Impact of soil pressure and compaction on tracheids in Norway spruce seedlings

We tested the effect of soil compaction on Norway spruce seedlings in terms of the size and theoretical volume flow rate of the tracheids. The results show that soil pressure limits growth in the diameter of the lumens of tracheids in all parts of seedlings studied. The tracheids of the roots with primary xylem had larger lumens than those of the roots and shoots with secondary xylem in both unloaded and loaded seedlings. This corresponds to the higher cumulative theoretical volume flow rate of the tracheids from roots with primary xylem than those from roots and shoots with secondary xylem. Although the volume flow rate of tracheids, according to the Hagen-Poiseuille law, was directly proportional to the quadratic power of the capillary diameter (tracheid lumen), the cumulative curve of the theoretical hydraulic volume flow rate was higher or relatively comparable in loaded seedlings. An explanation for these findings is that there were higher gradients of water potential values in roots and leaves in loaded seedlings because the lengths of the conductive pathways were 27% shorter than in unloaded seedlings. We hypothesise that trees have adapted to different stresses by shortening their conductive pathways to maintain a transpiration rate similar to that of non-stressed trees. These results concerning the impact of soil compaction on tracheid diameter and volume flow rate improve our understanding of the growth and functioning of different conifer organs and the mechanisms underlying the efficiency of water transport through the root xylem to the shoot.     

Seasonal changes of soluble carbohydrates in mycorrhizas of Norway spruce and changes induced by exposure to frost and desiccation

Mycorrhizas of six spruces of a stand were collected over two years and were analysed for soluble carbohydrates (sucrose, raffinose, glucose, trehalose and mannitol). The level of these carbohydrates was high during winter and low during summer. Upon exposure of excised mycorrhizas to frost or desiccation, the content of trehalose roughly doubled. A positive correlation was found between the trehalose content of mycorrhizas and their ability to survive frost or desiccation as determined by the electrolyte leaching method.     

Seasonal patterns of colonization of Norway spruce needles by Lophodermium piceae

Summary Colonization of living Norway spruce needles by Lophodermium piceae was studied in southern Sweden. One‐month to 3‐year‐old needles were assessed for frequency of infection. Needles appearing in the growing season of 1996 were monitored and sampled at 1 to 4 week intervals over a period of 14 months. Infection frequency of the current‐year needles increased from around 20% in July to approximately 40% in August 1996. This level had then doubled to approximately 80% after another growing season in August 1997. Multiple infections were rare in the needles during the first growing season, but after the second growing season at least 70% of the needles had multiple infections. The average number of infections in 9‐, 21‐ and 33‐month‐old needles were 1.3, 6.3 and 8.4, respectively. The size of the individual mycelia was estimated to be small in all needles, none of the individuals could be isolated from more than two adjacent needle sections, about 1.5 mm long. In conclusion, the number of L. piceae infections per needle increased during ageing of needles but mycelial growth is probably postponed until needle senescence and death.     

Inherited narrow crown form, harvest index and stem biomass production in Norway spruce, Picea abies

The effect of crown form on stem biomass production was investigated in an 18-19-year-old Norway spruce stand (Picea abies (L.) Karst.). The harvest index was 0.271 in pendula trees, which have a heritable narrow crown form, and 0.235 in normal-crowned trees and the dry weights of stem biomass were 2.57 kg and 3.37 kg, respectively. However, the production of stem biomass per crown projected area was more than twice as much in pendula trees as in normal-crowned trees. Results indicate that the crown form of pendula spruce is less plastic than that of normal-crowned spruce suggesting that such trees might maintain a high stemwood yield per unit ground area with increasing stand density. Because pendula spruce has a higher harvest index and a less flexible crown than normal-crowned spruce, it might be useful for crop tree ideotype breeding.     

Fresh-wood bending: linking the mechanical and growth properties of a Norway spruce stem

To provide data and methods for analyzing stem mechanics, we investigated bending, density and growth characteristics of 207 specimens of fresh wood from different heights and radial positions of the stem of one mature Norway spruce (Picea abies L. Karst.) tree. From the shape of each stress-strain curve, which was calculated from bending tests that accounted for shear deformation, we determined the modulus of elasticity (MOE), the modulus of rupture (MOR), the completeness of the material, an idealized stress-strain curve and the work involved in bending. In general, all mechanical properties increased with distance from the pith, with values in the ranges of 5.7-18 GPa for MOE, 23-90 MPa for MOR and 370-630 and 430-1100 kg m(-3) for dry and fresh wood densities, respectively. The first three properties generally decreased with stem height, whereas fresh wood density increased. Multiple regression equations were calculated, relating MOR, MOE and dry wood density to growth properties. We applied these equations to the growth of the entire stem and considered the annual rings as superimposed cylindrical shells, resulting in stem-section values of MOE, MOR and dry and fresh densities as a function of stem height and cambial age. The standing tree exhibits an inner stem structure that is well designed for bending, especially at a mature stage.     

