The effect of industrial pollution on copper,lead, zinc and cadmium concentration in xylem rings of resistant (Populus marilandica) and sensitive (P. balsamifera) species of poplar

Summary The patterns of radial distribution of copper, lead, zinc and cadmium in two species of poplar, growing under polluted conditions near a copper smelter, were investigated. Populus marilandica is very resistant to such pollution, but P. balsamifera is sensitive. Soils were examined for concentrations of these metals, and only a surface layer of soil was polluted with copper and lead. P. balsamifera absorbed much more cadmium than P. marilandica under the same conditions. P. marilandica shows a pattern of copper radial distribution with significantly increased concentration of the meral in the layers in the middle of the trunk compared to P. balsamifera. Zinc and cadmium have a pattern with increased metal concentration from the bark towards the pith of the tree. In the case of copper and lead the pattern indicates increased concentrations of these metals in the rings adjacent to the pith while the pattern in other parts of the tree remains unconfirmed due to poor precision. Poplar xylem seems to accumulate zinc and cadmium while the content of lead in xylem is much lower than in the adjacent soil.This paper is dedicated to the memory of Professor Stefan Bialobok, Institute of Dendrology, Polish Academy of Sciences in Kornik     

The ion concentration of dew condensed on Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) needles

Summary Dew droplets collected with pipettes from coniferous needles were analysed for their ionic composition. Almost all samples of dew taken from Scots pine trees (Pinus sylvestris) showed significantly higher ion concentrations than those taken from Norway spruce trees (Picea abies). This can be explained by the micromorphology of the needle surface. The higher microscale roughness of the wax layer of a pine needle causes a more efficient flux of atmospheric aerosol particles compared to the spruce needle surface. Dew on coniferous needles is shown to be capable of maintaining pH values below 3 for several hours.     

Dendrochronological research of Scots pine (Pinus sylvestris L.) radial growth in vicinity of industrial pollution

The aim of this research is to investigate changes in the annual radial increment of Scots pine (Pinus sylvestris L.) in the vicinity of intensive (3–10 km) and moderate (11–20 km) industrial pollution during different growth periods (growth promotion, inhibition, and recovery). Low level of emission was beneficial for tree growth during the growth promotion period, and the annual radial increment in the zones of intensive and moderate pollution increased by approximately 15–25% and 10%, respectively. Severe loss was reported to forests during the growth inhibition period when nitrogen and sulphur dioxide emissions were 37–40 thousand (thou.) tons per year. About 40–45% tree radial increment loss was observed in the stands closest to the pollution source, and 15–20% loss was observed for the most distant stands. The stabilization of radial growth decrease and the beginning of recovery of damaged stands began in 1988–1992, when the annual amount of industrial emissions and environmental pollution were considerably reduced. The stabilization of radial growth and the initiation of recovery after pollution reduction were high for the most damaged stands. Their radial increment was stable and close to that of the control stands in 2000–2011. Stands with less damage growing further from the pollution source were recovered earlier, and their radial increment stabilized near the control increment in 1995–1999. The results of linear regression analysis demonstrated that the impact of pollution is different for stands growing at different distances from the plant, and the impact decreases with distance (R² = 0.78 and R² = 0.75, respectively; p < 0.05).     

Topographic influences on radial growth of Scots pine (Pinus sylvestris L.) at small spatial scales

