Hydraulic and mechanical properties of young Norway spruce clones related to growth and wood structure

Stem segments of eight five-year-old Norway spruce (Picea abies (L.) Karst.) clones differing in growth characteristics were tested for maximum specific hydraulic conductivity (k(s100)), vulnerability to cavitation and behavior under mechanical stress. The vulnerability of the clones to cavitation was assessed by measuring the applied air pressure required to cause 12 and 50% loss of conductivity (Psi(12), Psi(50)) and the percent loss of conductivity at 4 MPa applied air pressure (PLC(4MPa)). The bending strength and stiffness and the axial compression strength and stiffness of the same stem segments were measured to characterize wood mechanical properties. Growth ring width, wood density, latewood percentage, lumen diameter, cell wall thickness, tracheid length and pit dimensions of earlywood cells, spiral grain and microfibril angles were examined to identify structure-function relationships. High k(s100) was strongly and positively related to spiral grain angle, which corresponded positively to tracheid length and pit dimensions. Spiral grain may reduce flow resistance of the bordered pits of the first earlywood tracheids, which are characterized by rounded tips and an equal distribution of pits along the entire length. Wood density was unrelated to hydraulic vulnerability parameters. Traits associated with higher hydraulic vulnerability were long tracheids, high latewood percentage and thick earlywood cell walls. The positive relationship between earlywood cell wall thickness and vulnerability to cavitation suggest that air seeding through the margo of bordered pits may occur in earlywood. There was a positive phenotypic and genotypic relationship between k(s100) and PLC(4MPa), and both parameters were positively related to tree growth rate. Variability in mechanical properties depended mostly on wood density, but also on the amount of compression wood. Accordingly, hydraulic conductivity and mechanical strength or stiffness showed no tradeoff.     

A model for spatial wood density gradients in Norway spruce stems and stochastic between-stem dissimilarities for basic and dry density

Density is a fundamental softwood quality trait. The aim of this paper is to identify a spatial model for wood density variation in spruce stems, with the main focus on basic density. Six thousand 20-mm-wide cubes systematically sampled from 85 trees in western Norway were analysed. The overall radial density pattern was the J shape with local maximum in the pith and increasing density outwards from a minimum at some distance from the pith. The minimum appeared closer to the pith further up in the tree. The stochastic nature of the six tree-wise parameters defining density gradients from pith to surface and from base to top was analysed and described. The results provide information needed to simulate density variation inside stems and between stems in a population. Considering the fundamental influence of density on a range of wood traits, such simulation should be of great significance for scientific and industrial analyses.     

Hydraulic and anatomical properties of light bands in Norway spruce compression wood

Compression wood (CW), which is formed on the underside of conifer branches, exhibits a lower specific hydraulic conductivity (k(s)) compared with normal wood. However, the first-formed tracheids of an annual ring on the underside of a conifer branch often share several properties with normal tracheids, e.g., thin cell walls and angular cross sections. These first-formed tracheids appear bright when observed by the naked eye and are therefore called light bands (LB). In this study, hydraulic and related anatomical properties of LBs were characterized and compared with typical CW and opposite wood (OW). Measurements were made on branches of Norway spruce (Picea abies (L.) Karst.). Specific hydraulic conductivity was measured with fine cannulas connected to microlitre syringes. Micro- and ultrastructural analysis were performed on transverse and radial longitudinal sections by light and scanning electron microscopy. Xylem areas containing both typical CW and LBs had a k(s) 51.5% that of OW (7.95 +/- 0.97 m(2) s(-1) MPa(-1) x 10(-4)), whereas k(s) of pure CW was only 26.7% that of OW. The k(s) of LBs (6.38 +/- 0.97 m(2) s(-1) MPa(-1) x 10(-4); 80.3% of OW) was estimated from these k(s) values because the cannulas were too wide to measure the k(s) of LBs directly. Mean lumen area of first-formed tracheids on the underside of branches was 65.7% that of first-formed tracheids in OW and about three times that of CW. Light-band tracheids exhibited a bordered pit frequency of 42.7 +/- 1.3 pits mm(-1), which was three times that in CW and 1.6 times that in OW. Bordered pit apertures in LB tracheids (9.15 +/- 0.60 microm(2)) were 1.7 times wider than those in CW and similar in aperture to those in OW. The high k(s) of LBs was correlated with their wide tracheid lumina, high pit frequency and wide pit apertures. We therefore suggest that LBs have a primarily hydraulic function within the mechanically optimized CW region. This might be important for supplying water to living tissues on the underside of branches, as well as to other distal areas along water transport pathways following the spiral grain of wood.     