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.     

Proteins in Norway spruce thermomechanical pulp

Two methods of cytochemical staining using Coomassie dye and Cu⁺-bicinchoninic acid, respectively, showed that there are proteins in thermomechanical pulp (TMP) of Norway spruce. Protein isolated from TMP was analyzed for amino acid composition. There was about twice the amount of acidic amino acid material compared with basic amino acids, and the presence of glucosamine indicated that the isolated polypeptides also contained glycoproteins. The presence of proteins in ray cells and fiber tracheids in TMP adds to the chemical heterogeneity of the structurally complex high-yield pulp.     

A method for quantifying changes in the epicuticular wax structure of Norway spruce needles

A method for quantifying changes in the epicuticular wax structure of spruce needles (Pieea abies) has been developed for the scanning electron microscope. This evaluation system, based on a five-point scale, classifies only the epistomatal waxes of current-year needles. Two criteria, the different crystal wax forms and the amount of changed wax as a percentage of the total stomatal area have been used for classification. Statistically significant results (accuracy below ± 10% at 5% error level) were obtained under the following conditions. Mixed needle samples were taken from two adjacent trees per site. These were in a dominant and predominant social position in the stand. The needles were collected from the 7^th whorl of sun crowns exposed to all wind directions. The wax quality was determined by evaluating three stomata per needle. One stoma was investigated at the needle base, one m the middle and the third at the apex of the needle. 24 needles were evaluated per site. SEM-Investigations of needle waxes of 25 Austrian forest sites with no local emission source (background sites) showed, that the mean wax quality ranged from 1.6 to 4.1, according to this classification method. Stepwise multiple regression analyses with site characteristics and element contents of the needles (nutrient elements, heavy metals) confirmed correlations between the wax quality and the manganese- and sulphur-content of the needles and also the average degree of crown defoliation.     

Boron retranslocation in Scots pine and Norway spruce

We previously traced 10B-enriched boric acid from shoots to roots to demonstrate the translocation of boron (B) in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. To gain a more detailed understanding of B translocation, we sought: (1) to demonstrate B retranslocation directly, by showing that foliar-applied 10B is located in the new growth after dormancy; and (2) to assess whether shoot-applied B affects growth in the long term. We applied 10B-enriched boric acid to needles of Scots pine and Norway spruce seedlings. After a dormancy period and 9 weeks of growth, small but significant increases in the 10B isotope were found in the new stem and needles of both species. In Scots pine, the total B concentration of the new stem was also increased. Both species contained polyols, particularly pinitol and inositol. Boron-polyol complexes may provide a mechanism for mobilizing B in these species. To determine the long-term effects of applied B, seedlings were grown for two growing seasons after the application of 10B to shoots. In Norway spruce, the proportion of 10B in the root systems and current needles of the harvest year was slightly higher than in the controls, and in Scots pine root systems, marginally so. The B treatment had no effect on growth of Norway spruce seedlings. In Scots pine seedlings, the B treatment caused a 33% increase in total dry mass and significantly increased the number of side branches.     

Fresh-stem bending of silver fir and Norway spruce

The bending and growth characteristics of large fresh stems from four silver fir (Abies alba Mill.) and three Norway spruce (Picea abies (L.) Karst.) trees were studied. Twenty logs taken from different stem heights were subjected to four-point bending tests. From the bending test records, we calculated stress-strain curves, which accounted for detailed log taper, shear deformation and self weight. From these curves we determined, among other parameters, the modulus of elasticity (MOE), the modulus of rupture (MOR) and the work absorbed in bending (W). No significant differences were found between species for the wood properties examined. Values of MOE, MOR and W generally decreased with stem height, with MOR in the range of 43 to 59 MPa and MOE ranging from 10.6 to 15.6 GPa. These MOE values are twice or more those reported for stems of young Sitka spruce (Picea sitchensis (Bong.) Carr.) trees. Based on the radial growth properties measured in discs from the logs, we calculated predicted values of MOE and MOR for the stem cross section. The predictions of MOE were precise, whereas those of MOR were approximate because of a complex combination of different failure mechanisms. Methods to test and calculate MOE, MOR and W for the stems of living trees are discussed with the aim of improving analyses of tree biomechanics and assessments of forest stability protection.     