Dendroecological and numerical methods were used to study the influence of topographic position on radial growth of Scots pine (Pinus sylvestris) stands exposed to soil dryness. The correlation structure of total tree-ring width and latewood width of eight scattered populations representing various topographic habitats (steep south-facing slopes, plateaus and hollows) within a rock-slide area (750 m a.s.l.) of about 1 km² was investigated by principal component analysis. Scatter plots of component loadings indicated that (i) total ring width and latewood width are influenced by various climatic factors, (ii) stands growing at similar topographic position show a high agreement in year-to-year variability of radial growth, and (iii) distinct effects of topographic features (slope aspect, slope magnitude) on tree growth are modified by local disturbances (erosion, grazing) and the age structure of stands. Furthermore, both the time series of component scores and non-metric multidimensional scaling of chronologies indicated years where extremely limiting or favorable climate conditions prevailed throughout the study area (pointer years). The influence of climate on tree growth in various topographic habitats was mediated through the influence of climatically stressful years. Because stands are located at sites with different levels of water stress, growth differences between chronologies are considered to be caused by site-specific susceptibility of tree growth to soil dryness. Significant correlations between precipitation in April to June and ring-width confirm that water availability is the primary growth-limiting factor within the study area. These small-scale variations in growth-climate relationships have significant implications for dendroclimatological studies. So paleoclimatic reconstructions based on tree rings will have to assure that an unbiased data set is used, which compensates for local growth-variabilities due to site related environmental stresses.     

Long-term implications of industrial pollution stress on lipids composition in Scots pine (Pinus sylvestris L.) roots

The influence of pollution stress (SO₂, Cu²⁺, Pb²⁺, Zn²⁺ and fluoride) on composition changes in cellular membranes of roots of three European of Scots pine (Pinus sylvestris L.) populations were examined. Plant material growing in three experimental areas: Kórnik relatively free of air pollution (control), Luboń: SO₂ and HF and Głogów: SO₂ and heavy metals. Analysis of total phospholipids and their composition indicates that the phosphatidylcholine (PC) and phosphatidylethanolamine (PE) were present in lower concentration in the root tissues from both polluted sites (Luboń and Głogów). The difference in PC:PE ratio between control and both polluted sites was greater in root of Scots pine population from Russia than in the population from Slovakia. Under pollution conditions the content of lipid soluble antioxidant α-tocopherol was lower about 220 %, in comparison to the control. The action of pollution stress also lead to lowering of unsaturated:saturated ratio of total fatty acid, and lower content of polyunsaturated fatty acids, linoleic acid (18:2) and eicosatrienoic acid (20:3). We concluded that the long-term pollution stress markedly inhibited lipid biosynthesis in root tissue of Scots pine and it is probably contribute to the reduction of productivity of forests. These results also suggest that lipid composition can be used as an indicator of changes in tissue roots of Scots pine caused by air and/or soil long-term pollution.     

Daily dynamics in xylem cell radial growth of Scots pine (Pinus sylvestris L.)

Daily dynamics of radial cell expansion during wood formation within the stems of 25-year-old Scots pine trees (Pinus sylvestris L.), growing in field conditions, were studied. The samples of forming wood layers were extracted 4 times per day for 3 days. Possible variations in the growth on different sides of the stem, duration of cell development in radial cell expansion phase and dynamics of cell growth in this phase were taken into account. The perimeters of tracheid cross-sections as a reflection of primary cell wall growth were the criterion of growth in a radial direction. For the evaluation of growing cell perimeters a special system for digital processing and image analysis of tracheid cross-sections of the forming wood was used. Growth rate for certain time intervals was estimated by the change in the relation of the perimeter of each observed cell in each of ten tracheid rows in each of 12 trees to the perimeter of the xylem cell of the same row before the expansion. Temporal differences in average values of the relations were estimated by Analyses of Variance. The existence of daily dynamics of Scots pine xylem cell radial growth has been proved. Intensive growth of pine tracheids has been shown to occur at any time of the day and to depend on the temperature regime of the day and the night as well as water supply of stem tissues. Moreover, reliable differences (P = 0.95) in the increment of cell walls during tracheid radial expansion have been found. Pulsing changes of the water potentials both of the cell and the apoplast, as the reason for the fluctuations of radial cell growth rate, were discussed.     

Concentration gradients of free sterols,steryl esters and lipid phosphorus in the trunkwood of Scot's pine (Pinus sylvestris L.)

