Age and weather effects on between and within ring variations of number,width and coarseness of tracheids and radial growth of young Norway spruce

Annual growth, fibre and wood properties of Norway spruce are all under strong influence from genetics, age and weather. They change dynamically, particularly at young ages. Most genetic research and tree improvement programs are based on data from this most dynamic phase of the life of trees, affected by differences in weather among sites and years. In the work presented, influences of age and weather were investigated and modelled at the detail of annual rings and at the sub-tree ring level of earlywood, transitionwood and latewood. The data used were analysed from increment cores sampled at age 21 years from almost 6000 Norway spruce trees of known genetic origin, grown on two sites in southern Sweden. The traits under investigation were radial growth, cell widths, cell numbers, cell wall thickness and coarseness as a measure of biomass allocation at cell level. General additive mixed models (GAMMs) were fitted to model the influences of age, local temperature and precipitation. The best models were obtained for number of tracheids formed per year, ring width, average radial tracheid width in earlywood, and ring averages for tangential tracheid width and coarseness. Considering the many sources behind the huge variation, the explained part of the variability was high. For all traits, models were developed using both total tree age and cambial age (ring number) to express age. Comparisons indicate that the number of cell divisions and ring width are under stronger control of tree age, but the other traits under stronger control of cambial age. The models provide a basis to refine data prior to genetic evaluations by compensating for estimated differences between sites and years related to age and weather rather than genetics. Other expected applications are to predict performance of genotypes in relation to site or climate and simulation of climate change scenarios.     

Histological response of resistant and susceptible white spruce to simulated white pine weevil damage

The traumatic wound response of families of white spruce, Picea glauca (Moench) Voss, resistant or susceptible to the white pine weevil, Pissodes strobi (Peck), were compared after simulated weevil damage. Leaders from 331 trees were wounded just below the apical bud in the spring, coinciding with the natural time of weevil oviposition. A portable 1-mm diameter drill was used to drill 24 holes per leader. Leaders were removed in the fall and examined for evidence of traumatic resin canal formation. Drilled trees had a traumatic wound response 8 times greater than that of undrilled trees; however, undrilled trees also formed some resin canals in response to unknown causes. In the drilled trees, the traumatic wound response extended into the lower part of the leader, where it could possibly affect older larvae. Trees from resistant families responded with greater intensity than trees from susceptible families, by producing multiple rings of traumatic resin canals. Trees from resistant families also responded more rapidly than trees from susceptible families based on number of cells to the first ring of traumatic resin canals. Trees from some resistant families exhibited no traumatic resin canal formation, showing considerable within-family variation and suggesting that other resistance mechanisms might be important. In the year after drilling, there was a reduction in tree diameter growth and trees suffered a reduction in constitutive resin canals in the bark, which suggests some energetic cost of traumatic resin production. There was no indication that the extent of constitutive defenses, as measured by density of cortical resin canals before wounding, was related to the ability to produce traumatic resin canals. Screening trees based on their capacity to produce traumatic resin canals may be useful in selecting genotypes resistant to white pine weevil.     

Fine root morphological adaptations in Scots pine, Norway spruce and silver birch along a latitudinal gradient in boreal forests

Variability in short root morphology of the three main tree species of Europe's boreal forest (Norway spruce (Picea abies L. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth)) was investigated in four stands along a latitudinal gradient from northern Finland to southern Estonia. Silver birch and Scots pine were present in three stands and Norway spruce was present in all stands. For three fertile Norway spruce stands, fine root biomass and number of root tips per stand area or unit basal area were assessed from north to south. Principal component analysis indicated that short root morphology was significantly affected by tree species and site, which together explained 34.7% of the total variability. The range of variation in mean specific root area (SRA) was 51-74, 60-70 and 84-124 m(2) kg(-1) for Norway spruce, Scots pine and silver birch, respectively, and the corresponding ranges for specific root length were 37-47, 40-48 and 87-97 m g(-1). The range of variation in root tissue density of Norway spruce, Scots pine and silver birch was 113-182, 127-158 and 81-156 kg m(-3), respectively. Sensitivity of short root morphology to site conditions decreased in the order: Norway spruce > silver birch > Scots pine. Short root SRA increased with site fertility in all species. In Norway spruce, fine root biomass and number of root tips per m(2) decreased from north to south. The differences in morphological parameters among sites were significant but smaller than the site differences in fine root biomass and number of root tips.     

