Organochlorine pollutants in Scots pine needles--biological and site related variation within a forest stand

To study the biological variation in the content of some persistent organic pollutants, viz hexachlorobenzene (HCB), alpha- and gamma-hexachlorocyclohexane (alpha- and gamma-HCH), and 1,1,1-trichloro-2,2-bis-(4-chlorophenyl)-ethane (DDT) were analysed in needles from Scots pine trees growing at an isolated peninsula south of Stockholm, Sweden. The concentration variations of each compound was evaluated by a nested analysis of variance (ANOVA) with sampling site specific compass direction, individual tree, and sampling height as factors. Two pair-wise post hoc tests were used to test significant results from the ANOVA. The hexachlorocyclohexanes showed no significant differences between sampling site, trees or sampling height. DDT concentration was significantly lower at the sampling site with the densest vegetation compared to at least three of the other sampling sites. HCB was significantly lower at the lowest sampling height (0.5 m).     

Growth variation of Scots pine across a pollution gradient on the Kola Peninsula, Russia

Decadal exposure to emissions from a non-ferrous smelter has damaged the forest ecosystems surrounding the city of Monchegorsk located on the Kola Peninsula in northwestern Russia. We use the methods of tree-ring analysis to study the areal extent and timing of recent growth reductions of Scots pine (Pinus sylvestris L.) in the region surrounding the smelter in Monchegorsk. Reduced growth of Scots pine was observed up to 30 km southwest of the smelter. This directional gradient of forest damage is related to the dispersal of pollutants which is influenced by the prevailing northern winds and local topographic features. Old Scots pines (age 200 years +) appeared to be more sensitive than younger ones: growth reductions of old trees had started earlier and reductions were observed farther from the smelter than for younger trees. The findings are compared to a classification which describes the state of forest ecosystems based on the occurrence of certain plant species; the classification matched well with the observed growth trends. Pollution-induced changes in the climatic signal in tree-rings are also studied. The strong dependence of growth on mid-summer temperatures, typical for Scots pine on high latitudes, proved to be insensitive to effects of pollution. Changes in the climate-growth relationship took place decades after growth trends had started to decline.     

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.     

The effects of multiple metal contamination on ectomycorrhizal Scots pine (Pinus sylvestris) seedlings

Experiments were conducted to investigate the effects of single and multiple metal contamination (Cd, Pb, Zn, Sb, Cu) on Scots pine seedlings colonised by ectomycorrhizal (ECM) fungi from natural soil inoculum. Seedlings were grown in either contaminated field soil from the site of a chemical accident, soils amended with five metals contaminating the site, or in soil from an uncontaminated control site. Although contaminated and metal-amended soil significantly inhibited root and shoot growth of the Scots pine seedlings, total root tip density was not affected. Of the five metals tested in amended soils, Cd was the most toxic to ECM Scots pine. Field-contaminated soil had a toxic effect on ECM fungi associated with Scots pine seedlings and caused shifts in ECM species composition on ECM seedlings. When compared to soils amended with only one metal, soils amended with a combination of all five metals tested had lower relative toxicity and less accumulation of Pb, Zn and Sb into seedlings. This would indicate that the toxicity of multiple metal contamination cannot be predicted from the individual toxicity of the metals investigated.     

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.     

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.     

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.     

High ambient concentrations of monoterpenes in a Scandinavian pine forest

Total monoterpene levels of the order of 100 μg m ⁻³ were assessed in the summer night air of a typical young planted forest of Scots pine (Pinus silvestris) in Sweden. These concentrations are much higher than those reported in previous studies of ambient monoterpenes. Predominant species were α-pinene and 3-carene. Camphene, β-pinene, myrcene, β-phellandrene and limonene were other prominent components determined. The monoterpenes were shown to originate almost exclusively from Scots pine and the proportions between them to be characteristic of this conifer species.     

