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.     

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.

     

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     

Cytology of Norway spruce needles

The structural changes in the albuminous, endodermal and mesophyll cells in the ageing needles of different aged healthy spruces (Picea abies [L.] Karst.) were studied on material collected in Germany and in Finland in winter. The results are considered as background information for the studies dealing with the structural changes in spruces suffering from biotic or abiotic stresses e. g., from the polluted air. Both light and electron microscopes were used.     

Crown morphology of Norway spruce from usual tree measurements

Intr0ducti0nNorwayspruce(PiceaabiesKarst)is0l1eoflI1e111ostimportal1tc011iferinFra11ce.lnl99l,tI1etotaIareaofNorwaysprLICewasestimatedt0be723000I11112.ItsvoIUmewasl27Inilli0I1m3.Tl1eanl1uaIvoILl111eiI1crc-mentwas5.74millionm3.ltiswideIyacceptedII1atIargeri11itialspacingssI1ouldbeusednow.l111940's,thedensity0ftl1epIantatiol1rangedfr0m5000to10000stems/hm2.Nowtl1einitiaIdensityvariesbetweenI000to250Ostems/I1m2.TI1eIargestspacit1g111aybe650~800stems/hm2.S0ithasasignifica11cetostLldyt…     

Choosing green sawing dimensions for Norway spruce from stochastic simulations

The high accuracy of log positioning and the stability of saw blades in breakdown machinery in modern sawmills have reduced the need to add margins for sawing variations. Oversize green sawing dimensions are still needed, but mainly to allow for drying shrinkage. This has put a new focus on better adapting green sawing dimensions to the shrinkage behavior of wood. In this study, a method for optimization of green sawing dimensions using stochastic simulation is presented. Normal distributions were generated for planed dry dimensions, kerf width, and target moisture content. The minimum share of boards exceeding the specified dry dimensions was decided, and deformations in boards from all positions in the cross section in a number of logs were simulated. The simulated shrinkage allowance from stochastic simulations was compared to experimental results from an industry test and to finite element results based on material data for Norway spruce. The results showed that the green width of the sawn boards should increase when the number of boards in the center yield increases. The green thickness of boards should be thinner for center boards and outer boards than for inner boards.     

Potential production of Norway spruce in Sweden

In this paper, the potential stem wood production for Norway spruce was estimated for different regions in Sweden. This was done by using basic physiological relationships of intercepted radiation versus biomass production and knowledge of how a water deficit reduces the potential production, derived from results of field experiments on nutrient optimisation. To scale these relationships up to regional and national levels, data of incident radiation and humidity during the growing season for all of Sweden were used. The figures for potential and attainable production indicate that the temperate to boreal climate allows considerably higher production than the current production, if availability of water and nutrients is non-limiting. In northern Sweden, the attainable production is ca. 300% higher than the current production and in southern Sweden, the yield can be increased by ca. 100%. In absolute numbers, as a mean for a whole rotation period, it is possible to achieve an annual stem-wood dry mass production of 7–9 Mg ha⁻¹ in southern and central and ca. 5–6 Mg in northern Sweden. This increased production would mean that rotation periods can be shorter than they are now by 20–30 years in southern Sweden and by ca. 50–60 years in northern Sweden.     

Pathogenicity of Ceratocystis resinifera to Norway spruce

The blue‐stain fungus Ceratocystis resinifera colonizes wounds on living Picea spp. and other conifers in Europe and North America. Little is known regarding the pathogenicity of this fungus and consequently, four Norwegian C. resinifera isolates were inoculated on to Norway spruce (Picea abies) using two different techniques. These included single‐point inoculations on young trees (two inoculations per tree on 14‐year‐old trees) and mass‐inoculations on older trees (～200 inoculations per tree on 34‐year‐old trees). In both experiments, C. resinifera induced minor symptoms that in most cases did not differ significantly from inoculation with sterile agar. The virulent blue‐stain fungus C. polonica, which was inoculated for comparative purposes, induced extensive symptoms, causing 83% dead cambium circumference and 82% blue‐stained sapwood, and long necrotic lesions in the phloem. The results suggest that C. resinifera is non‐pathogenic or only mildly pathogenic to Norway spruce and does not present a threat to these trees.     

