Response of needle sulphur and nitrogen concentrations of Scots pine versus Norway spruce to SO2 and NO2

The results of two field studies and an open-top chamber fumigation experiment showed that the response of mature Scots pine to SO(2) and NO(2) differed from that of mature Norway spruce. Moreover, the response of pine seedlings to SO(2) and NO(2) differed from that of mature trees. The greater increase in the needle total S concentrations of pine suggested more abundant stomatal uptake of SO(2) compared to spruce. Both pine seedlings and mature trees also seemed to absorb more N from atmospheric deposition. Mature pine was able to assimilate SO(4)(2-) derived from SO(2) into organic S more effectively than mature spruce at the high S and N deposition sites, whereas both pine and spruce seedlings accumulated SO(4)-S under NO(2)+SO(2) exposure. Spruce, in turn, accumulated SO(4)-S even when well supplied with N. Net assimilation of SO(4)(2-) in conifer seedlings was enhanced markedly by elevated temperature. To protect the northern coniferous forests against the harmful effects of S and N deposition, it is recommended that the critical level for SO(2) as a growing season mean be set at 5-10 microg m(-3) and NO(2) at 10-15 microg m(-3), depending on the 'effective temperature sum' and/or whether SO(2) and NO(2) occur alone or in combination.     

Effects of acid rain on growth and nutrient concentrations in Scots pine and Norway spruce seedlings grown in a nutrient-rich soil

The effects of artificially applied acid precipitation on growth and nutrient concentrations of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies [L.] Karst.) seedlings were investigated in a long-term acid irrigation experiment in field conditions. Seedlings of northern and southern origin were planted in boxes containing peat and composted soil rich in nutrients, and sprinkler irrigated with water acidified with nitric and sulphuric acids to pH 3 or pH 4 for periods varying from two to three and a half growing seasons during 1986-1989. Water irrigated (pH 5.4-7.6) and non-irrigated groups of seedlings were also included in the experiment. At the end of the experiment needles, main and lateral shoots and roots were collected from the seedlings for the determination of height growth and biomass partitioning, and for the analysis of S, N, Mg, P, K, Ca, Mn and Fe concentrations. The treatment effects compared to the irrigated control were studied using multivariate analyses of variance and covariance. In the pine seedlings the total dry matter production increased by 25-70% compared with the irrigated controls when the total wet deposition to the seedlings exceeded 67 kg S ha(-1) and 36 kg N ha(-1) (e.g. after two growing seasons' exposure of the pH 3 treatment). The increase was mainly due to an increase in needle dry weight (54-72% greater at pH 3) and root weight (20-65% greater at pH 3), whereas the height growth or shoot weight growth were less affected. The northern provenance pine seedlings responded more clearly to the pH 3 irrigation than the southern ones. The treatments had no consistent effects on any of the growth variables studied in the spruce seedlings, however. The pines had higher root and foliage Ca concentrations as a result of the acid irrigation, whereas in spruce, acid rain decreased the Ca concentration in needles and shoots. Root Mn and Fe concentrations were higher in both species as a result of the pH 3 treatment. A higher soil conductivity and Ca concentration resulted from the prolonged pH 3 treatment. The results strongly support the hypothesis that the long-term growth and nutrient allocation response of conifers to acid precipitation is dependent both on the tree species and on the nutritional status of the soil.     

Responses of spruce seedlings (Picea abies) to exhaust gas under laboratory conditions--I. Plant-insect interactions