Seasonal patterns of ascorbate in the needles of Scots Pine (Pinus sylvestris L.) trees: correlation analyses with atmospheric O3 and NO2 gas mixing ratios and meteorological parameters

In the present field study the role of ascorbate in scavenging the harmful atmospheric trace gases O3 and NO2 was examined. For this purpose ascorbate contents were determined in needles of adult Scots pine trees (Pinus sylvestris L.) during three consecutive years. Ascorbate contents were correlated with ambient tropospheric O3 and NO2 concentrations and with meteorological parameters. The results showed a strong correlation of atmospheric O3 but not of atmospheric NO2 concentrations with the apoplastic content of ascorbate during the seasonal course. Ascorbate contents in needle extracts did not correlate with ambient trace gas concentrations. In the apoplastic space, but not in needle extracts ascorbate contents correlate highly significantly with global radiation. From these results it is assumed that apoplastic ascorbate in Scots pine needles is adapted to the actual atmospheric O3 concentration to mediate immediate detoxification of O3, while the atmospheric O3 concentration itself is largely dependent on light intensity.     

Damage to Eastern White Pine by Sulfur Dioxide,Semimature-Tissue Needle Blight,and Ozone

Eastern white pine (Pinus strobus L.) forests were severely damaged by atmospheric sulfur dioxide up to distances of 25 miles northeast of large smelters located in the Sudbury mining district of Ontario. Damage to white pine was measured in terms of foliage, bark, and biological injuries, radial and volume growth decrement, and tree mortality. The foliar symptoms of sulfur dioxide injury on white pine trees sometimes resembles those caused by a physiogenic disease, semimature-tissue needle blight (abbreviated to SNB). Studies on the nature and etiology of SNB were conducted in white pine forests in the Upper Ottawa Valley, which are remote from smelter operations which might pollute the atmosphere. These studies included the determination of the role that naturallyoccurring atmospheric ozone plays in the occurrence of SNB. Differences between the symptoms of sulfur dioxide injury, SNB, and ozone damage are outlined.     

Relations between Scots pine needle element concentrations and decreased needle longevity along pollution gradients

Scots pine (Pinus sylvestris L.) shoots were sampled along transects near one urban pollution source and two smelters. Needle Mg, P and K concentrations decreased from the second to the fourth age class linearly with needle survival along the urban pollution gradient. Still, over 80% of the average concentration of these nutrients remained in the fourth needle age class. Decreased needle longevity was closely related to the increased heavy metal concentrations near the smelters. Near the urban pollution source, it was related to the increased annual needle mass and the increased needle nutrient concentrations. Decreased Mn accumulation along with needle age was detected near all pollution sources. Leaching of Mn from needles and especially from soil as a cause of decreased needle concentrations is 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.     

Contribution of ambient ozone to Scots pine defoliation and reduced growth in the Central European forests: a Lithuanian case study

The study aimed to explore if changes in crown defoliation and stem growth of Scots pines (Pinus sylvestris L.) could be related to changes in ambient ozone (O₃) concentration in central Europe. To meet this objective the study was performed in 3 Lithuanian national parks, close to the ICP integrated monitoring stations from which data on meteorology and pollution were provided. Contribution of peak O₃ concentrations to the integrated impact of acidifying compounds and meteorological parameters on pine stem growth was found to be more significant than its contribution to the integrated impact of acidifying compounds and meteorological parameters on pine defoliation. Findings of the study provide statistical evidence that peak concentrations of ambient O₃ can have a negative impact on pine tree crown defoliation and stem growth reduction under field conditions in central and northeastern Europe where the AOT40 values for forests are commonly below their phytotoxic levels.     

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.     

Effects of ozone on three open-pollinated families of Pinus taeda L. grown in two substrates

Seedlings from three open-pollinated loblolly pine (Pinus taeda L.) families grown in a mixture of commercial peat moss and grade 3 vermiculite (1:3 by volume) or a mixture of mineral soil and peat (1:1 by volume) were exposed to 0, 160 or 320 ppb ozone (O3) for 6h/day, 4 days/week for 8 weeks beginning 12 weeks after transplanting. Before exposures began, seedlings grown in the vermiculite-peat substrate were taller but smaller in diameter than those grown in the mineral soil-peat substrate. After 8 weeks of exposure, seedlings grown in the mineral soil-peat substrate were significantly larger in diameter and total biomass than those grown in the vermiculite-peat substrate. Primary needle and secondary needle injury increased with increasing O3 concentrations. Suppression of diameter growth, shoot weight and root weight was linear as O3 concentration increased. The effect of O3 on height or diameter growth or shoot biomass was not influenced by substrate type; but the suppression of root biomass due to O3 was dependent on substrate, with greater suppression in biomass occurring in the vermiculite-peat substrate. Foliar injury due to O3 was slightly greater in family 8-103, but growth suppression due to O3 was not significantly different among the families. Based on root biomass, response of seedlings to O3 was substrate-dependent.     