The amounts of free sterols, steryl esters and lipid phosphorus were determined in the sapwood and heartwood of mature, and in the outer and inner sapwood of young Pinus sylvestris trees. In the mature trees (up to 70 years old) the heartwood contains significantly higher amounts of free sterols than the sapwood. No radial gradient can be demonstrated in the amounts of steryl esters. Lipids extracted from the sapwood contain higher amounts of phosphorus than those from the heartwood. Stems of young Pinus sylvestris trees (up to 13 years old) show in the inner sapwood higher amounts of both free sterols and steryl esters than the peripheral younger wood zone. The inner sapwood of the young stems shows slightly higher amounts of lipid phosphorus than the outer sapwood. The results indicate that Pinus sylvestris accumulates both free sterols and steryl esters in the stems at a very early stage of the life cycle. Sterol accumulation in the innermost parts of the stems seems not to depend on heartwood formation.     

Effect of disturbed photoperiod on the surface structures of ripening Scots pine (Pinus sylvestris L.) seeds

 The role of disturbed photoperiod on the developing surface layers of ripening Scots pine (Pinus sylvestris L.) seeds was studied from August 14 to October 30. The embryo sizes of both the light-treated and the control seeds were examined by the X-ray method prior to germination tests. The anatomical details of resin-embedded seeds were examined by fluorescence and light microscopy. The timing of the ripening process of the surface structures was described and documented. The greatest anatomical changes in the ripening seeds occurred in the sarcotesta and in the nucellar layers. Maturation of the surface structures was essentially slower than could have been interpreted by the size of the embryo and responded clearly to the disturbance of photoperiod. Accumulation of phenolic substances and degeneration of cells, particularly at the chalazal region, advanced faster in the light-treated than in the control seeds up to mid-September. The ripening effect of the altered photoperiod diminished, however, after mid-September. This was also confirmed by the brown colour of the seed coat getting darker only in the control seeds at the end of the test period. Consequently, fully ripe structures were first found about a fortnight earlier in the control than in the light-treated seeds. The coincidental advancement of the anatomical potential examined by the X-ray method supported the role of the surface structures on the anatomical maturity of pine seeds and the timing of cone collecting. Received: 26 January 1998 / Accepted: 30 March 1998     

The effects of UV-B radiation on epidermal anatomy in loblolly pine (Pinus taeda L.) and Scots pine (Pinus sylvestris L.)

The effects of enhanced UV‐B radiation on the needle anatomy of loblolly pine (Pinus taeda L.) and Scots pine (Pinus sylvestris L.) were studied in the field under supplemental UV‐B radiation supplied by a modulated irradiation system. The supplemental UV‐B levels were designed to simulate either a 16 or 25% loss of stratospheric ozone over College Park, Maryland. Enhanced UV‐B radiation caused different responses in these two species. The needles of loblolly pine had larger amounts of tannin in the lumen of epidermal cells and more wall‐bound phenolics in the outer epidermal walls of UV‐B‐treated needles, whereas the most pronounced effect on Scots pine needles was increased cutinization. In both species, the outer epidermal cell walls thickened and the needle cross‐sectional and mesophyll areas decreased (statistically significantly only in Scots pine). This suggests that more carbon may have been allocated to the protection mechanisms at the expense of photosynthetic area. The difference in response between these species suggests that the response to UV‐B radiation is not mediated by a single mechanism and that no generalization with regard to the effects of UV‐B on conifers can be made.     

Food reserves of Scots pine (Pinus sylvestris L.)

Summary The amounts of starch, soluble sugars, triacylglycerols, diacylglycerols and free fatty acids were studied in Scots pine (Pinus sylvestris L.) during an annual cycle in current-year needles and in 1-, 2- and 3-year-old needles collected shortly after bud break. Determination of the compounds was performed using specific enzymatic assays, capillary gas chromatography and thin layer chromatography. Newly emerging needles contained relatively large amounts of starch, but only trace amounts of fat. During autumn and winter, fat content rose, while starch content decreased; amounts of both these reserve materials were very high the next spring shortly before bud break and decreased again during shoot elongation. Concentration of intermediates in triacylglycerol biosynthesis (diacylglycerols and free fatty acids), were low in summer and high in winter. The same pattern was observed for fructose and glucose (the predominant soluble sugars), galactose/arabinose and raffinose/melibiose. In contrast, sucrose concentrations were highest in spring and in autumn. Mature needles of different ages collected in May showed significant differences only in their triacylglycerol and starch content. Concentration changes of reserve materials are discussed in relation to season, mobilization and translocation processes, dormancy, frost resistance and the possibility of carbohydrate-fat interconversions.     