The effects of motor vehicle exhaust gas on Norway spruce seedlings (Picea abies (L.) Karst) and plant-insect interaction of spruce shoot aphid (Cinara pilicornis Hartig) was studied. The exhaust gas concentrations in the fumigation chambers were monitored and controlled by measuring the concentration of nitrogen oxides (NO(x)) with a computer aided feedback system. The concentrations of major exhaust gas components (black carbon [BC], fine particles, VOCs and carbonyl compounds) in the chamber air were also measured. Responses of Norway spruce seedlings to a 2 and 3-week exhaust gas exposure and subsequent performance of spruce shoot aphid were studied using realistic exposure regimes; 50, 100 and 200 ppb NO(x). The feedback control system based on NO(x) concentrations proved an adequate and practical means for controlling the concentration of exhaust gases and studying plant responses in controlled environment chambers. The exhaust exposure resulted in increased concentrations of proline, glutamine, threonine, aspartic acid, glycine and phenylalanine and decreased concentration of arginine, serine, alanine and glycine in young needles. No changes in soluble N concentrations were observed. The results are interpreted as a stress response rather than use of NO(x) as a nitrogen source. No changes in total phenolics and only transient changes in some individual terpene concentrations were detected. The exhaust gas exposure stressed the exposed seedlings, but had no significant effect on N metabolism or the production of defence chemicals. Aphid performance was not significantly affected. Soluble N, secondary metabolism and aphid performance were not sensitive to exhaust gas exposure during shoot elongation in Norway spruce.     

Norway spruce needles as bioindicator of air pollution in the area of influence of the Sostanj Thermal Power Plant, Slovenia

This paper reports the results of total sulphur content, photosynthetic pigments, ascorbic acid (vitamin C) and alpha-tocopherol (vitamin E) analysed in current-year needles of Norway spruce (Picea abies (L.) Karst.) in the area influenced by sulphur emissions from the Sostanj Thermal Power Plant (STPP), Slovenia, in the period 1991-2004. Ten differently polluted sampling sites in the emission area of STPP were selected. After desulphurization of emission gases from STPP total sulphur content in needles decreased and vitality parameters of needles increased. Moreover, a strong correlation between the average annual emissions of SO(2) from STPP and average annual sulphur content (increase) or average annual chlorophyll content (decrease) in current-year needles was found. The results showed that spruce needles may be an useful bioindicator for detecting changes in the emission rates of SO(2).     

Sulfur dioxide and nitrogen oxides emissions from U.S. pulp and paper mills, 1980-2005

Comprehensive surveys conducted at 5-yr intervals were used to estimate sulfur dioxide (SO,) and nitrogen oxides (NO.) emissions from U.S. pulp and paper mills for 1980, 1985, 1990, 1995, 2000, and 2005. Over the 25-yr period, paper production increased by 50%, whereas total SO, emissions declined by 60% to 340,000 short tons (t) and total NO, emissions decreased approximately 15% to 230,000 t. The downward emission trends resulted from a combination of factors, including reductions in oil and coal use, steadily declining fuel sulfur content, lower pulp and paper production in recent years, increased use of flue gas desulfurization systems on boilers, growing use of combustion modifications and add-on control systems to reduce boiler and gas turbine NO, emissions, and improvements in kraft recovery furnace operations.     

Norway spruce and spruce shoot aphid as indicators of traffic pollution

Two-year-old Norway spruce (Picea abies (L.) Karst) seedlings were exposed to traffic emissions along roadsides with three different traffic densities and speed limits; highway, street and a quiet local road. The responses of the exposed seedlings as a host plant and those of spruce shoot aphid (Cinara pilicornis Hartig) were studied. The concentrations of soluble N and free amino acids, defence chemicals (total phenolics, monoterpenes) were analysed, and aphid growth and reproduction were studied. Along the highway, street and at the local road control site, the atmospheric concentrations of black carbon (BC) and oxides of N (NO(x)) were measured for 1 week during the experiment. The BC data indicate deposition of organic particulate compounds along the highway and street. The NO(x) concentrations along the highway and street showed great diurnal variation, but the average NO(x) concentrations were relatively low. Thus, no changes in N metabolism or growth of the exposed Norway spruce seedlings were found. Along the street, the concentrations of many individual free amino acids, such as proline, as well as total amino acid concentrations, were lower than at the associated control site. Correspondingly, there was also no increase in spruce shoot aphid mean relative growth rate. The aphid reproduction, however, increased along the highway and is suggested to be due to more conducive microclimatic conditions at the exposure site or lack of natural enemies. No changes in defence chemicals (total phenolics, monoterpenes) in relation to the traffic exposure were found. Instead, the microclimatic conditions (temperature, solar irradiation) seemed to affect the concentration of total phenolics.     

