Growth responses of Scots pine to climatic factors on reclaimed oil shale mined land

Afforestation on reclaimed mining areas has high ecological and economic importance. However, ecosystems established on post-mining substrate can become vulnerable due to climate variability. We used tree-ring data and dendrochronological techniques to study the relationship between climate variables and annual growth of Scots pine (Pinus sylvestris L.) growing on reclaimed open cast oil shale mining areas in Northeast Estonia. Chronologies for trees of different age classes (50, 40, 30) were developed. Pearson’s correlation analysis between radial growth indices and monthly climate variables revealed that precipitation in June–July and higher mean temperatures in spring season enhanced radial growth of pine plantations, while higher than average temperatures in summer months inhibited wood production. Sensitivity of radial increment to climatic factors on post-mining soils was not homogenous among the studied populations. Older trees growing on more developed soils were more sensitive to precipitation deficit in summer, while growth indices of two other stand groups (young and middle-aged) were highly correlated to temperature. High mean temperatures in August were negatively related to annual wood production in all trees, while trees in the youngest stands benefited from warmer temperatures in January. As a response to thinning, mean annual basal area increment increased up to 50 %. By managing tree competition in the closed-canopy stands, through the thinning activities, tree sensitivity and response to climate could be manipulated.     

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.     

Adaptation strategies and referencing trial of Scots and black pine populations subjected to heavy metal pollution

The impact of industrial heavy metal pollution on Scots pine (Pinus sylvestris L.) and black pine (Pinus nigra Arn.) populations was investigated. Sampled pine stands, which were located in Upper Silesia (southern Poland) in an area strongly polluted by heavy metals, consisted of resistant and sensitive trees. To evaluate the adaptation process, genetic structure and diversity was tested using isozyme analysis. Higher levels of Zn, Pb, Cd and Cu were detected in needles of sensitive trees compared with resistant ones. With respect to morphology, Scots pines were more distinctly impaired than black pines. Although black pines had lower heavy metal concentrations, levels in 1-year-old needles, other than Cu, significantly exceeded “reference plant” values (Markert 1994). In both species, resistant trees demonstrated a lower degree of genetic variation than metal-sensitive trees with respect to some enzyme loci (SHDH A, PGI, PGM, MDH C and DIA). This observation was corroborated in sensitive trees by the smaller number of identified alleles and alleles per locus, absence of private alleles and significant excess of homozygotes in relation to expected Hardy–Weinberg equilibrium values. Assuming that only resistant trees of both species survive under conditions of prolonged soil contamination, the observed genetic structure implies that remaining populations will be depleted of some alleles of unknown adaptive value to future selection pressures. Genetic changes induced by heavy metals suggest an important role for specific enzymes—FEST, SHDH A and B, GOT B and PGI—in the adaptation process. Our results may serve as a basis for selection and propagation of individuals appropriate for re-cultivation of areas chemically degraded by industrial activity.     

Epiphyte recolonization of oaks along a gradient of air pollution in South-East England, 1979-1990

The abundances of epiphytes on free-standing oaks (Quercus robur) along a transect extending approximately 70 km SSW from Central London were redetermined annually from 1979 to 1990. During this period SO(2) levels in inner London fell to between one-quarter and one-seventh of levels occurring in the 1960s. The sample trees were also scored annually using Hawksworth and Rose's (1970) scale for predicting SO(2) levels from the epiphytes present. Little evidence was obtained for epiphyte recolinization of oaks at sites within the built-up area. At Epsom Common, on the urban fringe, cover of the pollution-tolerant lichen Lecanora conizaeoides fell, and in the inner suburbs, at Putney Heath, L. conizaeoides and Lepraria incana increased, while the alga Desmococcus viridis declined at this site. These changes might be due to a contraction inwards of zones of peak abundance of L. conizaeoides and D. viridis with decreasing SO(2) levels. However, it is also possible that the changes at Putney Heath resulted from scrub clearance activities and alteration to the microclimate. The cover of the moderately SO(2) tolerant foliose lichen Hypogymnia physodes remained low and static at Putney Heath during the 12-year period. Possible reasons for the poor recolinization of London's oaks by epiphytes are discussed. Particular emphasis is given to the highly acidic bark of oak in the London area (pH2.9-4.0) in comparison with other tree species and with oaks in unpolluted regions.     

