Impact of Contaminants From Oil Shale Processing on Forest Ecosystems

Oil shale mining and processing in northeast Estonia have brought about several ecological problems. The mined oil shale is used as fuel in power stations and in processing plants producing crude oil and about 40 manufactured articles. Pollutants emitted from oil shale processing and chemical plants include SO₂, CO, NO _x , oil shale fly ash, and organic compounds in which aromatic and aliphatic hydrocarbons, phenols, formaldehyde, etc., are represented. Pollution has caused changes in the condition of the forest ecosystem and the chemical character of soil and ground water. The condition of coniferous forest sites was investigated in 1995–1998. Because of the high concentration of alkaline fly ash in the air, the pH of rain water is somewhat elevated (pH = 7.0–7.1) and exceeds the level regarded as normal for rain water. The analysis of the soil samples showed that the concentrations of Ca, Mg and K, which dominate in the solid fraction of the pollutant mixture, are high, being respectively 18, 14, and 4 times as high as the control. The increases in the concentrations of K, Mg, Cu, Pb, and Ni in stemwood reflect increases in the regional oil shale fly ash deposition. Conifers influenced by high levels of air pollution emitted from the oil shale industry are characterized by retarded growth of needles and shoots and radial growth as a result of disturbances in their mineral nutrition and imbalance in their mineral composition.     

Uptake of benzyladenine in explants of Picea abies and Pinus sylvestris

Primary explants of Picea abies (L.) Karst. and Pinus sylvestris L. were incubated for various periods in solutions containing different concentrations of [¹⁴C]-benzyladenine. Time course analysis showed that uptake was linear for the initial 60 min; after this time linear uptake continued but at a much reduced rate. The amount of benzyladenine taken up by the explants saturated at a concentration about one third of that of the medium, Concentration dependence experiments showed that BA uptake was directly proportional to the external concentration. These results and those from experiments in which uptake was examined at different temperatures are consistent with a passive mode of BA uptake. The results are discussed with respect to in vitro micropropagation.     

Needle analyses and graphic vector analyses of Norway spruce and Scots pine stands

 The aims of this study were to test the nutritional condition of young stands of Norway spruce and Scots pine in southernmost Norway, where atmospheric inputs of anthropogenic nitrogen and strong acids, as well as deposition of magnesium by sea spray, are relatively high. There has been general concern about forest health in this region, specifically about nutrient deficiencies and nitrogen saturation caused by the atmospheric pollution. To expand our knowledge about these subjects, needle analyses and graphic vector analyses, as well as tree vigour index and height growth response to fertiliser application, were used as diagnostic tools. The overall conclusion is that phosphorous limitation is probably more frequent than expected in the coastal zone of southern Norway, especially in spruce on shallow soils. It is hypothesised that phosphorous limitation might be a more general problem or will arise as one in a 30 – 60 km coastal zone stretching from southernmost Sweden along the coast to the north and then westwards along the south coast of Norway. Nitrogen saturation in southernmost Norway might already occur on some sites with shallow soils. Needle analyses and graphic vector analyses can yield erroneous results if stress factors other than nutrient supply, e. g. severe drought, determine the growth rate of trees. In addition, the graphic vector analyses require that nutrients applied to the test plots are available to roots early in the growing season, and that needle nutrient concentrations in late autumn reflect those when the needles were formed. Received: 30 May 1995/Accepted: 27 December 1995     

Fractionated extraction and speciation of macronutrients from Scots pine and Norway spruce needles of different ages

Summary The relationship between proteins and the macroelements potassium, magnesium, calcium, sulphur and phosphorus was studied in homogenates of needles of different ages from Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.]. Complete extractions by acid digestion, protein extractions by a buffer-detergent and non-protein extractions by a buffer alone showed that most of the potassium and magnesium of the needles was soluble independent of the proteins. Only a minor part (50–60 ppm, dry weight) of the magnesium could be referred to the chlorophyll content of the needles. Both potassium and magnesium appeared independent of the protein gel filtration. This was also valid for the minor fraction of the total calcium content, which was extractable in a buffer. Heterogeneous calcium deposits are also suggested by extraction using a chelator (EGTA). Part of the buffer-soluble sulphur and phosphorus compounds occurred independently of solubilized proteins, and had complex distribution patterns after gel filtration, even outside the high molecular separation range. It is suggested that further analyses along the present lines, compared with conventional analyses on total extracts, may extend the usefulness of mineral nutrient analyses in plants.     

