Search

The paper presents an assessment of vegetation (composition and cover-abundance), nutrient supply, and especially sulfur accumulation in the trees foliage (birch and pine) used in reforestation and wood small-reed (Calamagrostis epigejos (L.) Roth) appearing in succession on reclaimed areas of the former Jeziórko sulfur mine (southern Poland, Tarnobrzeg region). In researched area, three categories of vegetation cover were determined: category D-degraded and unsuccessfully reforested plots, and two categories with successful reforestation: P-pine and B-birch stands. On each category, four study plots (4-6 areas each, depends on site category) were established. Soil and vegetation samplings (current year and 2-year-old pine needles, birch leaves, and wood small-reed foliage) were collected on the subplots established in regular grid square (10 × 10 m) in each category. Basic soil properties and nutrient content in soils and vegetation were analyzed. Trees grew well in areas where neutralization and reclamation treatments were carried out properly and showed a good supply of nutrients (exception of phosphorus and nitrogen), while on category D, only herbaceous vegetation with low cover-abundance and dominated by wood small-reed were noted. Linear correlations between the soil and trees nutrients content occurred, while the correlations between the soil and wood small-reed did not occur. Wood small-reed did not display increased sulfur uptake which may indicate a strategy of blocking pollutant uptake from the soil and may be recommended as a species resistant to sulfurous soils.

Climate change, higher levels of natural resource demands, and changing land use will likely lead to changes in vegetation configuration in the mountain regions. The aim of this study was to determine if the vegetation cover and composition have changed in the Swedish region of the Scandinavian Mountain Range, based on data from the long-term landscape biodiversity monitoring program NILS (National Inventory of Landscapes in Sweden). Habitat type and vegetation cover were assessed in 1740 systematically distributed permanent field plots grouped into 145 sample units across the mountain range. Horvitz-Thompson estimations were used to estimate the present areal extension of the alpine and the mountain birch forest areas of the mountain range, the cover of trees, shrubs, and plants, and the composition of the bottom layer vegetation. We employed the data from two subsequent 5-year monitoring periods, 2003-2007 and 2008-2012, to determine if there have been any changes in these characteristics. We found that the extension of the alpine and the mountain birch forest areas has not changed between the inventory phases. However, the total tree canopy cover increased in the alpine area, the cover of graminoids and dwarf shrubs and the total cover of field vegetation increased in both the alpine area and the mountain birch forest, the bryophytes decreased in the alpine area, and the foliose lichens decreased in the mountain birch forest. The observed changes in vegetation cover and composition, as assessed by systematic data in a national and regional monitoring scheme, can validate the results of local studies, experimental studies, and models. Through benchmark assessments, monitoring data also contributes to governmental policies and land-management strategies as well as to directed cause and effect analyses.

Trees in cutaway peatland are growing in difficult conditions. Fertilization with nutrient-rich wood ash helps improve growth conditions. Photosynthesis and carbohydrate concentration along leaf anatomy were studied on plots treated with 10 and 5 t ha(-1) wood ash (WA10 and WA5) and on untreated (Control) plot to explain the physiological background of the differences in tree growth. The leaves from WA10 had the largest leaf area, total thickness, the thickest mesophyll and also significantly larger average values of all anatomical parameters of the shoots. The photosynthetic assimilation was significantly higher on treated plots at 200 and 400 ppm CO2 levels. In leaves on the treated plots, the sucrose concentration was lower while that of starch was higher than in trees on untreated soil. The differences in the maximum photosynthesis were relatively small. At unit ground, the leaf area provided for a wood ash-treated tree an efficient surface for CO2 assimilation, light interception and some starch storage during the growing period.

The negative impacts of mammalian herbivores on plants have been studied quite extensively, but typically with only a single herbivore species at a time. We conducted a novel comparison of the browsing effects of voles, hares and cervids upon the growth and survival of boreal tree seedlings. This was done by excluding varying assemblages of these key mammalian herbivores from silver birch, Scots pine and Norway spruce seedlings for 3 years. We hypothesised that the pooled impacts of the herbivores would be greater than that of any individual group, while the cervids would be the group with the strongest impact. Growth of birch seedlings advanced when cervids were excluded whereas growth of seedlings accessible to cervids was hindered. Survival of all seedlings was lowest when they were accessible to voles and voles plus hares, whereas cervids seemed not to influence seedling survival. Our results show that the impact of herbivores upon woody plants can be potent in the boreal forests, but the mechanism and strength of this link depends on the tree and herbivore species in question. Risk of abated stand regeneration appears highest for the deciduous birch, though there is need for seedling protection also in coniferous stands. The clear cervid-mediated growth limitation of birch also indicates potential for a trophic cascade effect by mammalian top predators, currently returning to boreal ecosystems.