Measurement of ethylene emission from Japanese red pine (Pinus densiflora) under field conditions in NOx-polluted areas

Emission of ethylene from the needles of Japanese red pine, Pinus densiflora, was measured in air-polluted areas in Hiroshima, Japan. We applied a suitable protocol to determine the rate of ethylene emission from the excised needles. The influence of excision of needles on ethylene emission was not detected during the first 4 h of incubation at 20 degrees C. Ethylene emissions were low in the unpolluted (clean) areas regardless of the altitude or season. The emission of stress ethylene increased with the atmospheric NO2 concentration, suggesting that atmospheric NOx or related substances induced the higher ethylene emission in the polluted areas (near urban and industrial areas). In all cases, 1-year-old needles emitted significantly larger amounts of ethylene than the current needles. Ethylene emission did not increase evenly in the polluted areas, but the frequency of trees emitting high ethylene increased. Therefore, threshold rates for the baseline ethylene emission were proposed.     

Use of tree rings to investigate the onset of contamination of a shallow aquifer by chlorinated hydrocarbons

Oaks (Quercus velutina Lam.) growing over a shallow aquifer contaminated by chlorinated hydrocarbons were studied to determine if it was possible to estimate the approximate year that contamination began. The annual rings of some trees downgradient from the contaminant release site contained elevated concentrations of chloride possibly derived from dechlorination of contaminants. Additionally, a radial-growth decline began in these trees at approximately the same time that chloride became elevated. Growth did not decline in trees that contained smaller concentrations of chloride. The source of elevated chloride and the corresponding reductions in tree growth could not be explained by factors other than contamination. On the basis of tree-ring evidence alone, the release occurred in the late 1960s or early 1970s. Contaminant release at a second location apparently occurred in the mid- to late 1970s, suggesting that the area was used for disposal for at least 5 years and possibly longer.     

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.     

Reduction of stem growth and site dependency of leaf injury in Massachusetts black cherries exhibiting ozone symptoms

Leaf ozone symptoms in natural ecosystems are increasingly reported but ozone effects on tree growth and the mediation of site conditions are still little documented. This study tests two hypotheses: (1) leaf injury in black cherry is associated with decline in radial growth, (2) symptoms are more prevalent on mesic sites. On sites supporting black cherry across Massachusetts, tree growth and leaf ozone injury were surveyed in 1996 using a randomized plot network established in the 1960s. Forty-seven percent of 120 trees sampled for ozone symptoms were symptomatic with generally low levels of injury. Over a 31-year period symptomatic trees had 28% lower stem growth rates than asymptomatic trees. Ozone symptom expression was enhanced in well growing stands on moister, cooler and more elevated sites. Ozone appeared to increase environmental stress and had a more pronounced effect on growth in better growing black cherry stands. This complicates management decisions as thinning increases growth and moisture availability.     

Foliar sulphur species in pine: a new indicator of a forest ecosystem under air pollution stress

