Pine weevil (Hylobius abietis) damage to cuttings and seedlings of Norway spruce

Damage caused by pine weevil (Hylobius abietus L.) to planted seedlings and cuttings of Norway spruce (Picea abies (L.) Karst.) was studied at five clearcut sites in south-eastern Sweden. The main objective was to compare the two types of stock in terms of attack frequency and mortality due to pine weevil feeding. Cuttings and seedlings with the same initial stem-base diameter (4 mm) were compared. Two sites were harvested and scarified shortly before planting, two were harvested shortly before planting, but were not scarified, and one was harvested 2 years before and scarified the autumn before planting. The total mortality 5 years after planting was highest, greater than 90%, at the new, non-scarified sites, and lowest, 23%, at the old, scarified site. More than 90% of the mortality was caused by pine weevil feeding. Attack frequency and pine weevil induced mortality were significantly higher among seedlings than among cuttings. Mortality due to pine weevil damage was 4–43% higher in seedlings than in cuttings after the fifth year. Of the cuttings and seedlings that were attacked in the first year, a significantly higher frequency of the seedlings were girdled. The higher resistance of cuttings to pine weevil damage may partly explain the more rapid growth of cuttings reported in other studies. However, the causes of their higher resistance need to be further investigated. The thicker bark and needles on the stem base of the cuttings could be important in this respect.     

Growth of Mixed Birch-Coniferous Stands in Relation to Pure Coniferous Stands at Similar Sites in South-eastern Norway

Mean age, mean and top heights and yield were studied in 20 mixed stands of birch ( Betula pubescens Ehrh. and B. pendula Roth) and Picea abies (L.) Karst. and nine mixed stands of birch and Pinus sylvestris L. in south-eastern Norway. Each mixed stand and the adjacent pure coniferous stand (control) were growing under the same site conditions and had not been commercially thinned. There were no significant differences in mean age at breast height or in top heights between birch and conifers in the mixed stands, while mean height was significantly higher for birch than for spruce. A growth index was calculated based on total volume and age at breast height. For the spruce sites the growth in young mixed stands (birch < 17 m) was superior to that of pure spruce, while the difference was insignificant in older stands. The growth index correlated positively with the ratio between generatively and vegetatively regenerated birch trees, and negatively with the age of the oldest species in the mixture and with site quality. For the pine sites there was no significant difference in the growth index between mixed birch-pine and pure pine stands. A mixture effect of birch on the volume yield of spruce or pine could not be demonstrated.     

Temporal variation and efficiency of leaf area index in young mountain Norway spruce stand

Temporal variation of leaf area index (LAI) in two young Norway spruce stands with different densities was monitored during eight consecutive growing seasons (1998–2005). We focused on: (1) LAI dynamics and above-ground mass production of both spruce stands and their comparison, (2) leaf area duration (LADU), crop production index (CPI) and leaf area efficiency (LAE) evaluation, and (3) thinning impact on the above-mentioned parameters. Also, we tried to deduce the most effective LAI value for the Norway spruce forest investigated. The LAI values of both spruce stands showed a typical seasonal course. To describe the LAI dynamics of the stand, we recommend taking LAI measurements within short time intervals at the time of budding and needle expansion growth (i.e., in early spring) and close to the LAI peak, when the twig growth has been completed. The reason was that after reaching the seasonal maximum, no significant differences between subsequently obtained values were found in the following 2 months. Therefore, we recommend this period for the estimation of seasonally representative LAI values, enabling the comparison of various spruce stands. The maximum hemi-surface LAI value reached 12.4. Based on our results, the most effective LAI values for maximum above-ground biomass production were within the range of 10–11. We found an LAI over these values to be less effective for additional production of above-ground biomass. In forest practice, thinning intensity is mostly described by percentage of stocking reduction. We want to show that not only thinning intensity, but also the type of thinning is important information. The type of thinning significantly affected the stand above-ground biomass increment, canopy openness, stand LAI and LAI efficiency. The stimulating effect of high-type thinning was observed; the LAE as well as the CPI increased. Low-type thinning had no such effects on LAE increments compared to the high-type thinning with similar intensity.