Boron (B) deficiency is a common micronutrient deficiency that has been reported to affect the phenolic metabolism of plants. Thus, it may play a role in defense against herbivorous animals. However, the role of B in a plant's resistance to herbivores has not received any particular attention from researchers. In this study, we tested the effects of B nutrition 1) on the biochemical and mechanical defenses of birches and the growth of seedlings, and 2) the resistance of seedlings to larvae of the autumnal moth, Epirrita autumnata. Boron fertilization improved the resistance of birch, which was shown as reduced pupal weight of the herbivore. However, B fertilized trees suffered from heavier defoliation than unfertilized ones due to compensation feeding of larvae. The growth of the seedlings was diminished, and several biochemical changes occurred in leaves of herbivore seedlings, and B also played a role in these changes. Polyphenoloxidases (PPOs) and peroxidases (PODs) and their substrates, chlorogenic acids, were induced by herbivory in B fertilized seedlings but not in unfertilized seedlings. The lower pupal weights and increased consumption of the herbivores were probably linked to the plants' phenoloxidase-mediated production of reactive quinones, which decrease the nutritive value. Herbivory upon new stems led to an increase in the number of resin glands that provide defense against mammalian herbivores. Herbivory also had a substantially negative effect on B concentration in leaves of B fertilized seedlings. We postulate that B nutrition of trees may play a significant role in the induced defense of birches.

Spatial distribution of ectomycorrhizae-associated basidiomycetes was determined in oakbirch forest using terminal restriction fragment length polymorphism (T-RFLP) analysis. The data were correlated with actual soil humidity, pH, electric conductivity of the soil extract, absorbance A(465) and A(665) of water and alkali soil extracts and with the ratio A(465)/A(665) (parameter A4/A6). Natural non-homogeneity of the soil parameters was used as experimental gradient. Distance-based redundancy analysis of the T-RFLP data (with soil parameters being taken as environmental parameters) provided significant results when ITS1F-terminanted restriction fragments were analyzed. Among other fungi, a Mycena galericulata related fungus was observed to correlate negatively with A4/A6, indicating its association with highly humified soil organic matter. Positive association of other, unidentified fungi with A4/A6 was also observed. Several other unidentified fungi negatively correlated with electric conductivity of the soil extract. The results may explain nonhomogeneity of the spatial distribution of the fungi associated with ectomycorrhizae as a result of their interaction with non-homogeneous soil environment.

Environmental stress affects ectomycorrhizal communities (ECM), but it is not known how general the detected ECM responses are. We investigated ECM fungi on roots of mountain birch, Betula pubescens subsp. czerepanovii (Orlova) Hämet-Ahti, along three environmental gradients, two natural (altitude, seashore) and one human-induced (pollution), within the Kola Peninsula, NW Russia. Chlorophyll fluorescence of birch leaves indicated no environmental stress even in the conditions that were presumed most stressful in terms of abiotic environment, where the biomass and population density of birches were strongly reduced. Although neither overall ECM colonisation nor root fungal biomass showed stress-related patterns, colonisation by Cenococcum geophilum tended to decrease with abiotic stress. ECM morphotype diversity declined with abiotic stress, and along altitudinal gradient this decline was related to an increase in proportion of morphotypes with high fungal biomass. Polycormic birches had higher ECM colonisation than monocormic birches at high stress sites only. ECM morphotype diversity increased with foliar nitrogen concentration at low stress sites, but not at high stress sites. Birches with higher chlorophyll fluorescence had lower chitin concentration in their roots (indicating lower proportion of fungal structures) at high stress sites only. Our results suggest that at high stress sites (1) mechanical shelter created by polycormic trees may favour ECM fungi and (2) mountain birches maintain lower ECM diversity than at low stress sites.