An eleven-year foliar sulphur (S) monitoring program was carried out from 1976 to 1986 near a sulphur recovery-gas plant in west-central Alberta, Canada, as part of a case study designed to determine the effects of chronic, low concentration sulphur gas emissions on the forest ecosystem surrounding the gas plant. Measurements of both foliar total sulphur (ST) and foliar inorganic sulphur (SO4-S) concentration in lodgepole x jack pine trees at the end of each of the 11 growing seasons were taken to provide an indication of S loading of the forest from industrial sulphur emissions. To measure the state of the forest ecosystem, foliar ST was separated into foliar accumulated sulphur (inorganic sulphur or SO4-S) and foliar assimilated sulphur (organic sulphur or S0) and the ratio of SO4-S/S0 taken. Foliar S0 was calculated as the difference between foliar ST and foliar SO4-S. The median SO4-S/S0 ratio, with all three years of needles considered, varied from 0.29 at a reference location (AV) to 0.88 at the location with the highest stress (AI). The corresponding mean values ranged from 0.3 at the reference location to 2.2 at the location of highest stress. The mean seasonal photosynthetic rate of current year's foliage of the pine trees and soil pH were reduced at a stressed location (AI) compared to the reference location (AV), between 1976 and 1981. Over this same time period the mean foliar SO4-S/S0 ratio increased from 0.4 +/- 0.1 to 1.0 +/- 0.3 at the stressed location (AI) and remained nearly the same at the reference location (AV) at 0.3 +/- 0.1. This research suggests that the foliar SO4-S/S0 ratio is a useful indicator of the state of forest ecosystems under S air pollution stress. It is concluded that foliar S separated into various fractions has potential as an early warning environmental management tool.     

Effect of nitrogenous air pollutants on changes in protein spectra with the onset of winter in the leaves and shoots of the bilberry (Vaccinium myrtillus L.)

Electrophoretic profiles of 0.1 m Tris-HCl (pH 9.0)-soluble proteins in the leaves and stems of the bilberry (Vaccinium myrtillus L.) were investigated in the latter part of the growing season (July-October), in a highly-polluted industrial area with serious forest damage (main emissions NO(2), SO(2), NH(3), urea and fertilizer dust) in Oulu, northern Finland, and in an unpolluted control forest with the same climatic and edaphic conditions. About ten protein components in the leaves and seven in the stems increased in concentration in SDS-PAGE towards the autumn, while one to four other components decreased in concentration. The pollutants in the industrial area caused a considerable decrease in the concentrations of soluble proteins, particularly in the leaves in August and September. A decrease was obtained in six components in the leaves and three components in the stems, while increases were observed in one component in the leaves (10 kDa) and four components in the stems (10, 43, 50 and 65 kDa). One reason for the lowered cold-resistance in the bilberry caused by air pollutants could thus be the lowered concentrations of certain soluble cytoplasmic proteins that protect the plant against winter conditions.     

Conifer aphids in an air-polluted environment. II. Host plant quality

Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were transported to five forest sites at increasing distances from a pulp mill emitting mainly SO(2). Levels of compounds which may have nutritional or defensive value for aphids on pine and spruce seedlings were studied. Glucose and fructose concentrations were significantly increased in pine and spruce needles near the pulp mill. There were no changes in sucrose and starch concentrations. In pine shoots, total free amino acid concentration and the concentrations of ornithine, lysine, histidine and arginine were significantly negatively correlated with the distance from the pulp mill, while in spruce only the individual amino acids glycine, ornithine, lysine and histidine showed a significant negative correlation with distance. There were no changes in total phenolic, catechin, total monoterpene and total resin acid concentrations. However, in pine seedlings monoterpenes beta-pinene and sabinene and in spruce seedlings resin acid palustric acid were significantly correlated with the distance from the pulp mill. The results indicate that SO(2) disturbs carbohydrate metabolism in spruce and pine seedlings. The elevated concentrations of arginine may be the result of the combinations of SO(2), NO(3) and NH(3) emissions of the pulp mill. The emissions did not have any impact on total amounts of defensive substances in trees. Thus, the possible susceptibility of conifers to herbivores appears to be due to changes in nutritive value rather than to reduced chemical resistance.     