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Application of random amplified polymorphic DNA (RAPD) to detect genetic variation in Norway spruce

The RAPD assay is a screening method for genetic variation; in Norway spruce biology research, it may find many applications. We have investigated into its suitability for Norway spruce genetics by optimizing the protocol, testing for stability in clones, especially tissue culture clones, assessing the variation in a seed sample, and checking for correct Mendelian segregation in haploid megagametophytes. The RAPD assay produced numerous genetic markers quickly. The data obtained gave insight into the genetic make-up of clones and seed.

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Zusammenfassung Die RAPD Methode, ein Schnelltest für die genetische Variation, wurde für die europäische Fichte getestet, wo viele Aspekte der Biologie der Baumart damit untersucht werden könnten. Das Protokoll wurde optimiert, und genetische Stabilität in Klonen, besonders aus Gewebekultur, die genetische Variation in einer Samenprobe und die Mendel-konforme Aufspaltung in haploiden Megagametophyten wurde untersucht. Zahlreiche genetische Marker konnten gefunden werden, wodurch ein Einblick in die genetische Zusammensetzung des untersuchten Materials gegeben war.

     

Reduction of frost heaving of Norway spruce and Scots pine seedlings by planting in mounds or in humus

This study was carried out in northern Sweden to determine the effects of frost heaving on the establishment of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings in relation to planting methods. For this purpose, one year old containerised seedlings were planted on two sites and on two dates: during the spring (early planting) and during the fall (late planting). In each case, two planting depths (normal and deep planting) and four planting sites (in mineral soil in the depressions, in the scalp/trench area, on the top of the mound and in the untreated humus layer) were used. On each site, 50 seedlings were planted for each treatment. Frost heaving was observed and measured during two years. The amount of heaving was highest in the hole and almost insignificant on the top of the mound and in the humus layer. Planting depth influenced the degree of heaving only for Scots pine planted in the hole and was not related to the planting time.     

Influence of gaps on some selected tree characteristics of edge trees in Norway spruce plantations

The present study examined branch and stem characteristics of trees growing around gaps in Norway spruce plantations. Trees located at the edge of gaps with a radius of either 5 m or 7 m either unplanted or with supplementary planting were compared to trees in the original closed plantation. The experiment was carried out in two locations in the south of Sweden and the design included four blocks at each location and one replicate of each treatment in each block. The measurements were carried out on standing and felled trees in 2005 at the time of the first thinning. The results indicate a significant effect of gaps and supplementary planting on the diameter of the largest branch, the number of living branches close to breast height, branch basal area, height to the first living branch and taper. The effect of supplementary planting on branch and stem characteristics was greater in large gaps than in small ones. This study indicated that supplementary planting may be useful in affecting stem-form and branch parameters, although other studies have shown that the trees that result from supplementary planting contribute little to total production.     

间伐对祁连山青海云杉人工林土壤水分的影响

利用EM50土壤水分监测仪,在样地尺度上,测定了祁连山青海云杉天然林、无间伐和间伐强度为20%的人工林地生长季节的土壤水分,对比分析间伐对人工林土壤水分的影响。结果表明:未间伐人工林林地表层(10 cm)土壤含水量显著高于间伐强度为20%的人工林和天然林,间伐导致了人工林林地表层土壤水分下降;而对于深层土壤含水量而言,间伐措施又显著提高了深层60 cm处的土壤含水量。与天然林地土壤含水量相比,无间伐人工林深层60 cm和80 cm处的土壤体积含水量仅为天然林的49.7%和52.1%,深层土壤已经出现旱化现象,间伐措施能够减缓这种旱化现象。     

Effect of thawing regime on growth and mortality of frozen-stored Norway spruce container seedlings planted in cold and warm soil