漆树科4种植物次生韧皮部的解剖比较

【目的】以漆树科3属4个种(盐肤木、青麸杨、黄连木和黄栌)的次生韧皮部为研究对象,阐明各物种次生韧皮部组成细胞的结构特征和分布规律,为植物次生韧皮部组成细胞结构特征与分泌结构之间的关系研究提供理论依据,并为漆树科植物系统演化的相关研究提供参考依据。【方法】采集盐肤木、青麸杨、黄连木和黄栌的树皮样品(样品含形成层、次生韧皮部及周皮,不含次生木质部),采用比较解剖学的方法,对漆树科4种植物的次生韧皮部的结构特征、组成细胞的构成及其排列方式进行比较,并对4种植物的乳汁道结构进行研究。【结果】漆树科4种植物次生韧皮部均是由筛管分子和伴胞、韧皮薄壁组织细胞、射线细胞、乳汁道和韧皮纤维构成,呈切向带状相间排列。黄连木、青麸杨和盐肤木的具功能韧皮部和无功能韧皮部分界明显;青麸杨的射线细胞在无功能韧皮部中倾斜分布,其余3种植物均为整齐纵向排列;青麸杨的乳汁道为分枝型乳汁道,其他3个种的乳汁道均无分枝,仅有少量相邻乳汁道融合现象发生;黄连木和黄栌次生韧皮部内分布有大量的韧皮纤维,青麸杨韧皮部中较少,而盐肤木韧皮部中无韧皮纤维分布。【结论】漆树科4种植物次生韧皮部组成细胞及乳汁道结构存在一定差异,青麸杨乳汁道是目前为止漆树科植物中发现的较为稀有的分枝型乳汁道,研究结果能够为漆树科系统进化研究提供参考依据。     

Assessing the impacts of climate change and nitrogen deposition on Norway spruce (Picea abies L. Karst) growth in Austria with BIOME-BGC

The aim of this paper is to determine whether a detectable impact of climate change is apparent in Austrian forests. In regions of complex terrain such as most of Austria, climatic trends over the past 50 years show marked geographic variability. As climate is one of the key drivers of forest growth, a comparison of growth characteristics between regions with different trends in temperature and precipitation can give insights into the impact of climatic change on forests. This study uses data from several hundred climate recording stations, interpolated to measurement sites of the Austrian National Forest Inventory (NFI). Austria as a whole shows a warming trend over the past 50 years and little overall change in precipitation. The warming trends, however, vary considerably across certain regions and regional precipitation trends vary widely in both directions, which cancel out on the national scale These differences allow the delineation of 'climatic change zones' with internally consistent climatic trends that differ from other zones. This study applies the species-specific adaptation of the biogeochemical model BIOME-BGC to Norway spruce (Picea abies (L.) Karst) across a range of Austrian climatic change zones, using input data from a number of national databases. The relative influence of extant climate change on forest growth is quantified, and compared with the far greater impact of non-climatic factors. At the national scale, climate change is found to have negligible effect on Norway spruce productivity, due in part to opposing effects at the regional level. The magnitudes of the modeled non-climatic influences on aboveground woody biomass increment increases are consistent with previously reported values of 20-40 kg of added stem carbon sequestration per kilogram of additional nitrogen deposition, while climate responses are of a magnitude difficult to detect in NFI data.     

Xylem dysfunction in Yezo spruce (Picea jezoensis) after inoculation with the blue‐stain fungus Ceratocystis polonica

The blue‐stain fungus Ceratocystis polonica is pathogenic to Norway spruce (Picea abies) in Europe, as well as to Yezo spruce (Picea jezoensis) and Sachalin spruce (Picea glehnii) in Japan. The wilting mechanism in P. jezoensis saplings after inoculation with C. polonica was examined based on anatomical studies of the phloem and xylem of periodically harvested trees. In addition, the course of sap ascent in the trunks was traced by injection of acid fuchsin solution at harvest. As an initial external symptom, needle discolouration was observed. In dye conduction tests, xylem dysfunction in the xylem of inoculated trees became obvious. The dehydrated xylem area (dry zone) had extended more than 20 cm above the inoculation wounds, within 1 month after inoculation. When the sap flow to the branches had nearly stopped, the leaves began to discolour. Hyphae of C. polonica colonized the ray tissue around the inoculation wounds, but were absent at the front of the dry zones. Defence reactions occurred in ray parenchyma cells adjacent to the penetrating hyphae. It is suggested that secondary metabolites, which are formed by the ray cells and epithelial cells of resin canals, are involved in the obstruction of sap flow. Limited necrotic lesions of the phloem and cambium were not associated with foliar symptoms. It is proposed that the dry zone formation caused by C. polonica is the main mechanism leading to tree death.     