Development and variation of wood density, kraft pulp yield and fibre dimensions in young Norway spruce (Picea abies)

Important wood, pulp and fibre properties were investigated on small wood samples from two Rumanian, one German and one Norwegian provenance of Norway spruce (Picea abies) grown in Sørkedalen, Norway. Several samples were collected from inside each single tree, both in radial and transversal direction in the stem. Data were collected from a total of 59 trees, each 28 years of age. All investigated properties showed close relationship to ring number (RN) (cambium age). For basic density and fibre wall thickness (FWT), a fast decline was first observed when moving from pith to bark and a minimum value was found around RN 5–8. The declining trend was then followed by an increase. Kraft pulp yield, fibre length and fibre width (FW) also increased with RN, but the increase was most pronounced close to the pith. The pulp yield (PY) more or less stabilised outside of RN 5–6. FW showed a decrease outside of RN 10–12, but this was probably due to the remarkable simultaneous drop in ring width for the investigated trees. FW decreased, while basic density and fibre length increased with increasing height in the tree. PY and FWT were not affected by height in tree. Diameter at breast height (DBH) was the most important variable indicating differences between trees in the investigated material. Basic density, fibre length and FWT decreased, while FW increased with increasing DBH. Height to crown had a positive effect on basic density, but had no influence on any of the remaining properties. Differences between provenances were found for basic density, fibre length and FW. The analyses showed that it is possible to describe the variation inside and between trees satisfactorily for a range of important wood and fibre properties.     

Resin pocket occurrence in Norway spruce depending on tree and climate variables

Resin or pitch pockets cause a significant reduction in timber quality of Norway spruce (Picea abies (L.) Karst). Their occurrence is very variable within the stem and between trees and stands. Researchers have suggested several causes of resin pocket formation with a majority arguing in favour of storms as the main causing factor.     

Occurrence of microorganisms in the wood of Norway spruce trees from polluted sites

Bacteria, yeasts and other fungi were isolated from stem and root wood of Norway spruee trees with disease symptoms and were identified in order to evaluate a possible mierobial involvement in the widespread tree decline in Central Europe.     

Control of tracheid cross-sectional dimensions in Norway spruce and Scots pine wood raw material

Abstract

Wood properties, including tracheid cross-sectional dimensions, show a large degree of variation. To improve the properties of products made from wood, different methods to control variation have been developed. This study aims to determine the theoretical efficiency of three control strategies: the fractionation of pulped tracheids into earlywood and latewood, the separation of juvenile and mature wood, and sorting of logs according to tree size. The efficiency of each method was studied by first constructing virtual trees from measured tracheid cross-sectional dimensions, then simulating the efficiency of above-mentioned methods. The tracheid dimension data include Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.). The simulations show that separation into earlywood and latewood classes has the highest theoretical efficiency and yields the lowest variances in raw material. Classification into juvenile and mature wood groups is the second most efficient method, and the sorting of logs according to the size class of the tree is the least efficient method. It was also concluded that the variation in cell-wall thickness and radial diameter mainly originates from differences between earlywood and latewood, whereas the variation in tangential diameter mainly originates from differences between mature and juvenile wood.     