Long-term fertilization of acidic tussock tundra has led to changes in plant species composition, increases in aboveground production and biomass and substantial losses of soil organic carbon (SOC). Root litter is an important input to SOC pools, although little is known about fine root demography in tussock tundra. In this study, we examined the response of fine root production and live standing fine root biomass to short- and long-term fertilization, as changes in fine root demography may contribute to observed declines in SOC. Live standing fine root biomass increased with long-term fertilization, while fine root production declined, reflecting replacement of the annual fine root system of Eriophorum vaginatum, with the long-lived fine roots of Betula nana. Fine root production increased in fertilized plots during an unusually warm growing season, but remained unchanged in control plots, consistent with observations that B. nana shows a positive response to climate warming. Calculations based on a few simple assumptions suggest changes in fine root demography with long-term fertilization and species replacement could account for between 20 and 39% of the observed declines in SOC stocks.

A serious forest decline of Betula ermanii Cham. has been observed at Mt. Mae-Shirane, Oku-Nikko, Japan, where high ozone (O3) concentration and severe water deficiency have been measured. In order to consider the possibility whether O3 and/or water stresses could have been the causes of the forest decline of B. ermanii, plant growth experiments were conducted in environment-controlled growth cabinets. Two-year-old seedlings of B. ermanii were exposed to either charcoal-filtered air (O3 concentration <5 ppb) or 50 ppb O3 (daily average, ranging between 20-100 ppb) for 123 days at 20.0/12.5 +/- 1.0 degrees C (day/night) and 70/80 +/- 7% relative humidity (day/night). Simultaneously, seedlings were treated with three watering regime: 1.0 < pF < 1.8 (no water stress), 1.8 < pF < 2.5 (mild water stress) or 2.5 < pF < 3.0 (severe water stress). O3 exposure significantly reduced the dry weights of leaf, root and the whole plant, while water stress significantly reduced the dry weights of each organ and the whole plant. Significant reductions of net photosynthesis, transpiration and stomatal conductance were also observed under O3 and/or water deficiency treatments, while contents of RuBP carboxylase/oxygenase (Rubisco), chlorophyll( a+b ) and some essential nutrient elements (N, P, K, Mg and Ca) were not markedly changed. It was suggested that the decrease in net photosynthetic rate induced mainly by stomatal closure was the major cause of the growth reduction under O3 and/or water stresses. No significant interactions between O3 and water stresses were observed in terms of the depression of dry matter production, which suggested that simultaneous stress treatments of O3 exposure and water deficiency could affect the tree growth of B. ermanii additively.

Leaf litter and other organic resources returned to the soil are important regulators of ecological processes in forest ecosystems, and their ecological impacts may be strongly influenced both by their quality and by interactions between coexisting resource types. To date, most studies on effects of resource identity and mixing have only involved leaf litter, despite the fact that other resource types constitute a major input to the soil. We investigated how quality and heterogeneity of organic substrates found in boreal forests affects the activity and community structure of soil microbes, and plant growth. Six organic substrates (wood, charcoal, berries, sporocarps, vertebrate faeces and leaf litter) were added singly or in mixtures of two, three and six resource types to pots containing forest soil (with or without tree seedlings of Betula pendula Roth). The largest positive effects of single substrates on microbial basal respiration (BR), substrate-induced respiration (SIR) and microbial metabolic quotient (qCO(2)) were found for nutrient-rich substrates (faeces and sporocarps) or substrates with high sugar-content (berries). Mixing of substrates had no effect on BR or SIR, but decreased qCO(2) or altered the microbial community structure for specific combinations of substrates. In contrast to the niche complementarity hypothesis, microbial catabolic diversity was not stimulated by greater diversity of resources. Seedling growth responses to single substrates were neutral or negative; the inhibition of growth probably resulted largely from microbial competition for nutrients. Substrate mixing enhanced seedling nutrient-uptake and growth for all mixtures containing sporocarps and leaf litter. Overall, plants responded more strongly to resource heterogeneity than microbes, and synergistic effects only occurred when nutrient-rich substrates were present within the substrate mixtures. In particular, our results demonstrate a role for complex and non-additive interactions among previously overlooked resource types returned to the soil in influencing ecosystem functions such as nutrient cycling and plant productivity.

We studied the effects of the intensity of intraspecific competition, as indicated by seedling density, and competitive success within populations, as indicated by seedling size, on the secondary chemistry of the stems of silver birch seedlings and their palatability to field voles. We found that the size of seedlings and their total phenolic concentrations were inversely related to stand density. Voles, however, did not discriminate between seedlings grown at the densities studied. Variation in competitive success was associated with several phenomena. Seedling size was positively related to the concentrations of several secondary compounds and to vole preferences, but negatively related to nitrogen concentration. We suggest that the ecological consequences of variation in seedling size cannot be predicted without knowledge of the mechanism behind the variation.

