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.     

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.     

Predicting tree regeneration in <Emphasis Type="Italic">Picea abies</Emphasis> snag stands

A bark beetle (Ips typographus) infestation caused the death of almost all Norway spruce (Picea abies) trees in a mountain forest in the Swiss Alps. We developed a tree regeneration model, ‘RegSnag’ (=REGeneration in a SNAG stand), to project the future amount and height of tree regeneration in these snag stands. The model combines a height-class structured tree module with a microsite-based module of snag decay and ground-vegetation succession. Microsite-specific rates of germination, mortality and height growth were modelled for four tree species (Picea abies, Sorbus aucuparia, Acer pseudoplatanus and Betula pendula) in eight height classes (from seedlings to saplings 5 m tall) and on 26 microsite types (e.g. moss, grass). Model tests with independent field data from 8 years after the Picea die-back demonstrated that microsites had a considerable effect on the development of tree regeneration on both the montane and the subalpine level. With microsite-specific parameters, the height and frequency of Picea in each microsite could be simulated more accurately than without considering microsite effects (e.g. bias of 8 vs. 119 saplings ha⁻¹ on the montane level). Results of simulations 40 years into the future suggest that about 330–930 Picea saplings per ha out of those that germinated in 1994 and 1996 will reach a height of 5 m within 30–35 years after Picea die-back. This is due to differences in seed inflow and browsing intensities. Picea and not Betula or Sorbus trees will replace the current herbaceous vegetation in these snag stands.     

The onset of hazel wood formation in Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] Karst.) stems

Key message

Fungal infection was outlined as a potential reason for the onset of indented annual growth ring formation during the juvenile phase of hazel wood growth. Annual growth ring indentations resulted from the formation of disturbed zones which originated solely in close proximity to leaf traces.

Context

Hazel wood is an abnormal type of woody tissue that is formed as a result of exogenous stimuli that may trigger long-term responses in the cambium. Cambial responses produce anatomical alterations in the surrounding xylem tissue that can be observed as an indentation of annual growth rings. The chemical profiles of lignan hydroxymatairesinol may provide an indication of its possible role in the protection of a living tree against the spread of a fungal or microbial infection at the onset of indentation.

Aims

The objectives of this study were to reveal the anatomical differences in the altered woody tissue of Picea abies hazel wood at both the onset and the later stages of annual growth ring indentation and to determine the chemical profiles for hydroxymatairesinol upon elicitation by a fungal infection in the disturbed zones.

Methods

Light and scanning electron microscopy observations were carried out on radial, tangential, and cross sections of hazel wood zones separated from P. abies stems. Concentrations of hydroxymatairesinol were determined for both the disturbed zones and the non-indented zones using a gradient high-performance liquid chromatography.

Results

The formation of disturbed zones was accompanied by significant changes in both the direction and width of the tracheids which produced an abnormal formation of intertwined and twisted tracheids. Fungal hyphae, radial cell wall cracks, and unusually large cross-field pitting were all found in the tracheids of the disturbed zones.

Conclusion

The content of hydroxymatairesinol in the acetone extract determined from the disturbed zones was 3.4 times greater than that present in the non-disturbed tissues. By means of vascular dysfunction in the leaf traces, host trees responded to the fungal infection by plugging the lumens of conductive leaf trace tissue and filling the vascular pathway with polyphenolic compound deposits.

     

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.     

Optimum lifetime for Swedish <Emphasis Type="Italic">Picea abies</Emphasis> seed orchards

The optimum lifetime of Picea abies seed orchards in Sweden was calculated using a model which considers changes in seed yield over time, annual progress of genetic gain, establishment, management and seed costs, and both the monetary and genetic value of the seeds produced. The longer the seed orchard is active, the more genetically outdated the produced seeds will become, and thus their value will decline with time. A main scenario was constructed using the best available estimates for Swedish Norway spruce and was compared to various alternatives reflecting likely variations. The optimal lifetimes in all of these scenarios was in the order of 40 years. Sensitivity analyses showed that use of orchards for slightly more or less than their optimal lifetimes does not lead to large losses. A more effective tree breeding program could slightly reduce the optimal lifetimes. The optimal lifetime depends strongly on the relationship between monetary value and genetic gain. Thus, in cases where genetic progress is sufficiently rapid, the optimal lifetime may be less than 30 years.     