Scots pine needle surfaces on radial transects across the north boreal area of Finnish Lapland and the Kola Peninsula of Russia

To gain an understanding of the characteristics of the needle surfaces of naturally regenerated Scots pine (Pinus sylvestris L.) and their geographical distribution, eight physicochemical variables were investigated within the north boreal forest area. The visibly undamaged needles were collected in autumn 1990 from 114 plots (3-5 pines per plot) along radial transects from the Monchegorsk and Nikel smelters, emitting SO2 and heavy metals, on the Kola Peninsula, Russia, to Finnish Lapland. The needles were examined by scanning electron microscopy (SEM) and measured for surface wettability using the droplet contact angle (DCA) method. Significant geographical south-north and west-east patterns could be observed in needle surface characteristics, correlating with emissions from the smelters and to climate. Despite the slightly higher initial DCAs (61.4-87.6 degrees) towards the north, the pine needle surfaces of the northernmost transects exhibited higher annual needle wettability change (ANWC) and wax erosion rate (AWER), expressed as the reduction in DCAs and epistomatal wax tube distribution (WTD) during one year, respectively, but a lower occurrence of particles and fungal hyphae than those of more southern transects. The higher ANWC was related to higher atmospheric SO2 concentration, and to the lower long-term temperature sum, but not clearly to annual precipitation. In the Monchegorsk smelter area, the current needles exhibited, on average, a 15% higher WTD and seven degree larger DCA, resulting in more hydrophobic needle surfaces than in Finnish Lapland, but during their first year, both the AWER and ANWC, were greatly increased. In Finnish Lapland, 30%, of all the epistomatal wax tubes disappeared from the needle surface during the first year, the value being 70% for the pines located 8 km from the smelter. The mineral composition of the particles deposited on the needle surfaces mirrored that of the minerals being produced by the smelters (e.g. FexSx, CuFeS2, NixSx, FexOx). Stomatal densities were in the range of 71.1-141.7 stomata mm(-2). The lower densities of stomata on needles close to the smelters correlated with reduced number of needle age classes of the pines, higher dry weights, higher pollutant accumulation and lower Mn and Zn concentrations in the needles. The AWER and ANWC were able to indicate the most seriously deteriorated needle surfaces on a regional scale, e.g. including the surroundings of both smelters, although the plot-specific relationships between 'needle surface variables' and 'pollution variables' were generally weak. The present study showed that the exposure of pine needles to the combined effects of ambient pollutants and harsh climate led to a deterioration in the physicochemical characteristics of the epicuticular wax, which may have serious ecophysiological consequences in the long term.     

Seedling insensitivity to ozone for three conifer species native to Great Smoky Mountains National Park

Field symptoms typical of ozone injury have been observed on several conifer species in Great Smoky Mountains National Park, and tropospheric ozone levels in the Park can be high, suggesting that ozone may be causing growth impairment of these plants. The objective of this research was to test the ozone sensitivity of selected conifer species under controlled exposure conditions. Seedlings of three species of conifers, Table Mountain pine (Pinus pungens), Virginia pine (Pinus virginiana), and eastern hemlock (Tsuga canadensis), were exposed to various levels of ozone in open-top chambers for one to three seasons in Great Smoky Mountains National Park in Tennessee, USA. A combination of episodic profiles (1988) and modified ambient exposure regimes (1989-92) were used. Episodic profiles simulated an average 7-day period from a monitoring station in the Park. Treatments used in 1988 were: charcoal-filtered (CF), 1.0x ambient, 2.0x ambient, and ambient air-no chamber (AA). In 1989 a 1.5x ambient treatment was added, and in 1990, additional chambers were made available, allowing a 0.5x ambient treatment to be added. Height, diameter, and foliar injury were measured most years. Exposures were 3 years for Table Mountain pine (1988-90), 3 years for hemlock (1989-91), and 1 and 2 years for three different sets of Virginia pine (1990, 1990-91, and 1992). There were no significant (p<0.05) effects of ozone on any biomass fraction for any of the species, except for older needles in Table Mountain and Virginia pine, which decreased with ozone exposure. There were also no changes in biomass allocation patterns among species due to ozone exposure, except for Virginia pine in 1990, which showed an increase in the root:shoot ratio. There was foliar injury (chlorotic mottling) in the higher two treatments (1.0x and 2.0x for Table Mountain and 2.0x for Virginia pine), but high plant-to-plant variability obscured formal statistical significance in many cases. We conclude, at least for growth in the short-term, that seedlings of these three conifer species are insensitive to ambient and elevated levels of ozone, and that current levels of ozone in the Park are probably having minimal impacts on these particular species.     