One-year-old frozen-stored Norway spruce (Picea abies (L.) Karst.) container seedlings were planted in a controlled environment providing an air temperature of 22°C and soil temperature of 9±1 or 18±1°C. At planting the root plugs were either frozen or had been thawing for 4 days at 9°C. During a 5-week growing period, in both cold and warm soil the root growth and height growth were less in frozen-planted seedlings than in thawed seedlings. In addition, frozen-planting delayed bud burst and increased mortality. Soil temperature, however, had no effect on bud burst or mortality. Low soil temperature retarded root growth of seedlings thawed before planting but resulted in both retarded root growth and height growth if root plugs were frozen when planted. These results indicate that planting Norway spruce seedlings with frozen root plugs constitutes a considerable risk for successful forest regeneration at soil temperatures normally prevailing in Fennoscandia in spring or early summer especially if the soil is dry at the time of the planting.     

Fibril angle variability in earlywood of Norway spruce using soft rot cavities and polarization confocal microscopy

The main purpose of this study was to investigate the variability of the fibril angle of tracheids in earlywood of Norway spruce (Picea abies L. Karst.). Polarization confocal microscopy was chosen and compared with the method utilizing the orientation of soft rot cavities. There was a significant correlation between the soft rot and polarization confocal microscopy methods, which showed the same trend of high fibril angles in the first part of the earlywood followed by a decrease toward the end of earlywood. This declining trend was less pronounced in annual rings containing compression wood. Moreover, large variations in fibril angle occurred between neighboring tracheids. The investigation also emphasized the differences between X-ray diffraction and microscopic methods, as the large variation seen by the latter methods is not seen by the X-ray diffraction approach because of its large area of measurement. No correlation was found between fiber morphology (i.e., average length, width, density) and the average fibril angle in the investigated annual rings.     

Effect of temperature and compression on the mechanical behavior of steam-treated wood

The mechanical behavior of steamed spruce wood changes dramatically with compression along the grain, the change being much more moderate perpendicular to the grain. The stiffness decrement due to increased temperature is greatest in the tangential material direction. The stiffness decrement due to compression is greatest along the grain. Compression to 80% compressive strain at 131°C inverts the order of the material directions regarding stiffness, the stiffness being the least along the grain. Plastic strain due to compression is greater at higher temperatures. The compression-induced decrement of stiffness along the grain is greater at higher temperatures, but the off-axis decrement of stiffness is less at higher temperatures.     

Micromechanical properties of common yew (Taxus baccata) and Norway spruce (Picea abies) transition wood fibers subjected to longitudinal tension

The longitudinal modulus of elasticity of common yew is astonishingly low in light of its high raw density. At least this was found for specimens examined at the solid wood level and at the tissue level. However, to reveal if this low axial stiffness is also present at the cellular level, tensile tests were performed on individual yew fibers and on spruce fibers for reference. The results revealed a low stiffness and a high strain to fracture for yew when compared with spruce. This compliant behavior was ascribed to a relatively high microfibril angle of yew measured by X-ray scattering. It can be concluded that the high compliance of yew observed at higher hierarchical levels is obviously controlled by a structural feature present at the cell wall level. In future studies, the biomechanical function of this compliant behavior for the living yew tree would be of particular interest.     

Crystallinity of wood and the size of cellulose crystallites in Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis>)

X-ray diffraction was used to study variations in the crystallinity of wood and the average thickness and length of the crystallites of cellulose as a function of the number of the year ring in Norway spruce [Picea abies (L.) Karst.]. The crystallinity increased from ring 4 to ring 10 from the pith and was constant after ring 10. The crystallinity of mature wood was about 30% ± 5%. The average thickness and average length of the crystallites were 3.2 ± 0.1nm and 28 ± 2nm, respectively; and no systematic variation of these values with the number of the year ring was observed. The mean microfibril angle decreased near the pith but was constant in the mature wood.     