Tradeoffs between hydraulic and mechanical stress responses of mature Norway spruce trunk wood

We tested the effects of growth characteristics and basic density on hydraulic and mechanical properties of mature Norway spruce (Picea abies (L.) Karst.) wood from six 24-year-old clones, grown on two sites in southern Sweden differing in water availability. Hydraulic parameters assessed were specific hydraulic conductivity at full saturation (ks100) and vulnerability to cavitation (Psi50), mechanical parameters included bending strength (sigma b), modulus of elasticity (MOE), compression strength (sigma a) and Young's modulus (E). Basic density, diameter at breast height, tree height, and hydraulic and mechanical parameters varied considerably among clones. Clonal means of hydraulic and mechanical properties were strongly related to basic density and to growth parameters across sites, especially to diameter at breast height. Compared with stem wood of slower growing clones, stem wood of rapidly growing clones had significantly lower basic density, lower sigma b, MOE, sigma a and E, was more vulnerable to cavitation, but had higher ks100. Basic density was negatively correlated to Psi50 and ks100. We therefore found a tradeoff between Psi50 and ks100. Clones with high basic density had significantly lower hydraulic vulnerability, but also lower hydraulic conductivity at full saturation and thus less rapid growth than clones with low basic density. This tradeoff involved a negative relationship between Psi50 and sigma b as well as MOE, and between ks100 and sigma b, MOE and sigma a. Basic density and Psi50 showed no site-specific differences, but tree height, diameter at breast height, ks100 and mechanical strength and stiffness were significantly lower at the drier site. Basic density had no influence on the site-dependent differences in hydraulic and mechanical properties, but was strongly negatively related to diameter at breast height. Selecting for growth may thus lead not only to a reduction in mechanical strength and stiffness but also to a reduction in hydraulic safety.     

Automated image processing for quantification of blue-stain discolouration of Norway spruce wood

Bioincising is a promising method for enhancing liquid uptake (e.g. preservatives or wood-modification agents) in refractory wood. Incubation with the white-rot fungus, Physisporinus vitreus, which selectively degrades pit membranes, results in deeper and more homogeneous penetration of liquids. Conventional methods of assessing the degree of fungal discolouration of wood after treatment with preservatives (e.g. European standard EN 152) are partly based on a subjective rating scale, which gives a rough value of the surface colonisation by blue-stain fungi. Hence, an automated image processing (AIP) procedure was developed for standardised quantification of the segmentation thresholds of discolouration and tested against manual segmentation analysis. Using the red filter in the AIP method revealed high correlation (R ² 0.95) and allowed for more user friendly and objective determination of blue staining of wood.     

Ceratocysts polonica inoculated in Norway spruce: Blue-staining in relation to inoculum density,resinosis and tree growth

When Norway spruce trees were inoculated with Ceratocystis polonica, a dose-dependent response was recorded. Local resinosis near the inoculation sites decreased with increasing inoculum density; parallelled by an increasing degree of sapwood staining, and tree mortality. Suppressed trees, and trees showing growth decline appeared more susceptible than trees suffering less competition.     

Crown density changes of Norway spruce and the influence from increased age on permanent monitoring plots in Norway during 1988–1997

On the ‘local county monitoring plots’ located throughout Norway a reduction of crown density has been noted during the period 1988–1997. The aim of this study was to determine whether this change could be attributed to normal effects due to the increased age of trees in the plots. The dataset comprised around 580 plots and 27 000 single trees of Norway spruce (Picea abies), with 10 years of crown density measurements available for each tree. A two-step approach was used, first to search for an expectancy for normal reduction of crown density by age derived from the dataset, and then to compare this with the actual reduction. The interpretation was somewhat complicated as the various results were dependent upon each other. Highly significant correlations were found between crown density and age. The relationship indicated an annual reduction of crown density of around 0.12%. However, the relationship varied both between years and between regions, and it was not possible to definitely determine whether the relationship was best described by linear or nonlinear models. Of major importance here is that the relationship appeared to be influenced by the presence of stresses, whose effects tended to be more severe in old stands. On this basis it seems questionable whether an expectancy for normal ageing can be properly defined. In the present study, however, it could still be definitely determined that the mean crown density change of—0.41% annually was too negative to be attributed to normal ageing, as it was clearly below all the suggested expectancies from the various models. This suggests that the amount of stress in the period under study has been higher than normal, and this encourages the search for causal agents in further studies. Changes in silviculture may have had some influence. The results were valid for most of Norway, with the exception of western and northern regions. Crown density assessments are subjective, which may possibly give erroneous time trends. However, it is argued that this is less likely to be of major importance in the present data.     