A non-linear model to predict crown recession of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] Karst.) in Austria

In this study, a non-linear model was developed that predicts the five-year change of height to crown base (HCB) of Norway spruce (Picea abies [L.] Karst.). Data were available from the Austrian National Forest Inventory and comprised 2,419 trees from 1,637 permanent sample plots measured during 1981 and 2002. The dynamic model explained 36% of the variation in the observed change of HCB. It is well behaved and meets biological expectations. Based on five independent data sets, the predictive ability of the new dynamic model was compared to an already existing static crown ratio model. For this comparison, the models were applied as follows: the new ∆ HCB model was used to predict the change in HCB directly. For the semi-dynamic method, the static model was applied at the end and at the start of the growth period to obtain two estimates of HCB. The difference of these two estimates was then added to the initial HCB. For the static method, the model was only applied at the end of the prediction period to obtain the new estimate of HCB. Except for one plot, the new ∆ HCB model yielded the smallest BIAS and the highest precision, followed by the semi-dynamic and the static method. Because the independent data sets cover a broad range of age classes and thinning regimes, the validation results also indicate that the new ∆ HCB is robust and the effect of stand management is adequately represented.     

Adaptation to common optimum in different populations of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> Karst.)

Responses of Norway spruce populations to climatic transfer, in terms of growth and survival, were analyzed on the basis of a provenance experiment derived from the international provenance test IUFRO 1964/1968. The experiment comprises a series of five trial plots situated at contrasting elevations ranging from 484 to 1,275 m a.s.l., with 11 provenances represented at all trial plots that were used for the analysis. Transfer rates were defined as differences in altitudes or climatic variables between the site of plantation and the site of origin. Optimal transfer rates and optimal climates for individual provenances were derived from quadratic response functions. Spruce provenances generally responded positively by height and volume growth to transfer into lower altitudes, i.e., warmer conditions with less precipitations. The analysis at the level of provenances showed that optimal transfer rates were consistently negatively correlated with the underlying environmental variables and optimal climates were consequently nearly the same for all provenances irrespective of the response traits and ecodistance variables. Stability indices based on joint regression analysis indicate that provenances from higher altitudes, colder and wetter climates tend to be more stable, whereas provenances from lower altitudes, drier and warmer sites are more responsive to site quality. However, the differences in the stability are small and stability indices were generally close to 1. The results indicate that populations in different climates remain adapted to a common optimum and the extent of local adaptation is quite limited. Possible explanations of this observation are briefly discussed.     

Effects of <Emphasis Type="Italic">Ips typographus</Emphasis> (L.) damage on litter quality and decomposition rates of Oriental Spruce [<Emphasis Type="Italic">Picea Orientalis</Emphasis> (L.) Link.] in Hatila Valley National Park,Turkey

This study investigated the effects of Ips typographus (L.) damage on initial litter quality parameters and subsequent decomposition rates of oriental spruce tree species [Picea orientalis (L.) Link]. The needle litter was collected from highly damaged, moderately damaged and control stands on two aspects (north and south) and two slope position (top and bottom) on each aspect. The litter was analyzed for initial total carbon, lignin and nutrient (nitrogen, phosphorus, potassium, calcium, magnesium and manganese) concentrations. The variability in nitrogen and calcium concentrations and ratios of C:N, lignin:N and lignin:Ca was significantly affected by the insect damaged levels. While nitrogen concentrations in needle litter increased with increasing insect damage (and consequently the ratios of C:N and lignin:N decreased), calcium concentrations decreased (and consequently the ratio of lignin:Ca increased). Aspect and slope positions explained most of the variability in carbon, lignin, phosphorus, potassium, magnesium and manganese concentrations and lignin:P ratio between all studied stands. Litter decomposition was studied in the field using the litterbag technique. The litter from highly damaged stands showed highest decomposition rates followed by moderately damaged and control stands. The mass loss rates were significantly positively correlated with initial nitrogen concentration and negatively with C:N and lignin:N ratios. The effects of microclimate resulting from canopy damage on litter decomposition was also examined at the same time using standard litter with the same litter quality parameters, but they showed no significant differences among the insect damage levels indicating that alteration of the litter quality parameters produced by I. typographus damage played a more important role than altered microclimate in controlling needle litter decomposition rates. However, changes in microclimate factors due to topography influenced decomposition rates.     

