytotoxicity could not be identified.     

High nitrogen deposition causes changes in amino acid concentrations and protein spectra in needles of the scots pine (Pinus sylvestris)

Growth of the Scots pine (Pinus sylvestris) suffered considerably in forests close to fur farms in western Finland, with the occurrence of winter time dieback in the youngest shoots and leading to a bush-like, flat crown canopy. One reason for this growth disturbance may be a serious imbalance in nitrogen metabolism caused by the extra N supply, emitted as NH3 from the dung of the animals. Total N and NH4+ concentrations in the needles and soluble nitrogen concentration in the soil increased considerably in the vicinity of the fur farms. The extra N in the needles was bound in the first place in arginine, the concentration of which increased 10(2)-10(3) fold compared with control trees, and to a lesser extent in glutamine and other amino acids. Alterations in the quantitative and qualitative protein patterns of the needles were obtained. The extra N increased the concentration of total soluble proteins, although it inhibited the formation of certain polypeptides (particularly in the areas of 30, 38, 50 and 65-90 kDa) which were possibly essential for the normal wintering processes. One reason for the winter time dieback in the high N area could thus be found in the altered protein profiles.