Height growth of different European Scots pine Pinus sylvestris L. Provenances in a heavily polluted and a control environment

Results are presented of height measurements and degree of needle injury on five-year-old plants of Scots pine (Pinus sylvestris L.) growing near a phosphate fertiliser plant that emits SO(2) and fluorides. The populations of Scots pine represented in this experiment originate from 11 countries and were substantially differentiated in height growth and extent of needle necroses. Those populations which grew most rapidly were found to be the most sensitive to pollutant injury. The least productive provenances from the north of the range (Sweden, USSR) are at the same time characterized by lowest decline in height growth, lowest mortality and least extensive necroses. It is proposed that gene banks be established for the best genotypes likely to be eliminated in the heavily polluted conditions of Poland today.     

An Evaluation of the Effects of Ozone Injury on Radial Growth of Ponderosa Pine (Pinus ponderosa) in the Southern Sierra Nevada

Growth of ponderosa pines with visible symptoms of ozone injury was compared with that of asymptomatic trees in the southern Sierra Nevada, California. Time series analysis indicated that there was no significant reduction in annual radial increment of symptomatic trees during recent years compared to past growth and growth of asymptomatic trees. First order autocorrelation and climatic variables accounted for a large proportion of the variance in growth index, and winter precipitation was positively correlated with growth for all size and age classes. Although ozone concentrations are high enough to cause chlorosis and premature needle senescence in ponderosa pine, there has been no significant change in growth associated with ozone injury.     

Fluoranthene fumigation and exogenous scavenging of reactive oxygen intermediates (ROI) in evergreen Japanese red pine seedlings (Pinus densiflora Sieb. et. Zucc.)

Generation of reactive oxygen intermediates (ROI) such as O(2)(-), H(2)O(2), and *OH is known to be a major mechanism of damage in biological systems. This study investigated and compared effectiveness of scavenging ROI generated in fluoranthene (FLU) pre-fumigated Japanese red pine seedlings. Three kinds of eco-physiological assessments were used to express the impact of the different fumigants used inside the green house. Gas exchange measurements showed negative changes induced by 10 microM FLU on Japanese pine seedlings during a 10 d exposure period whilst no negative change was found during a 5 d exposure period. Moreover, during a 14 d FLU exposure incorporating ROI scavengers, results revealed that chlorophyll fluorescence, needle chemical contents and needle dry mass per unit area of the seedlings were affected. The negative effects of FLU on the conifer were dependent on both the dose and period of FLU fumigation. Peroxidase (PERO), superoxide dismutase (SOD) and mannitol (MANN) were all effective scavengers of ROI. MANN scavenged *OH, the most lethal of the ROI. For practicable use, MANN is more economical, and may be the best ROI scavenger among the three considered. It can be concluded that efficient scavenging of ROI in biological systems is important to mitigate the negative effects of FLU on Japanese red pine trees.     

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.     

Assessment of airborne heavy metal pollution by aboveground plant parts

Italian stone pine (Pinus pinea L.) and oleander (Nerium oleander L.) leaves, bark and wood samples were collected at different sites around an industrial area (Huelva, SW Spain) and compared with samples of the same species from a background site. Samples were analysed with respect to the following pollutants: Al, Ba, Cr, Cu, Fe and Pb by ICP-AES. The suitability of different plant parts as biomonitors of pollution was investigated. In pine samples from the polluted sites the ratio of concentrations between bark and wood was high for Al, Ba, Cu and Fe, whereas no differences were found in samples from the unpolluted area. No differences were detected in oleander for the same ratio. In the oleander species, the ratio between leaves and wood concentration allowed to distinguish between control and polluted sites. The ratio of the concentration between leaves and wood was elevated for Al, Ba and Fe in pine samples from the polluted sites. The ratio of the concentration in bark or leaves to their concentration in wood might be useful to detect inorganic atmospheric pollutants.     

