The effects of excess nitrogen deposition on young Norway spruce trees. Part II the vegetation

The effects of wet-deposited nitrogen on soil acidification and the health of Norway spruce were investigated in a pot experiment using an open-air spray/drip system. Nitrogen was applied as ammonium ((NH(4))(2)SO(4)) or nitrate (HNO(3)/NaNO(3)) in simulated rain to either the soil or the foliage. Symptoms of forest decline as observed in the field were not reproduced, and there was no evidence of direct toxicity. Treatments did, however, have significant effects on tree nutrition. Both NH(+)(4) and NO(-)(3) treatment applied to the foliage lowered foliar K concentrations. NH(+)(4) to a greater extent. Soil-applied NH(+)(4) reduced foliar Mg concentrations and increased foliar Al and Fe. Soil-applied NO(-)(3) significantly reduced foliar P concentrations, and at high doses prevented the alleviation of P deficiency by fertiliser. These effects could be important in some field situations. Ammonium deposition is predicted to be more damaging than nitrate deposition, although the latter may be critical for forests where P status is marginal, such as in parts of the British uplands.     

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.     

Physiology of young Norway spruce

In the Hohenheim experiment young spruce (Picea abies L. Karst.) were exposed to low levels of SO(2) and/or O(3) and acid precipitation. At the end of a five-year experimental period (1983-88) the following physiological parameters were examined: water soluble thiols, ascorbic acid, glutathionereductase activity and pigment content. Exposure to SO(2), leads to an increase in thiol content, to a slight decrease of ascorbic acid and to a pronounced decrease of pigments. O(3) exposure increases the content of ascorbic acid and decreases the thiols and the glutathione-reductase activity with no change in the pigment content. The combined exposure to SO(2), and O(3) results in the most distinct deviations compared to the control chamber response. These needles show the highest increase of ascorbic acid and thiols, the dry weight is decreased as is the glutathione-reductase activity and the pigment content is reduced. Consequences of these physiological alterations for the plant's health are discussed.     

Influence of soil substrate and ozone plus acid mist on the pigment content and composition of needles from young Norway spruce trees

The influence of an ozone + acid mist treatment on photosynthetic pigments has been examined with the needles of the (Picea abies) clones 11, 14, 16, and 133 by spectrophotometric analysis of the total pigment extract and of single components upon HPLC separation (Part A), and in terms of a detailed pigment analysis of the 1987 and 1986 needles of clone 14 by TLC (Part B). Clone 14 had been already analysed prior to the onset of the experiment. At the end of the 14-month experiment, which incorporated frost events during a simulated winter period, neither symptoms corresponding to those of spruce Type I or IV decline, nor those of ozone damage could be observed. However, the 1986 needles of the trees on soil 1, which exhibit an adequate nutrient content, showed zonal chlorosis independently of the ozone + acid mist treatment. Analysis of variance of chlorophyll contents and needle ages showed a clear reduction to nearly 50% in the 1986 needles of clone 11, soil 1, and clone 16, soil 2. In contrast, clones 14 and 16 (soil 1) formed significantly more chlorophylls during the shorter exposure time in the 1987 flush. The detailed analysis of the individual pigment components of clone 14 needles provided no evidence for a destructive influence of the treatment on the chlorophylls, xanthophylls and beta-carotene in the two needle generations which had developed during the experiment, in spite of the distinct K deficiency of the 1986 needles of the trees on soil 2 and the common chlorosis of the needles of the trees on soil 1. The observed increase in violaxanthin content upon O(3)-treatment observed in clone 14 can be considered as an expression of the protective function of the xanthophylls against photooxidative processes. In conclusion, the observed differences in the chlorophyll and carotenoid contents are better correlated with the individual clone and soil character than with the ozone + acid mist treatment. Comparing the results of the pigment analyses of the needles the differences in the pigment concentrations reflect the N and K contents (Pfirrmann et al., 1990), which differ significantly between the clones. Thus it is not possible to pool the pigment data of all clones without considering the different nutrient levels.     

Trichloroacetic acid in Norway spruce/soil-system. I. Biodegradation in soil

Trichloroacetic acid (TCA) as a phytotoxic substance affects health status of coniferous trees. It is known as a secondary air pollutant (formed by photooxidation of tetrachloroethene and 1,1,1-trichloroethane) and as a product of chlorination of humic substances in soil. Its break-down in soil, however, influences considerably the TCA level, i.e. the extent of TCA uptake by spruce roots. In connection with our investigations of TCA effects on Norway spruce, microbial processes in soil were studied using 14C-labeling. It was shown that TCA degradation in soil is a fast process depending on TCA concentration, soil properties, humidity and temperature. As a result, the TCA level in soil is determined by a steady state between uptake from the atmosphere, formation in soil, leaching and degradation. The process of TCA degradation in soil thus participates significantly in the chlorine cycle in forest ecosystems.     