Effects of thinning and fertilization on wood properties and economic returns for Norway spruce

The effects of silviculture on wood and tracheid properties, and economic returns of Norway spruce (Picea abies (L.) Karst.) were investigated in two case studies, one comparing different thinning intensity in southeastern Finland and the other considering the effects of optimal nutrient addition in northern Sweden. Models for predicting the wood and tracheid properties of Norway spruce were integrated into a distance-independent process-based growth and yield model. Increasing the thinning intensity resulted in a lower mean wood density, tracheid length, and latewood proportion in harvested wood. Wood density and tracheid length of harvested pulpwood slightly decreased in later thinnings and final cuts. Thinning regimes with high early growing stock and decreasing later growing stock were most profitable. Nutrient addition accelerated volume growth and increased the value growth. The increase in volume growth due to nutrient addition more than offset the economic influence of the loss in wood density and tracheid length.     

Comparison of colonization capacity by asexual spores of Heterobasidion species in Norway spruce wood

Two species of the tree pathogenic fungus Heterobasidion spp. exist in Sweden, Heterobasidion annosum s.s. and Heterobasidion parviporum. Both species are known to infect Norway spruce (Picea abies). The aim of the study was to examine the interspecific competition between H. annosum s.s. and H. parviporum as well as their colonization rate in fresh Norway spruce wood. Equal amount of conidiospores from each species was sprayed together on 30 fresh, previously uninfected, Norway spruce billets. After incubation in a greenhouse, the proportion of Heterobasidion spp. colonies belonging to each species was recorded. Of the 196 colonies isolated from the upper part of billets, 195 were H. parviporum. All isolated colonies further down in the billets were H. parviporum. To study the colonization rate, H. annosum s.s. and H. parviporum were sprayed alone on 30 spruce billets each, incubated and growth recorded both vertically and horizontally. H. parviporum grew further down in the billets (p = 0.008) and covered a larger area (p < 0.001) than H. annosum s.s. While H. annosum s.s. and H. parviporum both infected fresh Norway spruce wood H. parviporum outgrew and outcompeted H. annosum s.s during the early colonization stage.     

Chemical composition of earlywood and latewood in Norway spruce heartwood,sapwood and transition zone wood

This study focused on the distribution of wood components along a cross section of a spruce stem. Thin samples of earlywood and latewood were analysed by special micro-scale analytical techniques. Heartwood contained significantly more lignin and less cellulose than sapwood. The total content of hemicelluloses was the same along the radial direction, but the distribution of sugar units differed. The amounts of arabinoglucuronoxylan and pectins were larger in the heartwood. The transition zone between heartwood and sapwood had a specific composition, with less lignin and lipophilic extractives than heartwood and sapwood. For earlywood and latewood, significant differences were found in the distribution of sugar units in hemicelluloses. Latewood contained clearly more galactoglucomannan than earlywood, and conversely less pectins. The lipophilic extractives were also less concentrated in the latewood.Abbreviations EW or E earlywood - LW or L latewood - HW heartwood - SW sapwood - TZ transition zone wood - A.R. annual ring - AcBr Acetyl bromide - Ara arabinose - Xyl xylose - Gal galactose - Glc glucose - Man mannose - Rha rhamnose - GlcA glucuronic acid - MGlcA 4-O-methyl-glucuronic acid - GalA galacturonic acid - o.d. oven dry     

Radial mechanical properties of high-temperature dried Norway spruce (Picea abies) wood