Methyl jasmonate treatment of mature Norway spruce (Picea abies) trees increases the accumulation of terpenoid resin components and protects against infection by Ceratocystis polonica, a bark beetle-associated fungus

When conifers such as Picea abies Karst. (Norway spruce) are attacked by insects or pathogens, they often respond by producing increased quantities of terpenoid oleoresin. This response can be mimicked in young P. abies seedlings by treatment with methyl jasmonate (MJ). In this study, we determined the effects of MJ on terpenoids and other chemical defenses of mature P. abies, and investigated whether this treatment protected trees against attack by the blue-stain fungus Ceratocystis polonica (Siem.) C. Moreau, the most important fungal associate of the spruce bark beetle Ips typographus L. Methyl jasmonate treatment induced the formation of traumatic resin ducts in the developing xylem, enhanced resin flow and stimulated increased accumulation of monoterpenes, sesquiterpenes and diterpene resin acids. However, only minor changes were detected in terpene composition in response to MJ treatment and no changes in soluble phenolic concentration were measured. There was much variability in the timing and degree of response to MJ among clones. The observed chemical and anatomical changes in response to MJ treatment were correlated with increased resistance to C. polonica, suggesting that terpenoid oleoresin may function in defense against this pathogen.     

Microsite patterns of conifer seedling establishment and growth in a mixed stand in the southern Alps

In many forests of the Alps, permanent forest cover and, therefore, its continuous renewal is the main silvicultural goal. Regenerating these forests must be based on a sound understanding of the ecology of the tree species in question. The regeneration of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) in the upper montane zone of the southern central Alps has so far received little attention. The aim of this study was, therefore, to characterize the ecological niche for the establishment and growth of silver fir and Norway spruce seedlings in this zone. The study was conducted as a case study on a plot of 1.5 ha in the upper montane zone of southern Switzerland, at an altitude of 1380 m. The basal area of the mixed Norway spruce–silver fir stand was 43.7 m² ha⁻¹. Seedlings were censused on 375 plots of 1.0 m² area. Seedling density was 1.30 ± 4.25 m⁻² (mean ± standard deviation) for Norway spruce and 0.87 ± 1.50 m⁻² for silver fir. Logistic regression models were used to test the effect of microsite characteristics on seedling occurrence, and general linear models for effects on seedling height growth and biomass increment. Most seedlings received less than 10% light (photosynthetic photon flux density) as compared to values in the open. Silver fir occurrence was positively related to microsites at the edge of canopy trees, but unrelated to ground cover type, light and micro-relief. Norway spruce occurrence was only, and positively, related to the presence of mosses. Height growth and biomass increment of seedlings of both species were only loosely correlated with microsite conditions. Seedlings without canopy cover grew faster than those under canopy cover, probably as a result of light and moisture limitations under the canopy of adult trees. Diffuse radiation was positively correlated with average annual biomass increment of silver fir, but not of Norway spruce seedlings. In general, the results suggest that silver fir seedlings have less specific microsite requirements than Norway spruce seedlings in terms of ground cover. They are also more shade tolerant, and therefore, grow faster than Norway spruce in low-light environments of the upper montane zone of the southern Alps. While the microsite concept can be helpful in designing silvicultural operations, it has limitations when only patch characteristics that are easy to assess are used, and others neglected. Finally, our study suggests that counting the number of green shoots is a promising method to quickly and non-destructively estimate the biomass of a great number of small seedlings.     

Between- and within-site variation of density and bending properties of Picea abies structural timber from Norway

This study provides an analysis on the variability of structural timber of Norway spruce (Picea abies) grown in Norway. Density, modulus of elasticity (MOE) and bending strength were measured on 1188 boards from 205 trees, sampled from 14 sites throughout Southern Norway, Eastern Norway and Trøndelag. The area represents the procurement area for the majority of Norwegian sawmills. The variability of the timber properties was analysed in a linear mixed model where the random variance was divided into variance due to site, variance due to trees and within-tree variance. Models describing variance due to site based on site index, altitude and latitude were developed, and combined with data from the Norwegian National Forest Inventory to estimate mean values and variability of the timber properties. The results showed that major parts of the variance due to site are explained by altitude and site index, and for density also by latitude. Major parts of the variance due to site and the variance due to trees in bending strength and MOE were explained by density.     