Long-term exposure to enhanced UV-B radiation has no significant effects on growth or secondary compounds of outdoor-grown Scots pine and Norway spruce seedlings

The effects of long-term enhanced UV-B radiation on growth and secondary compounds of two conifer species were studied in an outdoor experiment. Scots pine (Pinus sylvestris) seedlings were exposed for two growing seasons and Norway spruce (Picea abies) seedlings for three growing seasons to supplemental UV-B radiation, corresponding to a 30% increase in ambient UV-B radiation. The experiment also included appropriate controls for ambient and increased UV-A radiation. Enhanced UV-B did not affect the growth of the conifer seedlings. In addition, neither the concentrations of terpenes and phenolics in the needles nor the concentrations of terpenes in the wood were affected. However, in the UV-A control treatment the concentrations of diterpenes in the wood of Scots pine decreased significantly compared to the ambient control. Apparently, a small increase in UV-B radiation has no significant effects on the secondary compounds and growth of Scots pine and Norway spruce seedlings.     

Phenolic compounds content in Pinus halepensis Mill. needles: a bioindicator of air pollution

Foliar phenol concentrations (total and simple phenols) were determined in Aleppo pine (Pinus halepensis Mill.) needles collected in June 2000, from 6 sites affected by various forms of atmospheric pollutants (NO, NO(2), NO(x), O(3) and SO(2)) monitored during two months. Results show an increase in total phenol content with exposure to sulphur dioxide and a reduction with exposure to nitrogen oxide pollution. p-Coumaric acid, syringic acid and 4-hydroxybenzoic acid concentrations increase with exposure to nitrogen oxide pollution, whereas gallic acid and vanillin decrease in the presence respectively of sulphur dioxide and ozone. This in situ work confirms the major interest of using total and simple phenolic compounds of P. halepensis as biological indicators of air quality.     

Effects of acid mist on needles from mature Sitka spruce grafts. Part II. Influence of developmental stage, age and needle morphology on visible damage

Mature grafts of five Sitka spruce (Picea sitchensis (Bong.) Carr.) clones were exposed to simulated acid mist comprising an equimolar mixture of H(2)SO(4) and NH(4)NO(3) (1.6 and 0.01 mol m(-3)) at pH 2.5 and 5.0. Mist was applied to potted plants growing in open-top chambers on consecutive days, four times a week, at a precipitation equivalent of 1 mm per day. The total exposure to polluted mist was equivalent to three times that measured at an upland forest in SE Scotland. The aim of the experiment was to characterize the response of juvenile foliage produced by physiologically mature grafts (on seedling root stock) and compare it with the behaviour of juvenile foliage on seedlings. Development of visible foliar damage was followed through the growing season. Measurements of needle length, diameter, weight, surface area, surface was weight and wettability were made on current year needles to determine whether particular foliar characteristics increased susceptibility to injury. Significant amounts (> 10%) of visible needle damage was observed on only one of the five clones. Damage was most severe on the clone with the most horizontal branch and needle habit, but over the five clones there was no relationship between angle of branch display and damage. Likewise no combination of needle characteristics (length, width, area, amount of wax) was indicative of potential susceptibility. A comparison with previous acid misting experiments using seedlings suggests that juvenile foliage on physiologically mature trees is equally susceptible to visible injury as juvenile seedling foliage. Data of budburst differed among clones, and in this experiment exerted the over-riding influence on development of injury symptoms. Foliage exposed to a combination of strong acidity and high sulphate concentrations over the few weeks immediately following budburst suffered most visible injury. The absence of significant amounts of visible damage in UK forests probably reflects the general low susceptibility to visible injury of Sitka spruce exposed to acid mist.     