Sensitivity of managed boreal forests in Finland to climate change, with implications for adaptive management

This study investigated the sensitivity of managed boreal forests to climate change, with consequent needs to adapt the management to climate change. Model simulations representing the Finnish territory between 60 and 70 degrees N showed that climate change may substantially change the dynamics of managed boreal forests in northern Europe. This is especially probable at the northern and southern edges of this forest zone. In the north, forest growth may increase, but the special features of northern forests may be diminished. In the south, climate change may create a suboptimal environment for Norway spruce. Dominance of Scots pine may increase on less fertile sites currently occupied by Norway spruce. Birches may compete with Scots pine even in these sites and the dominance of birches may increase. These changes may reduce the total forest growth locally but, over the whole of Finland, total forest growth may increase by 44%, with an increase of 82% in the potential cutting drain. The choice of appropriate species and reduced rotation length may sustain the productivity of forest land under climate change.     

Effect of acid rain on pine needles as food for capercaillie in winter

The effects of long-range air pollution on the chemical composition of needles of Scots pinePinus sylvestris and consequences for capercaillieTetrao urogallus feeding on the needles were studied. Samples of pine needles from localities receiving different amounts of acid rain were taken in early March. Concentrations of N and P were highest in needles from the most acidified areas, and N concentration was 35% higher in the most heavily polluted area than in the least polluted. Secondary chemicals decreased significantly with increasing acidification. Concentrations of Cd in pine needles were closely correlated with the acid deposition levels, with highest concentrations in the most polluted area. Al concentration also increased with increasing acidification. These results provide evidence that acid rain increases the nutritive value of pine needles through a fertilizing effect. Enhanced levels of certain metals are considered too low to be directly toxic to capercaillie. However, behavioural anomalies cannot be excluded.     

Climatic control of intra- and inter-annual wood-formation dynamics of Scots pine in northern Finland

Boreal forests are highly sensitive to climate and human impacts and therefore suitable as biological indicator for environmental changes. In this context, our study was aimed at getting deeper insight into the climate-dependence of the onset, intensity and end of wood formation of Scots pine during the growing season.We monitored the intra-annual growth dynamics of, on average, 42-year-old Scots pine trees over five consecutive years, 2000-2004, at two sites located 80 and 300 km south of the tree line in northern Finland. For that purpose, the cambium of the trees was weekly wounded with a pin and the resulting wound tissue, microscopically detectable in transverse thin-sections through the newly built wood, was taken as a time marker. During this 5-year study period, the intra-annual wood formation at the southern site was mainly positively associated with summer temperature. However, at the northern site such an association was either entirely missing or negative. At both sites, two thirds of the radial growth was produced within only 4 weeks from mid-June to mid-July, independent of whether the growing season started earlier or later.Moreover, we measured the widths of all tree rings from bark to pith (inter-annual growth) of the same study trees and assembled them to 51-year long tree-ring site chronologies. Since 1999, these two site chronologies - after having run fairly parallel over the preceding decades - were running in divergent directions thus corroborating our results derived from the intra-annual climate/growth analysis. Whereas the chronology of the southern site follows the average temperature of May and July very closely from 1961 up to 2004, the chronology of the northern site follows the July temperature, but only up to 1998, and from 1999 to 2004 is running just opposite to the distinctly rising July temperature. During the same period, there was - unlike in the years before - nearly no snow cover in May at the northern site, whereas at the southern site there was no change of the normally existing slight snow cover in May. This deviating weather situation may have led to a temperature-induced, temporary drought stress for the Scots pines at the northern site.     

Characterization of tree macroremains production in a recently burned conifer forest in northern Québec,Canada

The production of plant macroremains was studied in a conifer forest twomonths after it burned in 1996 in northern Québec. The proportions ofvarious types of charred and uncharred pieces (needles, cones andwood) produced by black spruce (Piceamariana) and jack pine (Pinusbanksiana) were determined by sampling around individual trees.Both species produced equivalent masses of charred material, but pieces of woodcharcoal from jack pine are generally larger that those of black spruce. Theproportion of charred versus uncharred needles is the bestindicator of the species dominance in the forest. Although the fall of uncharredneedles is delayed from the time of a fire, they contribute to more than half ofall remains produced. Jack pine cones remain on the tree for a long time after afire, while charred cones of black spruce are dehiscent (cones come off thebranches easily). Trees are poor wood charcoal producers compared toundergrowth shrubs. As a result, the macroremains assemblage associated with afire event is made up of large amount of uncharred material from trees and alarge proportion of charred pieces produced by undergrowth vegetation. Modernassemblages of plant macroremains indicate that in order to reconstruct pastvegetation associated with fire disturbance, it is important to distinguishbetween the various types of remains, because wood charcoal is mainly producedby material that is already dead.     