Genetic variation in foliar nutrient concentration in relation to foliar carbon isotope composition and tree growth with clones of the F1 hybrid between slash pine and Caribbean pine

The objectives of this study were: (1) to quantify the genetic variation in foliar nutrient concentration in relation to foliar carbon isotope composition (δ¹³C) and tree growth of 122 clones of ca. 4-year-old F₁ hybrids between slash pine (Pinus elliottii Engelm var. elliottii) and Caribbean pine (P. caribaea var. hondurensis Barr. et Golf.) grown at two experimental sites with different water and nutrient availability in southeast Queensland, Australia and (2) to examine the potential of using foliar nutrient concentration of the 4-year-old tree canopies for selecting elite F₁ hybrid pine clones with improved nutrient-use efficiency (NUE) and water-use efficiency (WUE), and ultimately enhanced tree growth under ambient growing conditions. There were significant differences in foliar nutrient concentrations between two canopy positions (upper outer and lower outer canopy) sampled, between summer and winter, and between the two sites. This highlights that foliar nutrient concentrations are influenced by sampling and environment. Significant genetic variations in foliar nutrient concentrations were detected between the clones, between the female parents, and between the male parents of the clones in both sampling seasons at both sites. Depending on the nutrient concerned, canopy position, season, and site sampled, the clones accounted for 4.7–33.9% of the total variation in foliar nutrient concentrations, the clone female parents for 0–25.1% and the clone male parents for 0–28.6%. The site-by-clone interactions were statistically significant for foliar N, P, Mg, Cu, Zn, Mn, Fe and mineral concentrations at the upper outer canopy in summer, and for foliar N concentration in winter. There were significant, positive correlations between clone means of foliar δ¹³C and N concentration at the upper outer canopy in summer for the wet site, while clone foliar δ¹³C was also positively related to clone foliar N concentration at both canopy positions in summer for the dry site. This suggests that clone WUE as reflected in foliar δ¹³C may be improved by selecting elite clones with higher foliar N concentration and increased photosynthesis, leading to enhanced tree growth when both water and N are the major growth-limiting factors. This is supported by the positive correlation detected between clone tree height and foliar N concentration at the upper outer canopy for both sites. Thus, foliar nutrient (particularly N) concentration, together with foliar δ¹³C, may be useful for assisting in selection of exotic pine clones with improved NUE and WUE, and enhanced tree growth under the nutrient- and water-limiting environments.     

Intra- and inter-provenance variability in phloem phenols of Picea abies and relationship to a bark beetle-associated fungus

One hundred Norway spruce (Picea abies (L.) Karst.) clones (three ramets per clone) were analyzed for phloem phenol composition and concentration before and 10 days after wound inoculation with sterile malt agar. Fifty clones (Experiment 1) belonged to the same provenance, whereas the remaining clones (Experiment 2) belonged to five provenances from three geographic areas. In Experiment 2, two additional ramets from the same clones were mass inoculated with Ceratocystis polonica (Siem.) C. Moreau to quantify the resistance of each clone. Tree response to wound inoculations was characterized by increased catechin concentration in both experiments, accompanied by increases in astringin and decreases in piceid in Experiment 1. In both experiments, we observed a diverse group of phenolic compounds whose concentrations increased (catechin, astringin) or did not vary (taxifolin glucoside) in response to wound inoculations, whereas concentrations of a homogeneous group of stilbene compounds decreased (piceid) or did not vary (isorhapontin, unidentified stilbene). In Experiment 2, provenances from the alpine and Hercynian-Carpatic areas differed from provenances from the Baltic area with respect to the relative importance of these two groups of compounds, further indicating that the two groupings of phenolic compounds structure the Norway spruce populations. Eighty days after mass inoculation, the percentage of healthy sapwood, which was taken as a measure of tree resistance, indicated that clones from the Baltic area were less resistant to mass inoculations than clones from the alpine and Hercynian-Carpatic areas. We conclude that the degree of resistance of Norway spruce trees to mass inoculations with a bark beetle-associated fungus can be predicted based on the diversity of constitutive phloem phenols and the ability to induce phenol synthesis in response to wounding.     