Biomass expansion factors for Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis> (Bong.) Carr.) in Ireland

The assessment of a forest resource in national inventories provides a firm basis for the calculation of biomass and carbon (C) stocks of forests. Biomass expansion factors (BEFs) and conversion factors provide a robust and simple method of converting from forest tree stem volume to total forest biomass. These factors should be constructed on the basis of nationally specific data in order to take account of regional differences in growth rates, management practices, etc. The objective of this study is to improve the accuracy of biomass estimation by calculating a range of age-dependant BEFs from representative data that more accurately describe the allometry of present forests. The results from this study show that the allocation of biomass to compartments in forest stands and throughout a rotation varies considerably, and that the use of BEFs for the calculation of C stocks in forests of sub-timber dimensions is highly impractical.

<Emphasis Type="Italic">Bursaphelenchus mucronatus</Emphasis> (Nematoda: Aphelenchoididae) vectored by <Emphasis Type="Italic">Monochamus </Emphasis><Emphasis Type="Italic">urussovi</Emphasis> (Coleoptera: Cerambycidae) in Hokkaido,Japan

Bursaphelenchus mucronatus Mamiya et Enda has been recovered for the first time from adults of the cerambycid beetle, Monochamus urussovi (Fischer), in Hokkaido, Japan. The nematode was also recovered from the inner bark of Picea jezoensis (Siebold et Zuccarini) Carrière and Abies sachalinensis (Fr Schmidt) Masters infested with M. urussovi larvae. PCR–RFLP analysis indicated that B. mucronatus in Hokkaido is the European type.     

Spatial pattern of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and oak (<Emphasis Type="Italic">Quercus pubescens</Emphasis> Mill.) seedlings in natural pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) woodlands

Spatial pattern of recruitment is an important factor influencing population dynamics of plant communities. The spatial pattern is determined by seed dispersal and by the spatial variability of germination and initial survival. In the process of forest expansion following farmland abandonment, mid- and late-successional species are often dispersed in pioneer forests by birds. This could result in an aggregated spatial pattern of seeds that could influence the dynamics of these species in mixed pioneer forests. In the sub-Mediterranean area, mid- and late-successional species such as Quercus pubescens (downy oak) and Fagus sylvatica (European beech) are expected to replace pioneer Pinus species. Using a point sampling method we demonstrated that beech and oak seedlings (height <2 m) have a clumped distribution in the understorey of pine. This could result from an aggregated dispersal by jays (dispersal effect) or from preferential recruitment in particular habitats (habitat effect). To test these hypotheses we proposed a statistical analysis of spatial patterns of regeneration of beech and oak. Ground cover variables (i.e. cover by rock outcrops, herbs, box shrubs, mosses or pine) did not differ significantly around seedlings as compared with random sample plots. Likewise, clumped seedlings had growth similar to isolated seedlings, thus refuting the hypothesis of preferential location in the most favourable microsites. Aggregated dispersal seems to be involved in the process of regeneration. Since beech and oak seedlings have contrasting ecological demands, we discuss the implication of this pattern for the replacement dynamics of pine by these species.     

Microfibril angle of Norway spruce [Picea abies (L.) Karst.] compression wood: comparison of measuring techniques

The structure of cellulose, especially the microfibril angles (MFAs), in compression wood of Norway spruce [Picea abies (L.) Karst.] was studied by wide- and small-angle X-ray scattering and polarizing microscopy. On the basis of the X-ray scattering experiments the average MF As of the cell wall layers S₂ and S₁ of the studied sample are 39 and 89, respectively; and the average diameter and length of the cellulose crystallites are 2.9 and 20.0nm, respectively. The average of the whole MFA distribution is shown to agree with the one obtained by polarizing microscopy of macerated fibers.     