Influence of soil substrate and ozone plus acid mist on the frost resistance of young Norway spruce

The needles of clonal Norway spruce grown in environmental chambers on two different soils (an acidic soil 1 and a calcareous soil 2) and exposed to two levels of ozone fumigation (a low level combined with neutral mist = control, and an elevated one combined with acid mist = treatment) were analyzed for their frost hardiness. No effect of ozone was observed on either the development of frost resistance during the hardening phase or on the decrease in frost resistance during the dehardening phase. The preliminary results of Brown et al. (1987) and Barnes and Davidson (1988), which indicated that ozone treatment predisposes plants to winter injury, could thus not be confirmed. Frost resistance was, however, distinctly influenced by the content of the mineral nutrients of the soils. The pronounced K(+) deficiency of the needles of the trees growing on the neutral soil (Alps) had less effect on the development of frost resistance than did the Ca(++) and Mg(++) deficiency of the needles of the trees grown on the acidic soil 1 (Bavarian Forest). The variability of frost resistance between the different clones on soil 1 was partly attributed to fluctuations in the mineral nutrient content of the needles, rather than to a genetic predisposition.     

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.     

Foliage responses of spruce trees to long-term low-grade sulfur dioxide deposition

Foliage on spruce trees (Picea rubens Sarg.) growing on dry SO(2) deposition zones (dry SO(2) deposition ranging from 0.5 and 8.5 S kg ha(-1) year(-1)) downwind from a SO(2) emission source was analyzed to assess chronic effects of long-term low-grade SO(2) deposition on net photosynthesis, stomatal conductance, dark respiration, stomatal antechamber wax structures, elemental concentrations in and on foliage (bulk and surficial concentrations), and types of epiphytic fungi that reside in the phylloplane. Elemental distributions on stomatal antechambers, on fungal colonies, and on smooth surfaces between stomates and fungus colonies were determined with a scanning electronic microscope (SEM) by way of X-ray scanning. It was found that net photosynthesis of newly developed spruce foliage (current-year, and 1-year-old) was not significantly affected by the local SO(2) deposition rates. Sulfur dioxide deposition, however, may have contributed to the gradual decrease in net photosynthesis with increasing needle age. Dark respiration rates were significantly higher on foliage taken from high SO(2) deposition zones. Stomatal rod-web structures deteriorated to flakes with increasing needle age and increasing SO(2) deposition. Further inspection of the needle surfaces revealed an increasing abundance of fungal colonies with increasing needle age. Many fungal taxa were isolated and identified. It was found that black yeasts responded positively, and Xylohypha pinicola responded negatively to high rates of SO(2) deposition. Surficial concentrations of elements such as P, S, K, Cl, Ca were about 10 times higher on fungal colonies than on smooth needle surfaces. Surficial Ca contents on 4 or 5-year-old needles decreased with increasing SO(2) deposition, but surficial S concentrations remained the same. In contrast, bulk foliar Ca and S concentrations increased with increasing SO(2) deposition.     

Response of mycorrhizal Norway spruce seedlings to various nitrogen loads and sources

Norway spruce seedlings were grown under greenhouse conditions in Rootrainers with a vermiculite-peat moss mixture under various N-regimes for 6 months. Either ammonium or nitrate was applied in loads of 100 or 800 kg N ha(-1) year(-1) to seedlings which were either non-mycorrhizal or inoculated with the mycorrhizal fungi Hebeloma crustuliniforme or Laccaria bicolor. The use of increasing N loads enhanced shoot and total biomass, whereas root/shoot ratio, number of short roots and mycorrhization decreased. A significant enhancement of the concentration and content was obvious for the element N, whereas a significant decrease was obvious for P and Zn concentrations. The use of ammonium, as opposed to nitrate, significantly enhanced the biomass and the numbers of short roots, and reduced the root/shoot ratios, but did not influence the mycorrhization. It further significantly enhanced the N concentrations in roots and shoots. Fungal inoculation with H. crustuliniforme or L. bicolor compared to non-inoculated controls significantly enhanced shoot and total biomass, but reduced root/shoot ratios. The mycorrhization further significantly enhanced N and P concentrations and contents, but reduced Mn. Overall, the mycorrhization improved the P nutrition of the seedlings independently on the applied N loads or N sources. Dose response curves using ammonium nitrate as N source with a maximum load of 1600 kg N ha(-1) year(-1) applied on seedlings associated with H. crustuliniforme revealed that the maximum growth was reached at a load of 800 kg N ha(-1) year(-1) with a simultaneous decrease of the mycorrhization. In both shoots and roots, N concentrations increased constantly with increasing N loads, while P, Ca, and Zn concentrations decreased constantly.     