Differences in soil properties in adjacent stands of Scots pine, Norway spruce and silver birch in SW Sweden

Soil properties were compared in adjacent 50-year-old Norway spruce, Scots pine and silver birch stands growing on similar soils in south-west Sweden. The effects of tree species were most apparent in the humus layer and decreased with soil depth. At 20-30 cm depth in the mineral soil, species differences in soil properties were small and mostly not significant. Soil C, N, K, Ca, Mg, and Na content, pH, base saturation and fine root biomass all significantly differed between humus layers of different species. Since the climate, parent material, land use history and soil type were similar, the differences can be ascribed to tree species. Spruce stands had the largest amounts of carbon stored down to 30 cm depth in mineral soil (7.3 kg C m⁻²), whereas birch stands, with the lowest production, smallest amount of litterfall and lowest C:N ratio in litter and humus, had the smallest carbon pool (4.1 kg C m⁻²), with pine intermediate (4.9 kg C m⁻²). Similarly, soil nitrogen pools amounted to 349, 269, and 240 g N m⁻² for spruce, pine, and birch stands, respectively. The humus layer in birch stands was thin and mixed with mineral soil, and soil pH was highest in the birch stands. Spruce had the thickest humus layer with the lowest pH.     

Regeneration niches of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] Karst.) saplings in small canopy gaps in mixed mountain forests of the Bavarian Limestone Alps

Natural regeneration measurements are the main silvicultural objective in overaged protective forests of the Bavarian Limestone Alps. While manifold problems with these stands, especially the impact of browsing, are widely recognised, the regeneration niches of Norway spruce (Picea abies [L.] Karst.) are insufficiently known. The purpose of this study was to determine favourable combinations of site factors for the development of spruce in small, unfenced canopy gaps, located on Aposerido-Fagetum caricetosum albae forest sites. We recorded the occurrence of spruce saplings (as dependent variable) and of six site factors (as independent variables) on 480 0.5 m²-subplots. In addition, we estimated the coverage of six acid adapted plant species to determine correlations with the humus depth. A binary logistic regression analysis was used to predict the probability of the occurrence of a spruce sapling in dependency of the different site factors. Supported by other studies, we assumed that the supply of solar radiation was adequate for the sufficient regeneration of spruce within the canopy gaps. Other site factors significantly determined the regeneration niches of spruce saplings. More spruce saplings were found near hindrances and on rough surfaces than would be expected from a random occurrence of saplings. These microsite types may have characteristics, especially protection against snow gliding that promotes spruce establishment. A calculated “hindrance index”, which accounted for the number, and the distance of surrounding hindrances might be a good specific value to describe the influence of hindrances on steep slopes. The sapling establishment decreased on thin humus layers. Our assumption for the sites was that thick organic layers might represent a good seedbed for spruce. Decayed dead wood was scarce, but was exceedingly favoured by spruce saplings. Results obtained suggest that the natural regeneration establishment of spruce on steep slopes can be successfully influenced by site factors which inhibit the influence of snow gliding. According to a “positive microsite” concept, we recommend for artificial regeneration measurements with spruce, microsites close to hindrances (e.g. stumps, downed trees) and Vaccinium myrtillus as a predictor for thick, acid humus layers.     

The effects of Douglas-fir on tree-specific arthropod communities in mixed species stands with European beech and Norway spruce