Total, organic and extractable-P in humus and soil beneath Sitka spruce planted in pure stands and in mixture with Scots pine

Total, organic and extractable P were measured in the humus and underlying soil to 10 cm depth beneath Sitka spruce (SS) and mixed Sitka spruce and Scots pine (SS+SP) stands planted on upland heath. The humus beneath SS+SP contained significantly (p<0.01) greater amounts of total and organic-P than that in SS and the mixed stands had more effectively retained approximately 87 per cent of previously applied fertilizer-P, totalling 100 kg P ha^–1, compared with 70 per cent in SS. Despite the larger amounts of total-P in the mixed plots 0.01 M CaCl₂ extractable molybdate reactive phosphorus (MRP) was significantly (p<0.05) greater in SS+SP humus only during March and April. Greater concentrations of MRP were released from the humus and soil during July and August at a mean rate of 58 g P ha^–1 day^–1. This coincided with drying of the soil during the summer and the rate of release, attributed to death of fine roots and microorganisms, was 4 to 30 times greater than reported values for rates of net mineralization of P from forest soils.     

A population ‘consensus’, partial linkage map of Picea abies Karst. based on RAPD markers

 We built a “consensus” partial linkage map based on RAPD markers using 48 sibships of eight megagametophytes each from a natural population of Norway spruce. A RAPD linkage map for a single individual from the same population had previously been constructed. Using 30 random decamers that had yielded 83 RAPD markers in the single-tree map, eight megagametophytes for each of the 48 sibships were screened. The linkage relationship among markers was estimated considering each family of eight megagametophytes as a progeny of a phase-unknown backcross mating between a heterozygous mother and a fictitious ‘recessive’ father. Markers were assigned to windows using LOD=2.0 and θ=0.4 as thresholds, and ordered using a criterion of interval support ≥2.0. For eight “windows” of recombination selected on the single-tree map, we investigated the consistency of marker order in the two maps. We adopted restrictive criteria for rejecting co-linearity between the two locus orders. For each window we imposed the most likely locus order obtained from one data set to the other (and vice versa), obtaining two symmetrical log-likelihood differences. We considered the hypothesis of co-linearity rejected when both symmetrical differences were significant (ΔLOD>3.0). By bootstrapping a subset of markers for each window (highly informative, ‘framework’ loci chosen on the previous single-tree map using a matrix correlation method) the sampling variability of the single-tree and population maps was estimated. As expected the population map was affected by a larger variability than the single-tree map. Heterogeneity in pairwise recombination fractions among groups of sibship revealed a (possible) alternative genomic arrangement detected within a single recombination window. Received : 4 January 1997 / Accepted : 24 January 1997     

Condition of Scots pine fine roots and mycorrhiza after fungicide application and low-level ozone exposure in a 2-year field experiment

 In a 2-year experiment at an open-air ozone fumigation field, the effects of fungicide application and low-level ozone exposure, single and combined, on fine root and mycorrhiza condition of Scots pine (Pinus sylvestris) seedlings were studied. Two different fungicides, copper oxychloride and propiconazole, with different modes of actions, were used. Propiconazole treatment reduced mycorrhizal infection in both years while copper oxychloride treatment and ozone exposure slightly stimulated mycorrhizal infection after the first year. Different mycorrhizal morphotypes showed different kinds of responses to the two fungicides. Light brown morphotype appeared to be the most sensitive one to propiconazole treatment. After the second year, ectendomycorrhizas disappeared in propiconazole treatment while in control treatment ectendomycorrhizas formed the majority of the light brown morphotype. The root biomass was not affected by fungicide treatments, but ozone exposure increased the total amount of short roots and the fresh weight of propiconazole treated roots. No significant differences in the concentrations of ergosterol, starch and total phenolics in pine roots between treatments were found. However, ergosterol concentration correlated positively with the mycorrhizal infection level. Both fungicides reduced the soil respiration compared to controls. At the ultrastructural level, both fungicides caused increased transparency and gradual granulation and degeneration of cytoplasm in the fungal symbiont of mycorrhizal short roots. Slightly elevated ozone did not have harmful effects on root ultrastructure. These results suggest that fungicides have deleterious effects on the quantity and quality of mycorrhizas in Scots pine roots and also side-effects on non-target soil fungi. Some of these deleterious effects were noticeable only at the ultrastructural level. Received: 23 June 1997 / Accepted: 11 December 1997     

Characterisation of the differential interference effects of two boreal dwarf shrub species