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.     

The effects of SO2 fumigation on the nitrogen metabolism of aseptically grown spruce seedlings

Aseptically grown spruce seedlings were cultivated in a hydroponic system, where the roots were separated from the shoots by a gastight, silicone material. The plants were fumigated with four SO(2) concentrations (93, 190, 270 and 530 microg m(-3)) for nine weeks. Up to 270 microg m(-3) of SO(2), an inhibition of nitrogen metabolism (enzyme activities of nitrate reductase (NR) and glutamine sythetase (GS) and nitrate content) in the shoot was compensated by a stimulation in the root, while nitrogen uptake was unaffected. Only the treatment with 530 microg m(-3) of SO(3) decreased enzyme activities, nitrate content in both roots and shoots as well as nitrate uptake, and inhibited the growth of plants. Increases in the content of thiols and superoxidismutase activity are discussed in terms of SO(2) detoxification.     

Interactive effects of nitrogen deposition and fire on plant and soil chemistry in an alpine heathland

The response of alpine heathland vegetation and soil chemistry to N additions of 0, 10, 20 and 50 kg N ha⁻¹ year⁻¹ in combination with simulated accidental fire (+/−) was monitored over a 5-year period. N addition caused rapid and significant increases in plant tissue N content and N:P and N:K of Calluna vulgaris, suggesting increasing phosphorus and potassium limitation of growth. Soil C:N declined significantly with N addition, indicating N saturation and increasing likelihood of N leakage. Fire further decreased soil C:N and reduced potential for sequestration of additional N. This study shows that alpine heathlands, which occupy the headwaters of many rivers, have limited potential to retain deposited N and may rapidly become N saturated, leaking N into downstream communities and surface waters.     

Persistent effects of ozone on needle water loss and wettability in Norway spruce

Four-year-old, seed-grown trees of Norway spruce (Picea abies (L.) Karst.) were exposed in open-top chambers to charcoal-filtered air (8 h daily mean 54 microg O(3) m(-3)) over three consecutive summers (1986-1988). In mid-May 1988, before the third season of fumigation and more than 7 months after exposure to ozone the previous summer had terminated, daily rates of transpiration from intact shoots and water loss from excised needles were measured together with the amount of wax on the needle surface. In mid-July, 92 days after the beginning of the third year of exposure, the wettability of needles was assessed by measuring the contact angle of water droplets on the surface of needles. Exposure to 156 microg O(3) m(-3) resulted in a 16% increase in daily transpiration in current year's needles and a 28% increase in 1-year old needles. These effects were associated with slower stomatal closure in response to increasing water deficit in the needles previously exposed to 156 microg m(-3) ozone. The long-lasting nature of such ozone-induced effects could predispose trees to drought and winter desiccation. No significant effects of ozone were found on the amount of wax covering the needle surface, but a marked increase in the wettability of needles exposed to ozone was observed. The far reaching physiological consequences of these effects in the field and the possibility that similar disturbances may contribute to the decline of high-altitude forests of Norway spruce in Europe are discussed.     

Effects of simultaneous ozone exposure and nitrogen loads on carbohydrate concentrations, biomass, and growth of young spruce trees (Picea abies)

Spruce saplings were grown under different nitrogen fertilization regimes in eight chamberless fumigation systems, which were fumigated with either charcoal-filtered (F) or ambient air (O3). After the third growing season trees were harvested for biomass and non-structural carbohydrate analysis. Nitrogen had an overall positive effect on the investigated plant parameters, resulting in increased shoot elongation, biomass production, fine root soluble carbohydrate concentrations, and also slightly increased starch concentrations of stems and roots. Only needle starch concentrations and fine root sugar alcohol concentrations were decreased. Ozone fumigation resulted in needle discolorations and affected most parameters negatively, including decreased shoot elongation and decreased starch concentrations in roots, stems, and needles. In fine roots, however, soluble carbohydrate concentrations remained unaffected or increased by ozone fumigation. The only significant interaction was an antagonistic effect on root starch concentrations, where higher nitrogen levels alleviated the negative impact of ozone.     