The ecological effects of planting exotic Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] in Central Europe are still poorly understood. The aim of this study was to answer the question of whether Douglas-fir affects tree specific arthropod communities in different mature forest types (Douglas-fir, spruce and beech dominated) in Southern Germany. Therefore, arthropod communities of stem and tree crown strata of Douglas-fir and spruce (Picea abies L.) were sampled in the years 1999–2001 using arboreal photo-eclectors and flight interception traps. Statistical analysis was conducted for all species and focused on conifer specialists at three levels: (1) species diversity, (2) guild structure and (3) community structure. Within the stem stratum, species diversity was significantly higher on spruce than on Douglas-fir independent of year and stand composition. This could not be explained by a single feeding guild, rather by species changing strata during the vegetation period. In contrast, species diversity in tree crowns was approximately the same for both conifer species. However, communities in Douglas-fir crowns were conspicuously different from those in spruce crowns, especially in the Douglas-fir dominated stand type. While zoophagous insects exhibited higher activity on Douglas-fir in 2000, xylophagous beetles were more abundant on spruce in 2001. In European beech stands with widely spaced Douglas-fir trees, the site specific and broad-leaved tree related fauna might be maintained. In addition, Douglas-fir with its resource of Adelges cooleyi and crowns that overtop the broad-leaved tree canopy, offer additional resources for several aphidophagous and thermophile species.     

Compositional changes of forest-floor vegetation in young stands of Norway spruce as an effect of repeated fertilisation

Forestry practices that aim to increase biomass production may mitigate climate change through increased carbon sequestration and the potential of substituting fossil fuels with renewable biofuels. Fertilising young stands of Norway spruce in Sweden have shown to increase tree growth by more than 200%. Fertilisation, however, also has other effects on forest ecosystems. Here, we studied the response of the species composition of forest-floor vegetation to three different frequencies of fertilisation in young stands of Norway spruce. Fertiliser was applied every year, every second year or every third year. The total amount of N ranged from 425 kg ha⁻¹ to 625 kg ha⁻¹, in combination with P, K, Ca, Mg, S, Mn, Zi, B and Cu. The largest effects of the fertilisation were found among bryophytes and lichens, which lost substantial cover. Unexpectedly, Deschampsia flexuosa, commonly known to be favoured by fertilisation, was negatively affected. Species that increased in frequency were Oxalis acetosella, Brachythecium sp. and Plagiothecium sp. Decreased availability of light, as an indirect effect of fertilisation through increased tree canopy cover, was found to be the most important factor behind the change in species composition of vascular plants. The total cover of bryophytes, however, did not show any significant response to the changes in canopy cover, indicating that the effects seen in this group may be a result of more direct effects of the fertiliser. Few significant differences were found between the two most intensive fertilisation frequencies, although fertilisation every third year was often distinguished from both the control and the other fertilised treatments. Even though the effects at the stand level were substantial, the effects on biodiversity and function of ecosystems on a landscape or regional level need further investigation.     

The influence of different forest organic matter on the growth of one-year old planted Norway spruce seedlings in a greenhouse experiment

A greenhouse trial was carried out to improve the knowledge of how forest organic matter could be utilized in site preparation and the choice of planting spot for Picea abies (L.) Karst. One-year old Norway spruce seedlings were grown for 26 weeks in pots containing pure mineral soil, forest organic matter in three different states of decomposition and combined treatments where the organic matter was mixed with or placed on the surface of the mineral soil (volume proportions of organic:mineral of 1:2). Watering was adapted to the water retention characteristics of each growing medium in order to keep the soil water potential between -4 and -5 kPa in all pots.In its pure form as well as combined with mineral soil, the moderately decomposed mor organic matter gave a higher seedling dry weight increment than the corresponding growing media containing the more decomposed humus. Adding mor or humus to mineral soil increased the seedling dry weight increment by 30–140% and 10–40%, respectively. The largest seedlings grew in the pure mor and humus. Mixing these forms of organic matter with mineral soil appeared to reduce seedling growth as compared to placing them on the mineral soil surface. The increment increase compared to plain mineral soil mostly consisted of proleptic growth and was probably due to the improved nutrient availability of the growing media. Fresh organic matter (chopped needles and twigs) seemed to impair seedling root function and reduced the dry weight increment by 30–50% as compared with pure mineral soil.The results suggest that as long as the water and temperature requirements are fulfilled planting Norway spruce seedlings without scarification and/or gathering extra mor and humus to the planting position should improve seedling growth as compared to the growth following scarification. Unmixed fresh needles and twigs or fresh needles and twigs combined with plain mineral soil should be avoided when planting Norway spruce seedlings.     