Optimal management of uneven-aged Norway spruce stands

An optimization model is specified to analyze forest management without any restrictions on the forest management system. The data on forest growth comes from unique field experiments and is used to estimate a nonlinear transition matrix or size-structured model for Norway spruce. The objective function includes detailed harvesting cost specifications and the optimization problem is solved in its most general dynamic form. In optimal uneven-aged management, stand density is shown to be dominated by limitations in natural regeneration. If the goal is volume maximization, even-aged management with artificial regeneration (and thinnings from above) is superior to uneven-aged management. After including regeneration and harvesting costs, the interest rate, and the price differential between saw timber and pulpwood, uneven-aged management becomes superior to even-aged management. However, in the short term the superiority is conditional on the initial stand state.     

Performance of full-sib families of Douglas-fir in pure-family and mixed-family deployments

A major objective of tree improvement programs is to identify genotypes that will perform well in operational deployments. Relatively little is known, however, about how the competitive environment affects performance in different types of deployments. We tested whether the genetic composition and density of deployments affect the performance of full-sib families of Douglas-fir (Pseudotsuga meniziesii), and whether traits related to crown morphology could help to explain differences in family performance under competition. Seedlings from eight families were planted in pure-family and three mixed-family composition treatments at high, medium, and low densities (11,954, 2988, and 747 trees·ha⁻¹, respectively). Height (HT), diameter at breast height (DBH), and volume·ha⁻¹ (VOLHA) were measured at ages 8 and 15 years. Significant differences were found among composition treatments in all traits other than VOLHA8 and significant interactions between composition and density treatments were found for all traits at age 15 years. Family ranks for DBH15 and VOLHA15 in pure-family treatments changed considerably among densities, but ranks were more stable for HT15. The performance of two mixed-family treatments differed significantly from the average performances of the same families in pure-family treatments for several traits. Differences in DBH15 among families in high-density, pure-family treatments could be explained in part by differences in crown morphology, with better performance among families with relatively narrow crowns, stout branches, and high leaf area relative to branch length. Our results suggest that the competitive environment has a considerable effect on family performance, and that incorporating crown morphology traits into selection criteria in tree improvement programs may lead to greater productivity of Douglas-fir.     

Structural features of water-soluble arabinogalactans from Norway spruce and Scots pine heartwood

 Isolated water-soluble acidic arabinogalactans from Norway spruce and Scots pine heartwood were analysed and compared to Siberian larch heartwood arabinogalactans. The carbohydrate monomer composition was determined by acid methanolysis and gas chromatography, while structural studies were performed by ¹³C NMR spectroscopy and methylation analysis. The main structural features were found to be the same in the three types of arabinogalactans. However, the structure of the arabinogalactans from spruce and pine were found slightly different from the structure of larch arabinogalactans. The amount of single unit side-chains, consisting of arabinose and glucuronic acid units, was higher in the spruce and pine arabinogalactans than in the larch arabinogalactans. The amount of glucuronic acid was higher in the spruce arabinogalactans than in the pine arabinogalactans. The pine arabinogalactans had a higher amount of side chains with more than two sugar units than the spruce arabinogalactans. Received 30 December 1999     

Early selection improves clonal performance and reduces intraclonal variation of Norway spruce plants propagated by somatic embryogenesis

Height growth during the first and second growth periods (i.e., the June-September period in consecutive years) and intraclonal variation were assessed in 13 Norway spruce (Picea abies (L.) Karst.) clones propagated by somatic embryogenesis. The plants were acclimatized and grown in a greenhouse until mid-July and then transferred outdoors. The clonal mean heights after the first and second growth periods were lower for somatic embryo plants than for seedlings from corresponding families sown at the time of somatic embryo plant ex vitro transfer, because a large proportion of somatic embryo plants were small. We determined whether certain selection criteria at ex vitro transfer can be used to identify somatic embryo plants with height growth characteristics comparable with those of seedlings. Epicotyl length and presence of lateral roots proved to be important parameters for selection, whereas main root length was less useful. A combined selection for somatic embryo plants with lateral roots and with an epicotyl length exceeding 8 mm resulted in taller plants and reduced intraclonal variation after the first and second growth periods. The growth of somatic embryo plants selected in this way was similar to that of seedlings from the corresponding families. We conclude that selection according to these criteria at ex vitro transfer can result in improved performance of clonal stock propagated by somatic embryogenesis.     