This study investigates variation in the secondary chemistry of the bark of three closely related, winter-dormant species of white birch (Betula resinifera, B. pendula, and B. platyphylla) at different ontogenetic stages by using different plant parts (top and base). The experimental birches were grown for 4 years in two growing conditions (pot and field) at different nutrient levels. There was considerable species-specific quantitative and qualitative variation in the secondary chemicals in bark, but this was also affected by fertilization and the age of the plant. In general, there was greater chemical diversity in saplings than in seedlings. The study revealed three new components, secoisolariciresinol 9-O-beta-glucopyranoside and two of its derivatives, that have not been reported previously for the bark of white birches. Principal component analysis showed that the species studied had a similar chemical composition at the juvenile stage, but as the plants grew, they became more clearly differentiated, which indicates that the species of older plants can be identified by chemotaxonomy. Evidently, the secondary chemistry of birches is under genetic control, but it is affected by properties of growing conditions and ontogeny.

The two forest-defoliating geometrid moth species Operophtera brumata and Epirrita autumnata are known to exhibit different altitudinal distribution patterns in northern birch forests. One possible explanation for this is that altitudinal climatic variation differentially affects the performance of two species through mismatching larval and host plant phenology. We explored this hypothesis by investigating the relationship between larval phenology and leaf phenology of Betula pubescens, which is the main host plant of both moth species, along ten replicate altitudinal transects during two springs with contrasting climate in northern Norway. There was a distinct monotonous cline in host plant phenology with increasing altitude in both years of the study, but the development of the leaves were generally 14 days later in the first of the 2 years due to cold spring weather. We found that larval development of both species closely tracked host plant leaf phenology independent of altitude and year. However, at the time of sampling, E. autumnata was approximately one instar ahead of O. brumata at all altitudes, probably reflecting that E. autumnata has faster early instar growth than O. brumata. The abundance of O. brumata was lowest at the altitudinal forest-line, while E. autumnata was lowest near sea level. Our results do not indicate that the altitudinal distribution patterns of the two moth species is due to any phenological mismatch between larval and host plant phenology. We suggest rather that natural enemies at low altitudes limit larval survival and thus abundance of E. autumnata, while an early onset of winter at the forest limit reduces survival of late eclosing adults of O. brumata.

The prevention of flower formation is important for avoiding the spread of transgenes from genetically modified plants into wild populations. Moreover, the resources not expended for the generation of flowers and fruits might be allocated to increased vegetative growth. We have been developing methods for preventing flower formation in silver birch (Betula pendula), a tree species of considerable economical importance in the boreal region. Here we study the suitability of the promoter of BpFRUITFULL-LIKE1 (BpFULL1, formerly BpMADS5) for tissue-specific ablation of inflorescences in Arabidopsis, tobacco and birch. With all these species, the development of inflorescences was successfully prevented. The results show that the BpFULL1::BARNASE construct has potential biotechnological applications in different plant species.

Boreal forest trees are highly dependent on root-colonizing mycorrhizal fungi. Since the maintenance of mycorrhizal symbiosis implies a significant carbon cost for the host plant, the loss of photosynthetic leaf area due to herbivory is expected to reduce the host investment in mycorrhizae. We tested this hypothesis in a common garden experiment by exposing ectomycorrhizal white birch (Betula pubescens Ehrh.) seedlings to simulated insect defoliation of 50 or 100% intensity during either the previous or the current summer or repeatedly during both seasons before harvest. The shoot and root growth of the seedlings were distinctly reduced by both 100% defoliation and repeated 50% defoliation, and they were more strongly affected by previous-year than current-year defoliation. The root to shoot ratio significantly decreased after 100% defoliation, indicating reduced proportional allocation to the roots. Ergosterol concentration (i.e. fungal biomass) in the fine roots decreased by 100% defoliation conducted either in the year of harvest or in both years. No such decrease occurred following the 100% defoliation conducted in the previous year, indicating the importance of current photosynthates for fungal symbionts. The trend was similar in the colonization percentage of thick-mantled mycorrhizae in the roots, the most marked decline occurring in the repeatedly defoliated seedlings. The present results thus support the prediction that the plant investment in ectomycorrhizae may decline as a response to foliage loss. Moreover, the colonization percentage of thick-mantled mycorrhizae correlated positively with the ratio of leaf to heterotrophic plant biomass in the defoliated birch seedlings, but not in the control ones. This tends to indicate a stronger carbon limitation of ectomycorrhizal colonization in defoliated seedlings.