Factors affecting the distribution of cadmium,copper and lead and their effect upon yield and zinc content in bush beans (Phaseolus vulgaris L.)

Summary Concentrations of Cd, Pb and Cu in the roots, stems and leaves of bulgarian bush beans (Phaseolus vulgaris L.) were determined for plants grown in various soils of increasing levels of contamination of these metals. Most of each heavy metal absorbed by plants was retained in roots. Concentrations of Cd, Pb and Cu in roots increased in response to soil concentrations, whereas, in stems, only Cd and Pb concentrations increased and Cu concentration was relatively constant. It is thought that Cu transport to the stele was metabolically controlled, whereas Cd and Pb reached the stem by leakage across non suberised areas of the endodermis. Uptake of heavy metals was associated with a decrease in zinc content in plants and a decrease in yield. By regression analysis decrease in both zinc content and plant yield could be best related to Cd content in stems. Possible reasons for these effects are discussed.     

Distribution of Cd and Zn in annual xylem rings of young spruce trees [Picea abies (L.) Karst.] grown in contaminated soil

Concentrations and total amounts of Cd and Zn in individual annual xylem increments of 5-year-old spruce trees (Picea abies) were investigated after one growing season in contaminated soils. The plants had been potted in soils amended with different concentrations of Cd and Zn and kept in open air conditions. In the outer xylem rings formed during the treatment period lowest concentrations of Cd and Zn were found. In inner rings concentrations of both elements increased and were highest at the pith. Total amounts of both elements showed reversed distribution patterns with highest values in the outermost rings. The observed increase of total dry matter of the rings from pith towards the outer rings was stronger than the parallel increase in total element amounts in the rings. Thus, concentrations dropped in younger rings. The time of the soil contamination event cannot be inferred from element concentrations in annual rings. The results challenge the concept of monitoring historical trends in trace element pollution using analytical data of tree rings of conifers.     

Climate-tree-growth relationships of Scots pine stands (Pinus sylvestris L.) exposed to soil dryness

 Dendroclimatological techniques were used to assess the impact of climatic factors on radial tree growth (total ring-width and latewood-width) of stunted Scots pine trees (Pinus sylvestris L.) exposed to soil dryness and nutrient deficiency on a dolomite substrate. The response of eight scattered populations representing various habitats, yet influenced by the same regional climate was investigated. Total ring-width and latewood-width were dated, standardized and several chronology statistics, which estimate the chronology signal strength and the potential climate signal in the series, were determined. Dendroclimatic analysis comprised evaluation of event/pointer years as well as response function elements. Response function analysis indicates that at most sites wide rings are significantly associated with high precipitation in April to June and cool conditions in May of the current year, and high precipitation in August to September of the preceding year. Latewood chronologies show a lower climate signal at all sites. Limited water availability causes a homogeneous relationship of radial tree growth to climate at all habitats, though site characteristics (slope magnitude, slope aspect, soil depth, vegetation cover) differ substantially. Cluster analysis of negative event/pointer years suggests that within the study area stands react to extreme climatic events depending on susceptibility to soil dryness, which is primarily determined by site topography. Received: 5 January 1998 / Accepted: 22 April 1998     

A statistical model for the prediction of the number of sapwood rings in Scots pine (Pinus sylvestris L.)