Abstract

Wood drying experiments were conducted in which the temperature and the drying rate were controlled independently. The mechanical properties of dried wood in radial loading were analysed in relation to drying parameters. Mass loss, due to thermal degradation of the wood structural components, occurred predominantly in slow high-temperature drying processes. Despite the higher mass loss, slowly dried wood showed similar radial strength and stiffness to rapidly dried wood. The formation of irreversible hydrogen bonds (hornification) within the wood structure may compete with the effects of mass loss on the radial mechanical behaviour of wood. However, both the mass loss and the hornification resulted in wood specimens with lower hygroscopicity. Application of slow high-temperature drying to reduce microscopic cell-wall damage, caused by anisotropic shrinkage of cell-wall layers, did not seem to affect the radial mechanical properties of wood. The effects of stress relaxation within the wood cell wall on the mechanical behaviour of wood may be offset by the degradation of structural components along with drying. Radial mechanical properties may be improved by rapid high-temperature drying up to high final dryness.     

Distribution of structure and lignin within growth rings of Norway spruce

A radial core from a Norway spruce (Picea abies (L.) Karst.) estimated to be about 107 years old was cut from a board and was analyzed for density and microfibril angle (MFA). Furthermore, cell geometry, wall thickness and lignin distribution were analyzed on three selected growth rings in detail. Intra-ring differences in the density profiles are also true for cell wall thicknesses as well as radial and tangential lumen diameters. A higher MFA was found for earlywood with a slow decrease toward the latewood region. The lignin was found to remain rather constant throughout the growth rings, which suggests a constant chemical composition of the cell wall material within the growth ring. From the recorded datasets on a cellular level, it can be concluded that the main adaptation regarding structure–property relationships toward the optimization of water transport and mechanical stability is mainly achieved at the cell level.     

Effects of genetic entry and competition by neighbouring trees on growth and wood properties of cloned Norway spruce (Picea abies)

? The aim of this work was to study both the effects of genetic entry and competition by neighbouring trees on growth, yield, wood density traits and fibre properties of 20 Norway spruce clones grown in an experimental trial located in southern Finland. The material included 10 Finnish clones, 2 Russian clones and 8 provenance-hybrids clones, the latter ones representing crosses between Finnish and foreign parent trees.

? We found that, compared to growth and yield, wood density traits and fibre properties showed, on average, lower phenotypic variations. Moreover, significant differences could be observed among the clones regardless of the trait (p < 0.05). Conversely, on average, no clear differences could be found between Finnish, Russian and provenance-hybrids clones in most of the studied traits.

? The marked differences among the clones, and lack of clear differences among the provenances suggested that any ranking, regarding different traits, should be based on individual clones. The phenotypic correlations between different traits ranged from moderate to strong suggesting that selection based on one trait would affect other traits. Regardless of clone, the growth of trees (e.g. breast height diameter) decreased and the wood density increased with increasing competition by neighbouring trees, which was expressed by competition index.

     

Variation in wood properties among five full-sib families of Norway spruce (Picea abies)

Genetic- and environmental variation and correlation patterns were characterized for modulus of elasticity (MOE), modulus of rupture (MOR) and related wood traits: latewood proportion, wood density, spiral grain, microfibril angle and lignin content in five full-sib families of Norway spruce. The families were evaluated on the basis of clearwood specimens from the juvenile -mature wood transition zone of 93 sampled trees at age 30 year from seed. Family-means varied significantly (p < 0.05) for all wood traits studied except lignin content. MOE varied between 7.9–14.1 GPa among trees and 9.4–11.0 GPa among families. MOR varied between 47–87 MPa among trees and 61–71 MPa among families. Families remained significantly different in an analysis of specific MOE (MOE/density) and MOR (MOR/density). Hence, solely relying on wood density as a wood quality trait in tree breeding would not fully yield the potential genetic gain for MOE and MOR. Correlations between wood structural traits and specific MOE and MOR are presented and discussed.