Stand density and growth of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> (L.) Karst.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.): evidence from long-term experimental plots

On the basis of nine Norway spruce (Picea abies (L.) Karst.) and ten European beech (Fagus sylvatica L.) thinning experiments in Germany, for which both residual and removed stock had been registered first during 1870, I scrutinize how moderate and heavy thinning from below (B-, C-grade) affects the production of merchantable volume compared with light thinning (A-grade). In relation to A-grade, cumulative merchantable volume (CV) of B- and C-grade amounts in average to 103–107% in juvenile and to 97–102% in mature Norway spruce stands. The corresponding findings for European beech are 101–106% and 94–102%. CV of individual stands varies between 89% and 130% for Norway spruce and 73% and 155% for European beech (CV of A-grade = 100%). These findings are substantiated by the relation between stand density (SDI) and periodic annual increment (PAI). On the B- and C-grade plots of spruce and beech, respectively, SDI was reduced down to 41–91% and 31–83% of the A-grade. When SDI is reduced in young stands, PAI follows a unimodal curve. Norway spruce’s PAI culminates in 109% if SDI is reduced to 59%; European beech’s PAI culminates in 123% when density is reduced to 50%. Whereas Norway spruce’s growth reacts most positively on thinning under poor site conditions and with increment reduction on favourable sites, European beech behaves oppositely. With stand development the culmination point of the unimodal relation moves towards maximum density, so that in older stands PAI follows the increasing pattern, which is the left portion of a unimodal curve. A model is presented which apparently unifies contradictory patterns of stand density–growth reactions by integrating relative stand density, average tree size and site fertility effects, and makes the findings operable for forest management.     

Effect of seed source elevation on bud opening of Yezo spruce (<Emphasis Type="Italic">Picea jezoensis</Emphasis>)

Spring frost damage is one of the obstacles to be overcome in establishing a Yezo spruce (Picea jezoensis Carr.) plantation. The timing of bud opening of Yezo spruce seedlings cultured from seeds collected at three different elevations (420, 700, and 1200m) in the Tokyo University Forest in Hokkaido was observed at two planting site elevations (610 and 750m) to investigate whether seed source elevation or planting site elevation affects the bud opening date. At both planting sites the bud opening dates were not significantly different among the seedling groups from different seed source elevations. On the other hand, bud opening dates of seedlings planted at the lower elevation site were significantly earlier than those planted in the higher elevation site. It could be deduced from this study that the environment of the planting site rather than seed source elevation affects the bud opening date of Yezo spruce.     

Frost hardiness of nutrient-loaded two-year-old <Emphasis Type="Italic">Picea abies</Emphasis> seedlings in autumn and at the end of freezer storage

The effects of nutrient loading (NLOAD) on the frost hardening and dehardening of Picea abies (L.) Karst. seedlings were investigated under nursery conditions. Before NLOAD, second-year container seedlings were either short-day (SD) treated for 3 weeks in July or left for the natural photoperiod (CO). By mid-September, after 5 weeks of NLOAD, the fertilization of three foliar nutrient concentration levels (low = L-SD, medium = M-SD, and high = H-SD) for the SD-treated seedlings and one (medium = M-CO) for the CO-seedlings was completed. The NLOAD resulted in foliar nitrogen concentration 10.6, 16.1, 22.3, and 17.5 g kg⁻¹ for L-SD, M-SD, H-SD and M-CO seedlings, respectively. The NLOAD had no effects on the morphology or dry mass variables of the seedlings, while SD-treatment reduced the dry mass of shoots, but not that of roots. The frost hardiness (FH) of different batches of the seedlings was assessed by the visual scoring of damage in their needles, stems and buds after their controlled exposure to freezing during frost hardening and dehardening. The low nutrient concentration in the SD-treated seedlings (L-SD seedlings) resulted in poor FH, to an even lower extent than that of the M-CO seedlings. The NLOAD did not affect the dehardening of the seedlings at the end of the freezer storage in the following spring.     

Characterization of lignin-carbohydrate linkages in the residual lignins isolated from chemical pulps of spruce (<Emphasis Type="Italic">Picea abies</Emphasis>) and beech wood (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>)

To investigate the linkage types between carbohydrates and lignin, residual lignins were isolated from three different unbleached pulps [kraft, alkaline sulfite anthraquinone methanol (ASAM), and soda with anthraquinone (AQ) and methanol] of spruce and beech wood and then characterized by oxidation with 2,3-Dichloro-5,6-dicyanobenzoquinone and followed by Prehm’s methylation. In residual lignins, sugar moieties were bound to lignins via benzyl ether bonds. In particular, galactose and mannose are predominantly linked to lignin fragments in residual lignins of spruce wood, while xylose and galactose are favored in the formation of LC bonds in the residual lignins of beech wood. In the case of hexoses, primary hydroxyl groups (C6 position) preferentially take part in benzyl ether linkages. Hydroxyl groups in the C2 and C3 positions of xylose participate in LC bonds and a small portion of arabinose was notably connected to lignin via the C5 position. Approximately seven or eight sugars were connected in soda/AQ/methanol residual lignin per 100 C9 lignin units, while the frequencies of LC bonds in kraft and ASAM residual lignins were distinctively less at one to three sugars per 100 C9 lignin units. The publication of this article was made possible by an Emachu Research Fund. The authors are grateful for the fund.     