Dendrochronology is a well-established dating method for wooden objects, but due to surface processing of construction timber or natural degradation the dating of historical wood often relies on a prediction of the number of missing rings based on sapwood statistics. Since Scots pine (Pinus sylvestris L.) is one of the most common tree species in north-western Europe, the absence of reliable sapwood statistics and models for the prediction of missing sapwood rings for pine samples is remarkable. We have therefore produced sapwood statistics based on data from 776 pine trees with ages from 15 to 345 years. The material consists of both living trees and historical timber, with varying growth rates, geographic settings, and from different soil types. When the whole material is considered, the average age of the trees is 103 years, and the number of sapwood rings is 54 ± 15 (1 SD), but range from 18 to 129. Trees less than 100-years in age contained 46 ± 11 (1 SD) sapwood rings and had an average tree-ring width (TRW) of 1.76 mm. With increasing age, the average TRW decreased while the number of sapwood rings increased. The average TRW of 101–200-year-old trees is 0.99 mm while the samples contained 63 ± 12 (1 SD) sapwood rings. For trees older than 201 years, the average TRW is 0.64 mm while the number of sapwood rings increased to 85 ± 16 (1 SD). The two most important factors in determining the number of sapwood rings for a given tree when only heartwood statistics are available proved to be (i) the number of heartwood rings and (ii) the average TRW of the heartwood rings. For incomplete samples, we have therefore developed a statistical model based on the sample’s heartwood rings (number and average width) to compute a prediction interval for the total number of rings. The sapwood and heartwood statistics suggest a statistical model for the number of sapwood rings with mean that increase with the number of heartwood rings. Furthermore, the average number of sapwood rings decreases with the mean width of the heartwood rings. However, the predictive power of the mean width is limited when the number of heartwood rings has already been taken into account. Thus, we suggest making predictions for the number of sapwood rings using only the number of heartwood rings. Predictions of the number of sapwood rings based on the statistical model where convincing in the case of the three different datasets that were analysed. The certainty in these predictions was such that the width of the 80% and 95% prediction intervals ranged 28–34 and 45–52 sapwood rings, respectively. Additionally, we demonstrate how make predictions when there is information about the number of remaining sapwood rings in a given sample. To make the sapwood model available, we present a free online R package for fitting our models and an online software dashboard.     

Influence of soil temperature on root freezing tolerance of Scots pine (Pinus sylvestris L.) seedlings

The influence of soil temperature on the root freezing tolerance of one-year-old containerized Scots pine (Pinus sylvestris L.) seedlings was investigated. In addition, the TTC and electrolyte leakage methods were evaluated in terms of their suitability for use in detecting damage to roots caused by freezing. In mid-August, seedlings were placed in three thermostat-controlled soil beds in a greenhouse with an initial soil temperature of 14.3 °C. Soil temperature was lowered in two of the soil beds, resulting in temperatures of 10.7 and 5.3 °C respectively. Each soil temperature, i.e. 14.3, 10.7 and 5.3 °C was maintained for eight weeks. Starting in early September, damage to roots induced by artificial freezing was estimated biweekly by measuring electrolyte leakage, triphenyl tetrazolium chloride (TTC) reduction and potential root growth in a three-week cultivation test. In addition, the root freezing tolerance of seedlings placed outdoors was tested. Measurements showed that these seedlings were exposed to soil temperatures ranging from 13.0 °C in mid-August to 0.5 °C in November. Generally, the development of root freezing tolerance was more pronounced for seedlings exposed to lower (0.5 and 5.3 °C) soil temperatures compared with those exposed to higher (10.7 and 14.3 °C) ones. Root freezing tolerance was highest among the seedlings placed outdoors which were also exposed to the lowest soil temperatures registered in the study. To examine the effect of a temporary warm period, the soil temperature in one treatment was increased from 5.4 °C to 13.9 °C, maintained at the latter temperature for two weeks in October and then lowered to 5.7 °C. Root freezing tolerance was reduced by exposure to the warmer soil temperature. However, after four weeks at the colder soil temperature, the tolerance of the seedlings had returned to the level measured prior to exposure to the warm soil temperature. Methods based on the measurement of root electrolyte leakage and TTC reduction were both found to have limitations when used to detect root freezing damages in containerized seedlings. This revised version was published online in June 2006 with corrections to the Cover Date.     