Monoterpenes emitted to air from industrial barking of Scandinavian conifers

Monoterpenes released to air were determined during single-log barking of timber at a saw-mill and during tumble drum barking of pulpwood at a TMP mill. Characteristic proportions of fourteen compounds were assessed by adsorbent sampling and gas chromatography. For Scots pine (Pinus sylvestris), the bicylic monoterpenes alpha-pinene and 3-carene predominated. For Norway spruce (Picea abies), alpha-pinene followed by beta-pinene were the major components. The proportions of beta-phellandrene and myrcene from spruce were remarkably high from barking compared to wood processing. Environmental aspects on the emissions are discussed.     

Sulphur dioxide adsorption in Scots pine canopies exposed to high ammonia emissions near a Cu-Ni smelter in SW Finland

Since 1994 the nickel-processing plant at the Cu-Ni smelter at Harjavalta, south-west Finland, has emitted considerable amounts of NH(3) into the atmosphere. The effects of NH(3) emissions on nitrogen and sulphur deposition in throughfall and the foliar nutrient status were investigated in a Scots pine stand at 0.5 km distance. Bulk deposition, stand throughfall and percolation water (20 cm depth) samples were collected at 4-week intervals during 1992-1998. pH and the Ca, Mg, K, NH(4) and SO(4) concentrations were determined on the samples. NH(3) emissions have strongly increased the scavenging of SO(2) from the air in the pine stand, and the increased levels of N and S deposition were clearly evident as increased foliar N and S concentrations and larger needle size. The increased input of SO(4) into the forest floor was not associated with an increase in the leaching of Ca and Mg from the surface soil layers.     

Performance of some growth variables

European beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Silver fir (Abies alba Mill.) were exposed to low concentrations of ozone (O(3)) and sulfur dioxide (SO(2)), alone and combined, and simulated acid rain (pH 4.0) in sheltered open-top chambers in Hohenheim (Southwest Germany) for almost five years. The concentrations of O(3) and SO(2) used were related to annual ambient average found in southern West Germany. Two control chambers were ventilated with charcoal filtered air and rainfall was simulated at pH 4.0 and 5.0. Because of large dense plant growth in the chambers it was only possible to measure uncompleted growth of shoots in the upper canopy. Therefore, growth analysis was restricted to this area. The treatment with acidic precipitation decreased the annual shoot growth of beech and reduced leaf surface area of those trees. Exposure to SO(2), O(3) alone and in combination resulted in further reduction of shoot length and leaf surface area. Fumigation with SO(2) and O(3) + SO(2) caused insignificant decreases of shoot length, total dry weight and needle surface area of spruce. The lateral leader shoot growth of spruce exposed to O(3) was significantly reduced only in the last year of the experiment. Growth rates of the spruce exposed to charcoal filtered air and non-acidic precipitation were reduced more than those of beech and fir. Growth variables determined for fir reflected different rates of incremental change. Exposure to O(3) resulted in the largest dry matter production of all fir groups but those exposed to charcoal filtered air and non-acidic precipitation responded with the best lateral leader shoot growth, lowest specific leaf area (SLA) and leaf area ratio (LAR) respectively indicating best metabolic efficiency. At the conclusion of this study a classification of sensitivity was developed for the tree species.     

Effects of acid mist on mature grafts of Sitka spruce. Part I. Frost hardiness and foliar nutrient concentrations