Regeneration in experimental gaps of subalpine <Emphasis Type="Italic">Picea abies</Emphasis> forest in the Slovenian Alps

In 1998, we analysed regeneration success in four 20 year old medium-sized (600–800 m²) and four small experimental gaps (100–200 m²) in subalpine Norway spruce forest in Triglav National Park, Slovenia. We assessed the influence of site (depressions, slopes), gap size, and position within gap (centre, gap south, and north edge) on regeneration success and interactions between regeneration density, height, height increment, direct and diffuse light, ground vegetation, soil depth, thickness of humus horizons, presence of woody debris, and microrelief. Seedling density was higher in depressions, where more seedlings developed in central positions of gaps in both gap sizes, and in microsites with deeper soils, less competition from ground vegetation and convex microsites. On slopes, a higher seedling density was found in small gaps, and at the northern edge of gaps regardless of gap size. Here, seedling density was negatively influenced by diffuse radiation and positively by soil depth. The results suggest that regeneration should be initiated from medium to large gaps in depressions and small gaps on slopes. Development of established seedlings was enhanced by higher radiation levels on both sites, therefore favourable extension of the gaps would be lateral to northwards.     

Nutrient cycling and soil leaching in eighteen pure and mixed stands of beech (Fagus sylvatica) and spruce (Picea abies)

Studies on the combined effects of beech–spruce mixtures are very rare. Hence, forest nutrition (soil, foliage) and nutrient fluxes via throughfall and soil solution were measured in adjacent stands of pure spruce, mixed spruce–beech and pure beech on three nutrient rich sites (Flysch) and three nutrient poor sites (Molasse) over a 2-year period. At low deposition rates (highest throughfall fluxes: 17 kg N ha⁻¹ year⁻¹ and 5 kg S ha⁻¹ year⁻¹) there was hardly any linkage between nutrient inputs and outputs. Element outputs were rather driven by internal N (mineralization, nitrification) and S (net mineralization of organic S compounds, desorption of historically deposited S) sources. Nitrate and sulfate seepage losses of spruce–beech mixtures were higher than expected from the corresponding single-species stands due to an unfavorable combination of spruce-similar soil solution concentrations coupled with beech-similar water fluxes on Flysch, while most processes on Molasse showed linear responses. Our data show that nutrient leaching through the soil is not simply a “wash through” but is mediated by a complex set of reactions within the plant–soil system.     

Variation of defence-related metabolites in the foliage of adult beech and spruce: a conceptual approach to approximating traded-off carbon

To prevent carbon (C) loss to consumers, trees need to defend their primary production. The tree-internal conflict in resource allocation between growth and defence demands has been the subject of various hypotheses but still requires quantification. A conceptual approach to approximating the C amount dispensable in favour of primary production at the expense of defence is demonstrated which is based on nine defence-related metabolite groups. Quantification is exemplified at the level of sun and shade leaves of adult Fagus sylvatica and Picea abies trees, two species contrasting in foliage type, under oxidative stress as induced by ozone exposure. The difference between maximum and minimum metabolite levels sampled several times throughout four consecutive growing seasons were conceived as dispensable between growth and defence-related metabolism and expressed in proportion of the mean annual gross primary production (GPP) of the foliage. In both species, this proportion amounted to between 2 and 5% of GPP (on a molar C basis). Remaining uncertainties are discussed as concerning functional overlap of substances between growth and defence-related metabolism, estimated classification of metabolite turnover rates and “third-party” trade-offs across C demands. Given the complexity of plant defence, simplification is needed for modelling allocation trade-offs in plants. The presented conceptual approach meets this need in approximating C transfer capacities between competing physiological demands and in stimulating empirical assessments towards mechanistic understanding. This article belongs to the special issue: “Growth and defence of Norway spruce and European beech in pure and mixed stands”.