Quality of timber products from Norway spruce

Summary The overall aim of this study and series of papers is to address the key variables for timber quality and to optimize the utilization of Norway spruce timber with respect to construction purposes. It is the end-user's degree of overall satisfaction that determines the quality of a product. Therefore, the performance of structural timber cannot solely be defined by mechanical properties. Geometric performance (warp) must be improved if timber is to continue as an important building material.An experimental study of the spatial variation in warp and bending properties of fast-grown Norway spruce is introduced. In this paper, the growth characteristics are presented as a function of stand and location in the tree. The knot area ratio (KAR) was considerably higher in the core (0.31) compared with timber closer to bark (0.21). The top log studs had higher KAR (0.38) than the corresponding butt log studs (0.31). The average grain angle was 3.5% ( 2°) and appeared not to vary radially. The presence of compression wood was much more common in the top log timber (75%) than in the butt log (44%). However, no consistent radial variation in compression wood was found.The authors gratefully acknowledge the support from EEC Forest programme, Contract No. MA2B-0024 and from Södra Timber AG from NUTEK project No. 9100554     

Quality of timber products from Norway spruce

Summary Strength and stiffness together with some properties characterizing the stand and the growth of trees were studied. Specimens (45 × 70 × 2900 mm³) were cut from different radial and longitudinal positions, from fast-grown trees from two stands in southern Sweden. These trees had relatively large annual rings (4–6 mm) and were not representative of Norway spruce in Sweden but are an example of the intensivelymanaged stands which will probably constitute a substantial part of the raw material supply in the future.The results indicate that the mean values for strength and stiffness were lowest for the core studs and increased further away from the pith. This radial variation in strength and stiffness appears to be associated with the variation in ring width. Density alone, on the other hand, does not explain the radial variation but should be used together with either ring width or knot area ratio to explain the stiffness and strength respectively. The increase in the strength and stiffness of the core studs from the butt logs to the top logs was significant. Density alone was found to be the best variable to explain the longitudinal variation between the butt logs and the top logs. The heartwood formation in the butt log juvenile core appeared not to have a positive effect on strength and stiffness. The occurrence of compression wood, the magnitude of grain angle and the margin knot area ratio had only a minor effect on strength and stiffness.The authors gratefully acknowledge the support received from the EEC forest research programme, Contract no MA2B-0024, from NUTEK, project no 9100554 and from Södra Timber AB     

Quality of timber products from Norway spruce

Summary In a previous paper in this series it was concluded that the warp (twist, crook and bow) of structural timber products is of utmost importance for end-user satisfaction. In this paper, a study of the spatial variation of warp of fast-grown Norway spruce is presented. The material, 372 studs (45 × 70 × 2900 mm²), was cut from different radial and longitudinal positions in trees from two stands.The results indicated that twist and crook were at maximum near the pith. However, twist decreased much more rapidly with the distance from pith compared with crook. Crook associated with core studs was larger in the butt log than in the top log, while twist and bow in core studs appeared not to vary longitudinally. Heartwood formation in the butt log juvenile core did not diminish warp propensity. Twist was rather well correlated to the distance between pith and centroid of cross section. The ratio of grain angle to pith distance was the best parameter to predict twist. Presence of compression wood increased bow and crook significantly, while ring width, density and knot area ratio did not substantially contribute to explain warp variation.A moisture content change from 18% to 12% decreased the overall acceptability, expressed by stud grade yield, from 76% to 43%. Thus, timber products should be delivered at a moisture level close to the one expected in the finalized building in order to avoid excessive warp after delivery.The authors gratefully acknowledge the support from EEC Forest programme, Contract No MA2B-0024, from NUTEK project No 9100554 and from Södra Timber AB     

Effects of genetic entry and competition on above ground biomass production of Norway spruce grown in southern Finland