Mature grafts of five clones of Sitka spruce (Picea sitchensis Bong. Sarg.) were exposed to simulated acid mist composed of an equimolar mixture of sulphuric acid and ammonium nitrate at pH 2.5 and pH 5.0 in open-top chambers from May to November 1991. Treatments were applied on consecutive days, four times a week. The pH 2.5 treatment provided an overall dose three times higher than that received by forests in upland areas of Britain. Frost hardiness was assessed in November by freezing detached current year shoots at a range of temperatures and assessing the rate of electrolyte leakage Foliar nutrient concentrations were determined on the same shoots. Acid mist at pH 2.5 significantly reduced frost hardiness in four of the five clones; the temperature causing 50% shoot death (LT50) was increased by 0 to 7 degrees C. The clones varied in their level of hardiness, one clone being exceptionally frost sensitive. The frost hardiness of the frost sensitive clone was found to be less perturbed by acid mist than the hardiness of the more frost resistant clones. Mature grafts showed a smaller reduction in hardiness at an equivalent dose than that found previously with Sitka spruce seedlings. Compared with seedlings, grafts had lower absolute concentrations of foliar sulphur. Exposure to acid mist at pH 2.5 increased %S in current year foliage by <0.05% compared with absolute increases of more than 0.10% in current year foliage of seedlings. We conclude that the effect of acid mist on frost hardiness is likely to be less on mature trees than on seedlings and that the increased frost risk to mature trees of Sitka spruce from occult deposition alone is small.     

The effect of papermill wastewater and organic amendments of sodium accumulation by potted cottonwoods

In this experiment, the impacts of pulp mill effluent irrigation, Fraser cottonwood (Populus deltoides 'Fraser') seedlings, and pulp sludge and manure soil amendments on sodium accumulation and distribution in the soil profile were evaluated during a 6 month greenhouse study. Sludge soil amendments and wastewater irrigation did not reduce stem biomass production of the cottonwood. Increased stem biomass production associated with manure soil amendments resulted in greater total uptake of sodium into stem material. This uptake was 0.002% of wastewater sodium inputs. In containers with seedlings, sodium concentrations were less in the surface horizon and more in the lower horizons than in containers with no seedlings. Infiltration rates and total sodium accumulation in the soil profile were not affected by the presence of Fraser cottonwood or the application of sludge or manure amendments to soil.     

Germination, growth and gas exchange of selected boreal forest seedlings in soil containing oil sands tailings

Greenhouse experiments were conducted to determine the effects of soil enriched in fine tailings (FT), produced by the oil sands extraction, on germination, seedling growth and physiology of several plant species of the boreal forest. The germination of seeds was initially delayed by 15% FT in dogwood (Cornus stolonifera Michx) and jack pine (Pinus banksiana Lamb) but not in white spruce [Picea glauca (Moench) Voss]. In the second set of experiments we showed that all dogwood seedlings survived 6 months of treatment with 15% FT while the survival rates of raspberry, jack pine and white spruce seedlings were reduced to 44, 55 and 94%, respectively. FT reduced root and shoot dry weights in raspberry seedlings and the number of lateral shoots in jack pine and white spruce seedlings. In raspberry and jack pine seedlings, reductions of gas exchange were recorded. The results of our study suggest that the modifications of soil chemistry, texture and structure by FT may all contribute to the observed phytotoxic effects.     

Ozone exposure-response relationships for biomass and root/shoot ratio of beech (Fagus sylvatica), ash (Fraxinus excelsior), Norway spruce (Picea abies) and Scots pine (Pinus sylvestris)

Current-year seedlings of beech, ash, Norway spruce and Scots pine were exposed during one growing season to different, but moderate, ozone (O(3)) scenarios representative for Switzerland (50, 85, 100% ambient, 50% ambient+30 nl l(-1)) in open-top chambers (OTCs) and to ambient O(3) concentrations in the field. Biomass significantly decreased with increasing O(3) dose in all species except for spruce. Losses of 25.5% (ash), 17.4% (beech), 9.9% (Scots pine) were found per 10 microl l(-1) h accumulated O(3) exposure over a threshold concentration of 40 nl l(-1) during daylight hours (AOT40). Ratios of root/shoot biomass (RSR) also significantly decreased with increasing AOT40 levels in beech and ash, but not in Norway spruce and Scots pine. The data show that the deciduous species beech and ash were more susceptible to O(3) with respect to RSR and biomass than the coniferous species Norway spruce and Scots pine.     