Monoterpenes emitted by the large pine weevil, Hylobius abietis (L.) feeding on Scots pine, Pinus sylvestris L.

Abstract. Attraction of the large pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae) to pine logs was enhanced by the presence of weevils already on the logs. We investigated the volatiles emitted by insects feeding on host trees to determine the mechanism of the attraction. Chemical analyses of the collected volatiles showed large amounts of monoterpenes. There were no qualitative differences in the collections, between Scots pine alone and pine plus feeding weevils, nor between males and females, mated or virgin. Strong quantitative differences in monoterpenes existed that were correlated to the surface of cut bark. Electroantennograms (EAGs) were recorded on male and female antennae stimulated by increasing doses of collected volatiles. EAGs for pine alone and pine plus weevils at the same stimulus loading in (a + P)-pinene gave superimposable curves. EAGs of male and female volatiles were also identical. These results suggest an attraction mediated by host plant allelochemicals rather than by an aggregation pheromone.     

Electrophoretic analysis of genetic linkage in Scots pine (Pinus sylvestris L.)

Eighty megagametophytes from each of 24 Scots pines (Pinus sylvestris L.) were subjected to horizontal starch gel electrophoresis. The trees were from crosses among widely separated provenances, and each was polymorphic for 8 to 14 loci. Evidence for linkage among 275 two-locus combinations was tested using chi-square analysis. Data from different trees were pooled to calculate map distances for the species. Nineteen of the twenty-nine loci tested were linked in one of six groups; the groups varied in size from two to seven loci. Similarities in linkage relationships among Scots pine, other pines, and other species within the Pinaceae support karyological research that suggests extensive conservation of the conifer genome.     

Seasonal changes in antioxidant level of scots pine (Pinus sylvestris L.) needles exposed to industrial pollution. I. ascorbate and thiol content

Current and previous year needles from three 16 years-old populations of Scots pine (Pinus sylvestris L.) trees were seasonally collected at the three experimental areas: Luboń- close to the phosphate fertiliser factory, Głogów — close to the copper foundry and Kórnik — control site. Głogów is the most polluted site, where at 1998 monthly mean daily concentrations of different pollutants were: SO₂ - 17 μg·m⁻³, NO_x - 12 μg·m⁻³ and dust containing heavy metals as Cu, Pb, Cd - 29 μg·m⁻³. Trees growing in Luboń were influenced for many years by high concentration of SO₂ and fluor compounds. A few years ago emissions were markedly reduced, but low pH of soil and high concentration of aluminium ions still influence the growth of trees. Seasonal changes of ascorbate and thiol content were observed in each needle class and population, with the maximum in the winter and minimum in the summer. In needles from trees growing on polluted sites higher level of ascorbic acid and thiols comparing to control site was observed. Significant differences appeared in each population of Scots pine growing under higher pollution stress in the Głogów site. In needles from trees growing in Luboń significant differences in ascorbic acid and thiols content were evidently less numerous. Needles from polluted sites in some seasons contained significantly more malondialdehyde (MDA) and those was more frequent in Głogów than in Luboń. The results indicated that in the Głogów site trees are more influenced by pollution stress than in Luboń and the defense reaction measured as an increase of the antioxidant level is more evident.     

Ectomycorrhizal development on Scots pine (Pinus sylvestris L.) seedlings in different soils

In the present study ectomycorrhizal development of Laccaria bicolor, Rhizopogon luteolus and Suillus bovinus associated with Scots pine (Pinus sylvestris) seedings was studied as affected by primary stand humus, secondary stand humus, podsolic sandy soil or peat in perspex growth chambers. After 9 weeks, ectomycorrhizal development with S. bovinus was significantly greater in peat and primary stand humus than in secondary stand humus or podsolic sandy soil. Ectomycorrhizal development with R. luteolus in secondary stand humus was higher than in primary stand humus. Degree of ectomycorrhizal development of L. bicolor, R. lutuelus and S. bovinus on Scots pine was related to potassium concentration, organic matter content and pH of the soils suggesting that chemical composition of the soils affects ectomycorrhizal development.