The effects of genetic entry and competition on the above ground dry biomass production (i.e. stem wood, needles, branches and harvest index) was studied in 20 Norway spruce (Picea abies (L.) Karst.) clones grown in southern Finland. Furthermore, the measured above ground biomass components were compared against the corresponding estimations based on biomass models developed previously. The clones included both Finnish and Russian clones, as well as provenance-hybrids clones. Differences existed between clones in stem dry mass production, but not in harvest index or in crown dry mass. However, the competition caused by neighboring trees also significantly affected above ground dry biomass, as well as the biomass of crown and stem separately. Differences in competition between the clones could not be found. Unlike the dry mass of branches, the dry mass of needles and stem could be estimated well for individual sample trees with the available biomass models. Moreover, the clone with the largest above ground dry biomass production had nearly doubled production, on average, than the average over all clones. Thus, some of the clones showed especially high potential for biomass recovery in energy wood thinning.     

Three-dimensional elastic behaviour of common yew and Norway spruce

In view of its high density, yew wood has a remarkably low longitudinal Young’s modulus, which makes it unique among coniferous woods. However, the elastic response of yew related to other load directions is largely unknown. Therefore, our goal was to comprehensively characterise the three-dimensional elastic behaviour of yew wood. To achieve this, we performed tensile tests on dog-bone-shaped yew specimens and determined the three Young’s moduli and six Poisson’s ratios using a universal testing machine and a digital image correlation technique. All tests were also applied to spruce as reference species. After including the shear moduli determined in a prior study by our group, all elastic engineering parameters of yew and spruce were ascertained. Based on these values, the three-dimensional elastic behaviour was describable with deformation bodies and polar diagrams. Evaluating these illustrations revealed that yew had a lower stiffness only in the longitudinal direction. In all other three-dimensional directions, spruce was clearly more compliant than yew. Particularly, in the radial–tangential plane, both species varied largely in their degree of anisotropic elasticity. All mentioned differences between yew and spruce originate at the microstructural level.     

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.     

Allometry varies among related families of Norway spruce

Key message

Slopes and intercepts of allometric equations for organs’ biomass varied among half-sib families of Norway spruce and between age categories in a family-dependent manner. Genetic variation should be accounted for when applying allometric analysis to mixtures of genetic groups.

Context

Genetic variation in relationships among plant biomass components was rarely addressed in trees, though depending on deployment strategies in tree improvement programs, variation among genetic groups in plant organs’ growth rates, and thus biomass allocation, would affect forest growth and carbon balance.

Aims

We investigated growth and biomass distribution in Norway spruce (Picea abies [L.] H. Karst) half-sib families. We assumed invariance among families in allometric relationships, and stability in growth rates of different organs between ages 2 and 8 years in the subset of those families. We also tested for ontogenetic trend in allometry using the independent dataset of biomass at age 21 years.

Methods

We analyzed allometric relationships among plant components using standardized major axis regression.

Results

Slopes and intercepts of allometric relationships varied among families, indicating variation in both organs’ growth rates and biomass partitioning at a given plant size. Variation in scaling exponents between age categories was also dependent on the family and plant organ considered.

Conclusion

Variation in slopes of allometric relationships indicates that a single scaling coefficient should not be applied when different genetic groups are compared. For the interpretation of age effect on biomass partitioning, both slopes and intercepts of allometric relationships should be examined.

     

Influence of planting density and rotation age on the profitability of timber production for Norway spruce in Central Europe

When forest stands are established by planting, the initial density is often chosen based on recommendations of specific plant numbers. Since forest owners typically face different alternatives for the employment of their opportunities and since market situations often change more rapidly and on a smaller scale than the characteristics of forest sites, the question arises whether these factors are reflected in one or within a small range of planting densities. Even though some advised densities might be profitable for some forest owners and market situations, the shift of profitable timber production as a result of changes in the circumstances surrounding the investment is unclear. In this paper, the influence of the planting density on the profitable production of timber is analysed with the explicit inclusion of price differentials as these offer incentives for the choice of the planting density in Central Europe. Since numerical calculations based on empirical data are required for an analysis of the complex facets of price differentials, the specific statements are restricted to the tree species Norway spruce (Picea abies Karst.). By means of typical calculation examples, the effects of changes in investment parameters are exemplified and discussed against the background of a sensitivity analysis. The results reveal the boundaries for the profitable production of high-quality timber and show that forest owners might compensate changes in the investment situation by adjustments of either or both the rotation age and the planting density.