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.     

Fertilization effects on mean stomatal conductance are mediated through changes in the hydraulic attributes of mature Norway spruce trees

Stomatal conductance was quantified with sap flux sensors and whole-tree chambers in mature Norway spruce (Picea abies (L.) Karst.) trees after 3 years of exposure to elevated CO(2) concentration ([CO(2)]) in a 13-year nutrient optimization experiment. The long-term nutrient optimization treatment increased tree height by 3.7 m (67%) and basal diameter by 8 cm (68%); the short-term elevated [CO(2)] exposure had no effect on tree size or allometry. Nighttime transpiration was estimated as approximately 7% of daily transpiration in unchambered trees; accounting for the effect of nighttime flux on the processing of sap flux signals increased estimated daily water uptake by approximately 30%. Crown averaged stomatal conductance (g(s)) was described by a Jarvis-type model. The addition of a stomatal response time constant (tau) and total capacitance of stored water (C(tot)) improved the fit of the model. Model estimates for C(tot) scaled with sapwood volume of the bole in fertilized trees. Hydraulic support-defined as a lumped variable of leaf-specific hydraulic conductivity and water potential gradient (K(l)DeltaPsi) -was estimated from height, sapwood-to-leaf area ratio (A(s):A(l)) and changes in tracheid dimensions. Hydraulic support explained 55% of the variation in g(s) at reference conditions for trees across nutrient and [CO(2)] treatments. Removal of approximately 50% of A(l) from three trees yielded results suggesting that stomatal compensation (i.e., an increase in g(s)) after pruning scales inversely with K(l)DeltaPsi, indicating that the higher the potential hydraulic support after pruning, the less complete the stomatal compensation for the increase in A(s):A(l).     

Importance of soil extractable phosphorus distribution for mature Norway spruce nutrition and productivity

Phosphorus is an essential nutrient for forest growth. In this study, we assessed the impact of soil extractable phosphorus using two simple extraction methods on nutrition and productivity of Norway spruce in sixteen mature forest stands on different bedrocks and soils in Bavaria, Southern Germany. Representative trees were sampled for needles, twigs, branches, stem bark, and stem wood. Total phosphorus content in the tree parts and soil phosphorus stock extractable with citric acid and sodium bicarbonate up to a soil depth of 80 cm were determined. We found that easily soil extractable phosphorus is a suitable indicator for estimating phosphorus uptake and stand productivity in Norway spruce. In contrast, organic layer phosphorus showed no significant correlation with aboveground biomass phosphorus contents. In the biomass, the highest phosphorus contents were measured in young needles and twigs, but the highest correlation with soil phosphorus was detected for phosphorus contents in needles and bark. The stock of phosphorus extracted by citric acid down to 40 cm soil depth revealed the best correlation with phosphorus in needles and bark. Therefore, as a supplemental or alternative method to needle analysis, our study suggests the use of phosphorus contents in stem bark to evaluate tree phosphorus nutrition. These results highlight the suitability of the citric acid soil extraction method to characterize plant available phosphorus in Norway spruce ecosystems.     

Comparison of methods to quantify loss of hydraulic conductivity in Norway spruce

? Percent loss of hydraulic conductivity (PLC) is an important measure in plant water relations, but the available methods differ and results have rarely been compared.

? We compared PLC measured in Norway spruce (Picea abies) by quantifying conductivity before and after removing emboli, either by flushing with high water pressure or by infiltration under a partial vacuum, with relative water loss and staining conductive xylem to test advantages and possible problems of commonly applied methods.

? Infiltration removed nearly all gas from sections of drought-stressed and unstressed twigs, and flushing and infiltration efficiently removed emboli. Infiltration appears less prone to producing clogged xylem elements than high pressure flushing. Relative water loss (RWL) and the proportion of xylem stained with phloxine B (PSX) was also highly correlated with PLC, the latter can be quantified by image analysis, and also shows the pattern of xylem dysfunction. Loss of conductivity was more common in the inner annual rings, in early wood within an annual ring, and in compression wood, though pattern differed substantially between branches.