Successful <Emphasis Type="Italic">Agrobacterium</Emphasis>-mediated transformation of <Emphasis Type="Italic">Populus tomentosa</Emphasis> with apple <Emphasis Type="Italic">SPDS</Emphasis> gene

The problem of salinized soils has become one of the most serious constraints to agricultural and forest productivity. With the purpose of enhancing salt stress tolerance of Populus tomentosa, we transformed this tree species with spermidine synthase （SPDS） genes derived from an apple by an Agrobacterium-mediated method. Four transgenic clones were confu＇med by PCR and Southern blot analysis. As well, the expression of introduced SPDS genes was analyzed by real-time quantitative PCR.     

Bioremediation of CCA-treated wood by brown-rot fungi <Emphasis Type="Italic">Fomitopsis palustris</Emphasis>, <Emphasis Type="Italic">Coniophora puteana</Emphasis>, and <Emphasis Type="Italic">Laetiporus sulphureus</Emphasis>

This study evaluated oxalic acid accumulation and bioremediation of chromated copper arsenate (CCA)-treated wood by three brown-rot fungi Fomitopsis palustris, Coniophora puteana, and Laetiporus sulphureus. The fungi were first cultivated in a fermentation broth to accumulate oxalic acid. Bioremediation of CCA-treated wood was then carried out by leaching of heavy metals with oxalic acid over a 10-day fermentation period. Higher amounts of oxalic acid were produced by F. palustris and L. sulphureus compared with C. puteana. After 10-day fermentation, oxalic acid accumulation reached 4.2 g/l and 3.2 g/l for these fungi, respectively. Fomitopsis palustris and L. sulphureus exposed to CCA-treated sawdust for 10 days showed a decrease in arsenic of 100% and 85%, respectively; however, C. puteana remediation removed only 18% arsenic from CCA-treated sawdust. Likewise, chromium removal in F. palustris and L. sulphureus remediation processes was higher than those for C. puteana. This was attributed to low oxalic acid accumulation. These results suggest that F. palustris and L. sulphureus remediation processes can remove inorganic metal compounds via oxalic acid production by increasing the acidity of the substrate and increasing the solubility of the metals.An erratum to this article can be found at     

Impacts of soil conditions and light availability on natural regeneration of Norway spruce <Emphasis Type="Italic">Picea abies</Emphasis> (L.) H. Karst. in low-elevation mountain forests

? Key message

Natural regeneration of P. abies (L.) H. Karst. may reach high densities in lower mountain elevations. The highest densities were found in sites with moderate light availability, with low pH, and not near the riverbank. However, age-height classes differed in the predicted magnitude of response, but were consistent in response directions. Mosses and understory species typical of coniferous forests were positively correlated with regeneration density.

? Context

Norway spruce Picea abies (L.) H. Karst. in Central Europe is at risk under climate change scenarios, particularly in mountain regions. Little is known about the impact of environmental factors on the natural regeneration of P. abies in low-elevation mountain forests.

? Aims

We aimed to assess impacts of distance from the riverbank, soil pH, and light availability on natural P. abies regeneration. We hypothesized that (1) natural P. abies regeneration would depend on light availability and soil pH and (2) there are understory plant species which may indicate the microsites suitable for natural regeneration of P. abies.

? Methods

The study was conducted in the Sto?owe Mountains National Park (SW Poland, 600–800 m a.s.l.). We established 160 study plots (25 m²) for natural regeneration, light availability, soil pH, and understory vegetation assessment.

? Results

The highest densities of natural regeneration of P. abies were observed in sites with moderate light availability (0.1–0.2 of open sky) and low pH (3.5–4.5), and located relatively far from the riverbank. Cover of 22 understory plant species were correlated with natural P. abies regeneration densities, mostly positively.

? Conclusion

Different stages of natural regeneration of P. abies revealed different regeneration niches. Most understory plant species (bryophytes and herbs typical of coniferous forests) do not compete with natural regeneration of P. abies.