If herbivory is unevenly distributed among different types of plants, or if individual plants differ in their response to herbivory, herbivores may affect seedling growth and survival, and consequently plant population structure. In this study we examined variation in resistance to insect herbivory and in growth responses to feeding among 20 silver birch (Betula pendula Roth) genotypes representing variation within a natural population. Birch seedlings were grown in dense stands in random arrangement so that seedlings of different genotypes competed with each other. On insect exposure plots natural colonization of insects was allowed, and insect removal plots were sprayed with insecticide. Resistance to insect herbivory was measured as the inverse of leaf damage, and growth responses of seedlings to feeding were determined as the change in seedling height relative to the amount of damage. Resistance varied significantly among genotypes, but growth responses to feeding did not. In fertilized seedlings, resistance correlated negatively with height, indicating a trade-off between resistance and growth. The absence of such a correlation in non-fertilized seedlings is a sign of environmental effects on the cost of resistance. Growth responses to feeding did not correlate with either resistance or growth. Nevertheless, different effects of the actual damage on height increase changed the positions of the genotypes in the size hierarchy of the experimental populations. Thus, even moderate levels of insect herbivory can change the outcome of competitive interactions between birch genotypes.

Leaf maturation in mountain birch (Betula pubescens ssp. czerepanovii) is characterized by rapid shifts in the types of dominant phenolics: from carbon-economic flavonoids aglycons in flushing leaves, via hydrolysable tannins and flavonoid glycosides, to carbon-rich proanthocyanidins (condensed tannins) in mature foliage. This shift accords with the suggested trade-offs between carbon allocation to plant defense and growth, but may also relate to the simultaneous decline in nutritive leaf traits, such as water, proteins and sugars, which potentially limit insect growth. To elucidate how birch leaf quality translates into insect growth, I introduce a simple model that takes into account defensive compounds but also acknowledges insect demand for nutritive compounds. The effects of defensive compounds on insect growth depend strongly on background variation in nutritive leaf traits: compensatory feeding on low nutritive diets increases the intake of defensive compounds, and the availability of growth-limiting nutritive compounds may modify the effects of defenses. The ratio of consumption to larval growth (both in dry mass) increases very rapidly with leaf maturation: from 2.9 to 9.8 over 2 weeks in June-July, and to 15 by August. High concentrations in mature birch leaves of "quantitative" defenses, such as proanthocyanidins (15-20% of dry mass), presumably prevent further consumption. If the same compounds had also protected half-grown leaves (which supported the same larval growth with only one third of the dry matter consumption of older leaves), the same intake of proanthocyanidins would have demanded improbably high concentrations (close to 50%) in young leaves. The model thus suggests an adaptive explanation for the high levels of "quantitative" defenses, such as proanthocyanidins, in low-nutritive but not in high-nutritive leaves because of the behavioral responses of insect feeding to leaf nutritive levels.

Hemiparasitic angiosperms concentrate nutrients in their leaves and also produce high quality litter, which can decompose faster and release more nutrients than that of surrounding species. The impact of these litters on plant growth may be particularly important in nutrient-poor communities where hemiparisites can be abundant, such as the sub-Arctic. We tested the hypothesis that plant growth is enhanced by the litter of the hemiparasite Bartsia alpina, in comparison with litter of co-occurring dwarf shrub species, using a pot based bioassay approach. Growth of Betula nana and Poa alpina was up to 51% and 41% greater, respectively, in the presence of Bartsia alpina litter than when grown with dwarf shrub litter (Vaccinium uliginosum, Betula nana and Empetrum nigrum subsp. hermaphroditum). The nutrient concentrations of Betula nana plants grown with Bartsia alpina litter were almost double those of plants grown with dwarf shrub litter, and a significantly greater proportion of biomass was allocated to shoots rather than roots, strongly suggesting that nutrient availability was higher where Bartsia alpina litter was present. The presence of litter from dwarf shrubs, or the moss Hylocomium splendens, did not reduce the positive effect of Bartsia alpina litter on plant growth. E. nigrum litter did not appear to affect plant growth substantially differently from litter of other dwarf shrub species, despite earlier reports of its allelopathic action. The enhanced nutrient uptake and growth of plants in the presence of Bartsia alpina (and potentially other hemiparasitic species) litter could have important implications for communities in which it occurs, including enhanced survival of seedlings of co-occurring species and increased resource patchiness.