Growth, respiration and nitrogen content in needles of Scots pine exposed to elevated ozone and carbon dioxide in the field

Single Scots pine (Pinus sylvestris L.) trees, aged 30 years, were grown in open-top chambers and exposed to two atmospheric concentrations of ozone (O3; ambient and elevation) and carbon dioxide (CO2) as single variables or in combination for 3 years (1994-1996). Needle growth, respiration and nitrogen content were measured simultaneously over the period of needle expansion. Compared to ambient treatment (33 nmol mol(-1) O3 and 350 micromol mol(-1) CO2) doubled ambient O3 (69 nmol mol(-1)) significantly reduced the specific growth rates (SGRs) of the needles in the early stage of needle expansion and needle nitrogen concentration (N1) in the late stage, but increased apparent respiration rates (ARRs) in the late stage. Doubled ambient CO2 (about 650 micromol mol(-1)) significantly increased maximum SGR but reduced ARR and N1 in the late stage of needle expansion. The changes in ARR induced by the different treatments may be associated with treatment-induced changes in needle growth, metabolic activities and turnover of nitrogenous compounds. When ARR was partitioned into its two functional components, growth and maintenance respiration, the results showed that neither doubled ambient O3 nor doubled ambient CO2 influenced the growth respiration coefficients (Rg). However, doubled ambient O3 significantly increased the maintenance respiration coefficients (Rm) regardless of the needle development stage, while doubled ambient CO2 significantly reduced Rm only in the late stage of needle expansion. The increase in Rm under doubled ambient O3 conditions appeared to be related to an increase in metabolic activities, whereas the decrease in Rm under doubled ambient CO2 conditions may be attributed to the reduced N1 and turnover rate of nitrogenous compounds per unit. The combination of elevated O3 and CO2 had very similar effects on growth, respiration and N1 to doubled ambient O3 alone, but the interactive mechanism of the two gases is still not clear.     

Ozone-induced changes in the chloroplast structure of conifer needles, and their use in ozone diagnostics

Ozone induces characteristic symptoms in the chloroplasts of the needles of several coniferous species. Chloroplasts are (1) reduced in size and (2) the stroma is electron dense. Moreover, (3) these chloroplast alterations are more pronounced in the outer mesophyll cell layers and in the upper side of the needle compared to the inner layers and lower side. The syndrome, including the three symptoms (1)-(3), is found in the green needles of Scots pine and Norway spruce not only in the experimental fumigations, but also in mature trees in the field, and has potential for diagnosis of ozone stress. For sound ozone diagnostics all three symptoms must be present in the samples studied. The symptoms in relation to needle anatomy and physiology is discussed, and recommendations for sampling and analysis are given.     

Long-Term Monitoring of Scots Pine Litter Decomposition Rates Throughout Sweden Indicates Formation of a More Recalcitrant Litter in the South

Decomposition studies were carried out at sites throughout Sweden, including the four Integrated Monitoring sites. Scots pine needle litterbag weight loss measurements over 3 or 5 years were determined at 26 sites and repeated up to 27 times, depending on the site. Humus layer respiration rates were determined for 20 sites in 1987–1989 and repeated in 2007–2008. Partial Least Squares (PLS) regression was used to elucidate the relative importance of climatic and soil factors. Annual needle weight losses decreased only slowly (20–10%) over 3–5 years for all northern (>60°N) sites but decreased sharply from 30 to 10% in the third year in southern (<60°N) sites. Respiration rates of southern sites were less (40% on average) than those of northern sites. Humus layer N was positively correlated to needle weight loss during the first and the second years, but negatively correlated in the third year and to respiration rates. The results indicated that litter formed in southern Sweden became more recalcitrant in later stages of decomposition compared to litter produced in northern Sweden.