Quantifying ozone uptake at the canopy level of spruce,pine and larch trees at the alpine timberline: an approach based on sap flow measurement

Micro-climatic and ambient ozone data were combined with measurements of sap flow through tree trunks in order to estimate whole-tree ozone uptake of adult Norway spruce (Picea abies), cembran pine (Pinus cembra), and European larch (Larix decidua) trees. Sap flow was monitored by means of the heat balance approach in two trees of each species during the growing season of 1998. In trees making up the stand canopy, the ozone uptake by evergreen foliages was significantly higher than by deciduous ones, when scaled to the ground area. However, if expressed per unit of whole-tree foliage area, ozone flux through the stomata into the needle mesophyll was 1.09, 1.18 and 1.40 nmol m(-2) s(-1) in Picea abies, Pinus cembra and Larix decidua, respectively. These fluxes are consistent with findings from measurements of needle gas exchange, published from the same species at the study site. It is concluded that the sap flow-based approach offers an inexpensive, spatially and temporally integrating way for estimating ozone uptake at the whole-tree and stand level, intrinsicly covering the effect of boundary layers on ozone flux.     

Growth variation of white oak subjected to historic levels of fluctuating air pollution

Growth of white oak (Quercus alba L.) trees was examined, using tree-ring analysis, at three sites near a small, remote coal-fired power plant in central Pennsylvania, USA. Forests immediately adjacent to the power plant have been subjected to power plant emissions since the power plant initiated operation in 1954. However, localized, ground-level fumigations have been gradually reduced over the years due to a series of construction projects resulting in increased stack heights. Comparisons of growth were made among the white oaks growing at the three close-in sites, as well to the growth of white oak at three control sites located 10-50 km from the power plant, during periods of differing stack heights. White oak exhibited reduced growth at two of the close-in sites during the time period when historical ground-level air pollution exposures were assumed to be greatest due to low stack heights. White oak growth at the third close-in site was not substantially reduced during this time period. In 1976, taller stacks were implemented at the power plant to reduce local, ground-level concentrations of air pollutants. The recovery of tree growth at the two close-in affected sites, and increased synchronous growth responses from 1976-85 among all three close-in sites, indicates that implementation of taller stacks in 1976 reduced ground-level pollutant levels to such dosages that growth was not impaired at any site. Also, growth rates after 1976 were comparable to the growth rates of the white oaks growing on the control sites. A possible interacting factor was a severe drought that occurred in the mid-1960s in central Pennsylvania.     

Isozyme markers associated with O(3) tolerance indicate shift in genetic structure of ponderosa and Jeffrey pine in Sequoia National Park, California

Effects of canopy ozone (O(3)) exposure and signatures of genetic structure using isozyme markers associated with O(3) tolerance were analyzed in approximately 20-, approximately 80-, and >200-yr-old ponderosa (Pinus ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. & Balf.) in Sequoia National Park, California. For both species, the number of alleles and genotypes per loci was higher in parental trees relative to saplings. In ponderosa pine, the heterozygosity value increased, and the fixation index indicated reduction of homozygosity with increasing tree age class. The opposite tendencies were observed for Jeffrey pine. Utilizing canopy attributes known to be responsive to O(3) exposure, ponderosa pine was more symptomatic than Jeffrey pine, and saplings were more symptomatic than old growth trees. We suggest that these trends are related to differing sensitivity of the two species to O(3) exposure, and to higher O(3) exposures and drought stress that younger trees may have experienced during germination and establishment.     

Reproduction of mountain birch along a strong pollution gradient near Monchegorsk, Northwestern Russia

We explored effects of severe pollution on sexual reproduction of mountain birch, Betula pubescens subsp. czerepanovii, by counting catkins on sample branches and weighing both somatic and generative structures of both short and long shoots from birches growing at 21 sites around large nickel-copper smelter at Monchegorsk, Northwestern Russia. Proportion of reproducing trees, production of catkins, shoot and catkin weight, as well as the relative difference in weight of somatic structures of generative and vegetative shoots, were generally independent of pollution load; in 2003 birches growing in industrial barrens produced more catkins than birches growing in unpolluted forests. Thus, we found no support for the hypothesis that reproductive allocation should decrease with decrease in environmental capacity. Absence of adverse effects might indicate that long-lasting pollution impact already eliminated the most sensitive individuals from the affected birch populations.