Modeling individual tree height growth of Norway spruce and Scots pine from national forest inventory data in Norway

We developed individual tree height growth models for Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in Norway based on national forest inventory data. Potential height growth is based on existing dominant height growth models and reduced due to competition by functions developed in this study. Three spatially explicit and two spatially non-explicit competition indices were tested. Distance effects and diameter ratio effects were estimated from the data simultaneously with parameters of the potential modifier functions. Large height measurement errors in the national forest inventory data caused large residual variation of the models. However, the effects of competition on height growth were significant and plausible. The potential modifier functions show that height growth of dominant trees is largely unaffected by competition. Only at higher levels of competition, height growth is reduced as a consequence of competition. However, Scots pine also reduced height growth at very low levels of competition. Distance effects in the spatially explicit competition indices indicated that the closest neighbors are most important for height growth. However, for Scots pine also competitors at larger distance affected height growth. The five competition indices tested in this study explained similar proportions of the variation in relative height growth. Given that unbiased predictions can only be expected for the same plot size, we recommend a spatially explicit index, which describes the distance function with a negative exponential, for use in growth simulators.     

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.     

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.     

Isolation and characterisation of water soluble polysaccharides from Norway spruce and Scots pine

Water soluble polysaccharides from Norway spruce, Scots pine, and Siberian larch were compared. For all species the total amount of polysaccharides isolated from the heartwood was higher than that from the sapwood. The heartwood polysaccharides had a high content of galactose and arabinose units, and some glucuronic acid units, suggesting the presence of acidic arabinogalactans. The total amounts of recovered water-soluble arabinogalactans were 1.9 mg/g for spruce heartwood, 5.3 mg/g for pine heartwood, and as much as 106 mg/g for larch heartwood. The other water-soluble polysaccharides were mainly glucomannans. The average ratio of Gal:Ara:GlcA in the water-soluble arabinogalactans of spruce heartwood was about 4.3:1:1, pine heartwood about 4.5:1:0.2, and larch heartwood about 6.7:1:0.1. The corresponding molar ratios then being about 3.6:1:0.8 for spruce, 3.8:1:0.2 for pine, and 5.6:1:0.08 for larch. Thus, the content of glucuronic acid units was especially high in the spruce heartwood arabinogalactans. The content of arabinose was slightly higher in spruce and pine than in larch heartwood arabinogalactans.     

Growth of advance regeneration of Norway spruce after clear-cutting

We developed a basal area growth model for recovery of advance growth of Norway spruce trees after clear-cutting. Stem diameter growth at ground level and needle-mass characteristics were measured on permanent sample plots in Estonia. Both tree ring analysis (destructive sampling on one sample plot) and yearly repeated measurement data (two plots) were used to quantify advance growth. Basal area growth of small trees was estimated by multiple regression analysis. Previous-year basal area of the tree and basal area growth explained tree performance the next year. Tree needle-mass variables characterizing the acclimation status of the tree were included in the model as explanatory factors. Needle samples (one shoot from the upper third of each tree crown) were collected each year after the growth period from all sample trees. Needle masses of shoots from consecutive years were correlated and this variable was used as a predictor in the simulation model. Accelerating growth was observed in trees that exceeded the growth threshold in the year after release: the greater the needle mass per shoot, the greater the acceleration in growth. Competition among advance regeneration trees was included in the model: small trees under taller neighbors exhibited reduced growth. We found that trees released from a long period of heavy shade can survive, but the time needed for acclimation and resumption of competitive growth rates is considerably longer than for trees released from light shade. Such trees can be used for forest regeneration, but competition control (particularly reducing the proportion of fast-growing hardwoods) is required.