A laboratory investigation of the routes of HNO3 dry deposition to coniferous seedlings

White pine, Norway spruce and red spruce seedlings were exposed to nitric acid vapor concentrations of 10 to 120 ppb in constant stirred tank reactors. Nitric acid dry deposition rates were determined from both the change in nitric acid concentration in the reactor flow stream and from the amount of nitrogen recovered from the seedlings. Nitric acid labeled with 15N was used to distinguish dry-deposited nitrogen in the plant from the nitrogen that was already present. It was found that dry deposition occurs via three routes: surface deposition, trans-cuticular deposition, and stomatal deposition. Resistance to surface deposition is very low (< 4.8 m2-s mol(-1)) for a freshly washed surface, but increases as the surface adsorption sites are occupied. Resistance to trans-cuticular uptake averaged 206 m2-s mol(-1). Stomatal resistance can be calculated from the rate of water diffusion out of the plant. Eighty per cent of the nitric acid deposited via the trans-cuticular and stomatal routes was assimilated by the plant. However, none of the nitric acid deposited on the surface was assimilated. In rural areas with coniferous forests, the combination of low ambient nitric acid concentrations and low initial surface resistance means that most nitric acid will be dry deposited on the tree surface, and thus will not be directly assimilated.     

Container-grown tree seedling responses to sodium chloride applications in different substrates

Seedlings of Norway maple (Acer platanoides), silver birch (Betula pendula), Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) were grown in selected sodium chloride (NaCl) concentrations, soil types and under different watering regimes. Plants were raised from seeds, except for Scots pine plants which were obtained from a commercial source. Among the plant species tested, Scots pine was the most tolerant to soil salinity, while Norway spruce was the most susceptible. For both Norway maple and Norway spruce some half-sib families were more tolerant than others. No significant correlation was found between the tolerance of different half-sib families and the tolerance of mother trees observed in the field. The extent of leaf necrosis correlated significantly with the leaf concentrations of sodium (Na) and chloride (Cl). Among half-sib families within the species no such correlation was found. On the other hand, the least injured progeny of Norway maples had the highest concentrations of NaCl. The extent of salt-induced leaf necrosis varied with soil type, and a significant interaction between species and soil type was observed. Seedlings of Norway spruce grown in sand showed more severe necrosis and significantly higher concentrations of Na and Cl than seedlings grown in loam, silt loam, and peat. The severity of salt-induced leaf injury varied with the watering regime. Silver birch was the most affected species by drought and autumn watering treatments. Plants of silver birch subjected to drought showed increased leaf necrosis compared to the non-treated plants, and autumn watering treatment reduced the severity of leaf necrosis.     

Organism-induced accumulation of iron, zinc and arsenic in wetland soils

Four year old spruce (Picea abies (L.) Karst.) seedlings were planted in sand pots and supplied with nutrient solution. Three groups were formed, differing only in manganese nutrition (0.5 ppm, 2.5 ppm, 12.5 ppm, respectively). After three months, five individuals of each group were transferred to a dew chamber. For the next seven weeks the trees were sprayed in the evenings, the relative humidity overnight was kept high and the droplets were collected directly from the needles in the mornings. The trees were sprayed with HNO3 (pH 3.4) during the first three weeks to reduce the natural buffering capacity of the needles. After this time, the trees were sprayed with KCl (1 mM) solution, and NaHSO3 was added to the chamber resulting in SO2 concentrations usually between 50 and 150 microg m(-3). Needles and water samples were analysed. Foliar Ca seemed to be only a short-time buffer even under optimal Ca supply. A highly significant influence of managanese supply on manganese in needles and droplets was observed, as well as on sulphate, H+ and calcium concentrations in the droplets. The SO2 flux to trees treated with 12.5 ppm Mn was about twice as high as to trees treated with 0.5 ppm Mn. The conclusion is that this is due to a synergism between manganese leaching and catalysis of the SO2 oxidation by the leached Mn2+ ions. The results suggest a positive feedback between (moderate) acidification of soils and SO2 and NH3 inputs to terrestrial ecosystems.