Rapporteurs' report on discussions at the workshop on excess nitrogen deposition, Leatherhead, September 1987

This paper reports major points from the discussions which took place during the Workshop on Excess Nitrogen Deposition.     

Ecosystem effects of atmospheric deposition of nitrogen in The Netherlands

Atmospheric deposition of inorganic N, mainly ammonium volatilized from manure produced in intensive stockbreeding, on sensitive terrestrial and aquatic ecosystems in The Netherlands is in the order of 40 to 80 kg ha(-1) year(-1). Proven effects of this deposition are (i) eutrophication with N, leading to floristic changes (ii) acidification of base-poor sandy soils and of moorland pools, leading to higher concentrations of dissolved, potentially toxic metals such as Al3+, and (iii) increased levels of nitrate in groundwater below woodlands. In acid forest soils, but not in soils under heathland, nitrification and leaching of nitrate is common. However, in very poor sandy forest soils and at very high ammonium inputs, nitrification may be too slow to prevent the development of high concentrations of ammonium. Both excessive acidification and excessive levels of ammonium probably play an important role in the general forest decline, which is most severe in the southern and central parts of the country, where ammonium inputs are highest.     

The effects of atmospheric nitrogen deposition and soil chemistry on the nutritional status of Pseudotsuga menziesii, Pinus nigra and Pinus sylvestris

The ammonium content and the base cation content, expressed relative to ammonium, are enhanced in the soil of Dutch forests, due to the extremely high deposition of ammonium to the forest floor. A nation-wide investigation was carried out to establish whether and how these changed nitrogen fluxes in deposition and soil affect the nutritional status of coniferous trees. The chemical composition of needles of Douglas fir, Scots pine and Corsican pine showed a regional trend similar to that of deposition and soil solution. Particularly nutrients, expressed relative to nitrogen, decreased from North to South. Of the macronutrients phosphorus was most often deficient and therefore probably limiting in the Douglas fir stands. Many pine trees suffered from relative magnesium shortages. In all stands, magnesium and, in Douglas stands, also phosphorus contents of the needles were negatively correlated with ammonium and ammonium/cation ratios in deposition. However, in contrast to pine trees, nutrient contents in needles of Douglas fir showed correlation with nitrate rather than with ammonium in the soil solution. Correlation analyses indicate that nitrogen fluxes in the soil, indirectly affect the nutritional status of coniferous trees.     

Uptake,translocation and fate of trichloroacetic acid in a Norway spruce/soil system

Trichloroacetic acid (TCA) is a secondary atmospheric pollutant formed by photooxidation of chlorinated solvents in the troposphere--it has, however, recently been ranked among natural organohalogens. Its herbicidal properties might be one of the factors adversely affecting forest health. TCA accumulates rapidly in conifer needles and influences the detoxification capacity in the trees. The aim of the investigations--a survey of which is briefly given here--was to elucidate the uptake, distribution and fate of TCA in Norway spruce. For this purpose young nursery-grown plants of Norway spruce (Picea abies (L.) Karst.) were exposed to [1,2-14C]TCA and the fate of the compound was followed in needles, wood, roots, soil and air with appropriate radio-indicator methods. As shown by radioactivity monitoring, the uptake of TCA from soil by roots proceeded most rapidly into current needles at the beginning of the TCA treatment and was redistributed at later dates so that TCA content in older needles increased. The only product of TCA metabolism/biodegradation found in the plant/soil-system was CO(2) (and corresponding assimilates). TCA biodegradation in soil depends on TCA concentration, soil humidity and other factors.     

Selection effects of air pollution on gene pools of Norway spruce, European silver fir and European beech.

The effects of industrial pollution on allelic and genotypic structures of Norway spruce. European silver fir and European beech were investigated by means of isozyme analysis. In a mixed Norway spruce-silver fir forest stand in an area heavily polluted by sulphur dioxide and heavy metals in the region of Spis (eastern Slovakia), pairs of neighbouring damaged and apparently healthy trees were selected in two replicates (44 and 69 pairs in a heavily and moderately damaged stand, respectively). Pairwise sampling of trees with contrasting vitality was applied to reduce potential effects of site heterogeneity on the vitality of sampled trees. No significant differences in allelic and genotypic frequencies were found between sets of healthy and declining trees. There were differences in the single-locus heterozygosities, but these were not consistent between the replicates. However, the set of damaged trees exhibited higher levels of genetic multiplicity and diversity, possibly due to the deleterious effect of rare alleles under the conditions of air pollution. Consequently. following the decline of pollutant-sensitive trees, the remaining stand will be depleted of a part of alleles with unknown adaptive value to future selection pressures.     

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.