? Advantages and potential problems of these methods are discussed and it is suggested that RWL or PSX may be used as proxy measures for PLC in species when the correlations have been established.

     

Effects of experimental drought on the fine root system of mature Norway spruce

Norway spruce (Picea abies (L.) Karst.) is an economically important, but relatively drought-sensitive tree species that might suffer from increasing drought intensities and frequencies, which are predicted to occur in parts of central Europe under future climatic change. In a throughfall exclusion experiment using sub-canopy roofs, we tested the hypothesis that enhanced drought leads to an increase in fine root mortality, and also to a higher, subsequent fine root growth. Fine root production and mortality were assessed using two independent approaches, sequential soil coring (organic layer) and direct observations in minirhizotrons (organic layer plus upper mineral soil). Six weeks of throughfall exclusion resulted in mild drought stress, i.e. a reduction in average soil moisture from 20 to 12 vol.% during the treatment. Based on the sequential coring data, experimental drought did not result in significant changes in fine root biomass during the 6-week treatment period, but caused an increase in fine root mortality by 61% in the 6 weeks following the drought treatment. Remarkably, fine root production showed a synchronous increase in this period, which more than compensated for the loss due to increased mortality. The minirhizotron data confirmed that the drought treatment increased fine root loss in the organic layer. Based on this method, however, root loss occurred during the drought period and was not compensated by increased root production. The mild drought stress was mainly restricted to the organic layer and did not significantly influence fine root dynamics in the mineral soil. We calculated that the drought event resulted in an extra input of about 28 g C m⁻² and 1.1 g N m⁻² to the soil due to increased fine root mortality. We conclude that even periods of mild drought significantly increase fine root mortality and the associated input of root-derived C to the soil organic matter pool in temperate Norway spruce forests.     

Hydraulic and stomatal adjustment of Norway spruce trees to environmental stress

A study of how the water conducting systems of 30-50-year-old Norway spruce (Picea abies (L.) Karst.) trees growing at three sites adjust to shade and waterlogging indicated that water relations characteristics varied with the life histories of the trees. Xylem was more efficient at conducting water and stomata were more sensitive to atmospheric evaporative demand in trees subjected to favorable growth conditions (control trees) than in trees growing in shade or waterlogged conditions. At the same soil water availability, shade-grown trees suffered more severely from water deficit than control trees. Under conditions of high atmospheric vapor pressure deficit, foliage of shade-grown trees exhibited low water potentials, as a result of low hydraulic conductance of the vascular system and inefficient stomatal control. Because of the increased internal resistance to water flow, more negative leaf water potentials (Psi(x)) must be reached to provide an adequate water supply to the foliage. It is concluded that dynamic water stress is one of the main causes of the continuing growth retardation in suppressed Norway spruce trees after their release from the overstory. Trees growing in waterlogged soil (bog-grown trees) were characterized by weak stomatal control, resulting in large water losses from the foliage. Although bog-grown trees exhibited uneconomical water use, they possessed mechanisms (e.g., osmotic adjustment) that allowed leaves to tolerate low Psi(x) while stomata remained open. Under conditions of sufficient soil water availability and moderate atmospheric vapor pressure deficit, soil-to-leaf conductance was highest in bog-grown trees (1.45 +/- 0.06 mmol m(-2) s(-1) MPa(-1)), followed by control and shade-grown trees (1.04 +/- 0.04 and 0.77 +/- 0.05 mmol m(-2) s(-1) MPa(-1), respectively). The lowest soil-to-leaf conductance (0.45 +/- 0.04 mmol m(-2) s(-1) MPa(-1)) was recorded in control trees at high atmospheric evaporative demand, and was probably caused by tracheid cavitation.     

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.     