Variability of wood properties in two wild cherry clonal trials

There has been an increase in wild cherry cultivation over recent decades, revealing a need for improvements in planting material to be used for the production of high quality timber. To date, trial tests have been carried out mainly on growth and tree architectural traits, while no investigations have focused on wood properties. The present study investigated the variability of the growth traits and the physical and mechanical properties of wild cherry wood, both within the tree, within and among clones and between sites. The aims of the study are to provide useful information on the genetics of wood traits in valuable hardwoods and to discuss how within-tree variation can affect early selection of genotypes. The results suggest that site was the most important source of variation in growth traits and in the proportion of heartwood, although differences among clones were highly significant. The number of sapwood rings was very homogeneous both between sites and among clones. Concerning the wood traits, most variation was detected within the tree, rather than within or among clones or between sites. Within tree variability seems to be mainly due to heartwood/sapwood presence. However, since there is a high correlation between heartwood and sapwood properties, a quite efficient selection made on the basis of sapwood should provide an efficient means of selecting heartwood traits.     

Genetic parameters of growth and wood quality traits in Picea abies

Genetic parameters were estimated for wood and growth traits in two 19-yr-old clonal trials and a 40-yr-old full-sib progeny trial of Norway spruce [Picea abies (L.) Karst.]. In the clonal trials high (>0.4) broad-sense heritabilities were found for wood density traits, lignin content, number of internal cracks, growth traits, spiral grain and number of resin canals. Moderate (0.2–0.4) heritabilities were found for tracheid lumen diameter and cell wall thickness, microfibril angle and tracheid length, while low heritabilities (<0.2) were found for pulp yield, fibre strength, wood stiffness and wood colour. Lignin content and pulp yield showed low genetic variation, whereas the genotypic coefficient of variation for most other traits ranged between 5 and 15%. Most traits showed low levels of genotype by environment interaction. Among the wood properties, latewood proportion, earlywood density and ring density showed significant, adverse correlations with volume in both clonal trials.     

Initiation and development of resin ducts in the major organs of Pinus massoniana

The structure, distribution and patterns of resin ducts in processes of its initiation and development were studied with the methods of thin section and ultrathin section. This paper emphasized the ultrastructural changes during canal development by a ring of the live epithelial cells, and the epithelial cells were usually surrounded with one or two layered sheath cells, which were normal parenchyma cells in some primary resin ducts and became dead cells with thick walls in other primary and secondary resin ducts. The resin ducts were found to occur in almost all organs, except in cotyledon. The resin ducts were formed by schizogeny and their development can be divided into three stages (e.g., initial stage, formation stage and mature stage). At the initial stage, the initial cells had many plastids without integral membrane structures, which contain one or two starch grains in them, and there are a few black osmiophilic droplets on the endoplasmic reticulum and membranes. A small number of osmiophilic droplets were present in the plastids. At the formation stage, the number of plastids, mitochondria and Golgi bodies in epithelial cells increased. The plastids were commonly surrounded by endoplasmic reticulum sheath. The larger osmiophilic droplets in cytoplasm and the smaller osmiophilic droplets on the plastids envelope, mitochondrion envelope and Golgi vesicles obviously increased in number during canal developing. At the mature stage, the cytoplasm of epithelial cells became thin with small nucleus. The number of mitochondria and Golgi body decreased, but numerous plastids still existed. Osmiophilic droplets were abundant in epithelial cells as in previous status. Taken together, the structures of plastids in epithelial cells gradually became well developed and the synthesis of resin was remarkably enhanced during resin duct formation and plastids should be the main site for resin synthesis.     