Our purpose was to characterize the relative competitive and phytotoxic potential of two closely related dwarf-shrub species, Empetrum nigrum and E. hermaphroditum, which form clones in a mosaic pattern in post-fire successions of the boreal forest of northern Sweden. We determined morphological and growth parameters of both species, performed bioassays and chemical analysis, and established field experiments to explore possible differing interference effects on trees by the two species. Both Empetrum species had very similar morphological and growth characteristics. E. hermaphroditum exerted considerably greater negative effects than E. nigrum against Pinus sylvestris and Populus tremula seed and seedlings. These negative effects were related to the different substitution of a bibenzyl in the two species. The effect on seed germination of the two bibenzyls isolated from E. nigrum and E. hermaphroditum was compared with that of other simple phenolics; the latter were found to be inactive, indicating a specific phytotoxic component in the bibenzyls. P. sylvestris seeds planted into clones of both species in the field revealed that E. hermaphroditum had much stronger inhibitory effects than did E. nigrum and the addition of activated carbon partially reversed these effects. Seed germination, biomass and survival of P. sylvestris after four seasons were significantly lower in E. hermaphroditum- than in E. nigrum-dominated plots. We conclude that while both Empetrum species are superficially very similar morphologically, they have vastly different effects on tree seed germination, seedling establishment and growth. These effects appear to be due, at least in part, to the different chemical profile of the two species. Received: 2 April 1999 / Accepted: 2 November 1999     

NMR imaging of roots: effects after root freezing of containerised conifer seedlings

Seedlings of Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L) Karst.] were subjected to low root temperatures, and 10 days later the roots were examined by NMR imaging. The amount of NMR detectable roots decreased with decreasing temperature, with the signal from the younger roots at the bottom of the container being the first to disappear. The origin of the loss of NMR signal is unclear but may be due to changes in the NMR properties of root water after cold damage. A recent method is discussed for obtaining unbiased estimates of root lengths from a series of total vertical projections; the method is particularly suited to evaluating NMR projection images. Since NMR imaging methods can apparently distinguish between control and cold damaged roots, it may be possible to design more routine applications using low resolution NMR methods.     

Pollen competition and seed-siring success in Picea abies

The aim of the present work was to study pollen-tube competition in Picea abies. Controlled crossings were performed with pollen mixtures including pairs of pollen lots with fast and slowly elongating pollen-tubes. Paternity analysis using isozyme markers was performed on the progenies in order to study whether the in vitro pollen-germination vigour corresponds to the proportion of seeds sired by the pollen donor. Paternal success was found to be unequal, 15 out of 23 crossings producing progeny that differed significantly from the hypothetical ratio of 1:1. The paternal contribution in the majority of the crossings was as expected: the pollen parent with more-vigorous in vitro germination sired more seeds than the less-vigorous pollen. In the case of two pollen mixtures, however, the seed-siring success summed over the maternal trees was the opposite to the expected value. Despite these aberrations, the results support the hypothesis that pollen-tube competition is one of the factors contributing to male fitness in P. abies. However, when all the other factors affecting pollination and seed set under natural conditions are taken into account, it is clear that the seed-siring success of a particular paternal genotype cannot be predicted reliably by measuring only the in vitro pollen vigour. Received: 2 July 2001 / Accepted: 7 August 2001     

Effects of ozone and mild drought stress on gas exchange, antioxidants and chloroplast pigments in current-year needles of young Norway spruce [Picea abies (L.) Karst.]

 To investigate the effects of ozone exposure and soil drought, singly and in combination, on gas exchange, antioxidant contents and pigments in current-year needles of Norway spruce [Picea abies (L.) Karst.] 4-year-old seedlings were fumigated in growth chambers with either charcoal-filtered air or with 100 nl l^–1 ozone for 106 days. After 3 weeks a 20% reduction in gas exchange was observed in ozone-treated seedlings. However, no further decrease occurred in spite of continued ozone exposure. Whole needle ascorbate and apoplastic ascorbate increased until the end of the experiment and contents were 62% and 82%, respectively, higher than in ozone-free controls. This increase in ascorbate might have protected net photosynthesis from further decline. Ozone pre-treated plants and ozone-free controls were subjected to soil drought for 38 days which caused stomatal narrowing. Thereby ozone uptake was reduced when compared to well watered seedlings. At the end of the experiment drought alone, and even more in combination with ozone, had also caused an increase in ascorbate. Glutathione increased only in drought-stressed seedlings. The redox states of the ascorbate and the glutathione pools were not affected by any treatment. Superoxide dismutase activity declined under both stresses but was most reduced by ozone alone. While chlorophyll and neoxanthin contents remained unchanged, carotenes were significantly decreased upon drought. The combination of O₃ and drought induced increased lutein contents, an increased pool size of the xanthophyll cycle as well as an increased epoxidation status of the xanthophyll cycle. These results suggest that spruce needles seem to be able to acclimate to ozone stress but also to drought stress by increasing their ascorbate pools and protecting pigments. Received: 15 September 1997 / Accepted: 24 March 1998