Nutrient accumulation and role of atmospheric deposition in coniferous stands

Three stand types on drained wetlands, all 31 years old, were studied. The stands were: (1) Scots pine, unfertilized; (2) Scots pine, fertilized; and (3) Norway spruce, fertilized. Amounts of nutrients (N, K, Ca, Mg, P, S, B, Fe, Mn, Zn, Cu) in above-ground biomass for all three stand types could be simulated precisely by a curvilinear regression model, with stand volume on bark as regressor. Net H⁺ production of the fertilized pine was estimated to be 661 mol H⁺ ha⁻¹ year⁻¹ from establishment to 31 years of age. The corresponding value for spruce was 1232 mol H⁺ ha⁻¹ year⁻¹. Atmospheric inputs to the pine and spruce sites were 695 and 516 mol H⁺ ha⁻¹ year⁻¹, respectively. Atmospheric input of N was 2.3 and 1.3 times the accumulation in the biomass of unfertilized and fertilized pine, whereas the value for spruce was 0.7. The corresponding ratios for S were 43, 19, and 11.     

不同产脂量云南松树脂道解剖学比较研究

为确定高产脂云南松的早期选择指标,采用石蜡切片法对不同海拔、不同产脂量云南松的针叶、树干韧皮部和木质部的组成性树脂道特征进行解剖观察,结果表明:高产脂与低产脂云南松的针叶树脂道个数、树干木质部树脂道密度、韧皮部树脂道大小及单位长度树脂道数量存在极显著差异(P＜0.01),但不同海拔之间无显著差异;高、低海拔高产脂云南松针叶树脂道数量分别比低产脂的高38.89％、30.34％,树干木质部树脂道密度分别高58.06％、48.48％,韧皮部树脂道大小分别高32.20％、29.73％,韧皮部单位长度树脂道数量分别高21.95％、38.51％.说明针叶树脂道数量、树干木质部树脂道密度及韧皮部树脂道大小3项指标,也可作为云南松产脂力评价的部分指标.     

Seedling emergence in uneven-aged Norway spruce stands in Finland

The emergence of Norway spruce seedlings on different types of germination sites in uneven-aged Norway spruce dominated stands four years after selection harvesting was studied. The data were collected from 20 stands in southern Finland. In each stand, ground cover was measured from a set of 64 one square metre subplots on a permanent experimental plot of 1600?m². Correlation analysis, direct comparison of subplots with and without seedlings and mixed-effects models were used to estimate the influence of ground and tree cover on seedling occurrence. Grass cover, dwarf shrub cover, cover of certain moss species and close vicinity of trees decreased the occurrence probability and number of seedlings, while greater stand and local basal area around the subplots increased the seedling number. The cover of vegetation free surfaces, usually considered as good germination sites, was rather low. Variation in seedling emergence between plots was high. We concluded that spruce regeneration was slightly more abundant in stands with greater overall stand density and local density within a stand in postharvest conditions.     

Prediction of individual tree growth in managed stands of mixed Picea abies (L) Karst,and Betula pendula Roth & Betula pubescens Ehrh.

The individual tree growth in stands of mixed Norway spruce (Picea abies (L.) Karst.) and birch (Betula pendula Roth & Betula pubescens Ehrh.) is estimated using basal area and height growth functions for each species separately. The individual tree growth models are distant dependent and the number, size and proximity of neighbours are expressed as size‐ratio competition indices. The competition indices were calculated using a basal area factor gauge to define competitors. The tree growth functions are based on data from nine mixed stands of Norway spruce and birch. The recursive multivariate regression approach is used. The growth functions have standard deviation about the function/standard deviation about the mean (sf/sm) values between 31 and 61% and the evaluations made with root mean square error (rmse) give estimates which vary between 8 and 45 % of the observed mean value. These values are comparable with the precision reported in other studies. In the present investigation the distant dependent indices are important independent variables. It seems suitable to describe the change in growth conditions from retarded to released growth by means of size ratio competition indices. For birch, a positive effect on growth is obtained the more the total competition consists of Norway spruce. For Norway spruce a negative effect on growth is obtained the more the total competition consists of Norway spruce. The lower competitors have a positive effect on the growth of the spruce trees. The interpretation should be that it is better to have a small competitor than a larger one, not that small competitors as such have a positive effect on growth of Norway spruce.     

Intersterilitätsgruppen und Klone von Heterobasidion annosum-Isolaten aus Koniferen- Wurzel- und -Stammfäulen

Intersterility groups and clones of Heterobasidion annosum isolates from root and butt rots of conifers 69 H. annousum heterokaryons from scots pine, Douglas fir, Norway spruce and larch belonged to the P group, three heterokaryons from Norway spruce to the S group. The results of a study with clones of H. annosum isolates from roots of Scotch pine trees in close neighbourhood suggest colonisation by H. annosum (partly) via root contacts.