Interactions between soil scarification and Norway spruce seedling types

Interactions between scarification treatments and seedling types of Norway spruce were examined at two different locations in southern Sweden. The scarification treatments were not scarified control, mounding and soil inversion and the seedling types were a 10 week-old containerized seedling (mini seedling), a 2 year-old containerized seedling and a 2 year-old hybrid seedling, grown as a containerized seedling the first year and as a bare-root seedling the second year. Site preparation intensity had a greater effect on the mini seedlings. The results indicated that mini seedlings established faster in the soil inversion treatment compared to the larger seedling types. At the same seedling age, the mini seedlings had faster or similar growth rate as the containerized and the hybrid seedlings regardless of scarification treatment. Although mini seedlings grew as well as or even better than larger seedlings if they were successfully established, they were more sensitive to their planting environment and proper handling was critical. Problems with frost heaving and competing vegetation have to be taken into consideration when choosing site preparation method and seedling type.     

Growth recovery of mature Norway spruce and European beech from chronic O<Subscript>3</Subscript> stress

Elevated O₃ levels can strongly impair the health and vitality of forest ecosystems. Free-air exposure systems reveal that forest tree and stand growth can be reduced strongly under chronic O₃ stress. Detailed knowledge of the effect of O₃ exposure on photosynthesis, carbon sequestration, allometry and growth during chronic stress is available. However, knowledge of growth response after O₃ reduction is scarce. Here, we analyse the growth of mature Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) in the free-air O₃ fumigation experiment at Kranzberg Forest. We compare tree growth over a 9-year period (2008–2016) after exposure to O₃ (2000–2007). During 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech decreased by 24 and 32%, respectively. After cessation of 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech not only recovered but exceeded the growth of the trees in the control condition by 14 and 24%, respectively. The growth resilience and resistance of trees previously exposed to 2?×?O₃ towards drought stress and late frost was hardly lower than that of the trees in the control condition. The capacity for growth recovery even after long-term chronic O₃ stress emphasizes the strong beneficial effect of air pollution control on the health of forest ecosystems and on the global land carbon sink.     

Influence of nitrogen and phosphorous availability and ozone stress on Norway spruce seedlings

Four-year-old Norway spruce (Picea abies L. (Karst.)) seedlings were exposed to ambient and elevated (1.5 x ambient in 1997 and 1.6 x ambient in 1998) ozone concentrations [O3] and three nitrogen (N) and two phosphorus (P) availabilities: "optimal" values (control); 70% of the control N and P values (LN and LP); and 150% of the control N value (HN). Treatments were applied in an open-field ozone fumigation facility during the 1997 and 1998 growing seasons. Effects on growth, mineral and pigment concentrations, stomatal conductance and ultrastructure of needles were studied. The HN treatment increased growth significantly, whereas elevated [O3] had a slight or variable impact on growth and biomass allocation in all N treatments. Although there were no significant effects of the LP treatment on plant growth during the second year, there was a reduction in 1-year-old shoot dry mass in the elevated O3 + LP treatment at the end of the experiment. There were no significant treatment effects on mineral concentrations of current-year and 1-year-old needles at the final harvest. In response to the HN treatment, chlorophyll a and b and carotenoid concentrations increased significantly in current-year needles. Chlorophyll a/b ratio decreased in response to elevated [O3] alone, but increased in seedlings in the O(3) + LP treatment. Stomatal conductance of current-year needles decreased with increasing N availability, but increased in response to elevated [O3]. However, the O3-induced increase in stomatal conductance was less in the LN and LP treatments than in the control treatment. In chloroplasts of current-year needles, increased N availability decreased mean starch grain area, but increased the number of plastoglobuli. We conclude that Norway spruce seedlings are relatively tolerant to slightly elevated [O3], and that nitrogen and phosphorus imbalances do not greatly affect the influence of O3 on this species when the exposure lasts for two growing seasons or less.     

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     

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.     

Differences in wood properties between juvenile wood and mature wood in 10 species grown in China

 This study examined the intrinsic differences in various wood properties between juvenile wood and mature wood in 10 major reforestation species in China. Comparisons between juvenile wood and mature wood were made in both plantation- and naturally-grown trees. Considerable differences in most wood properties were found both between plantation-grown juvenile wood and mature wood, and between naturally-grown juvenile wood and mature wood. This suggests that wood properties of plantation-grown trees, to a large extent, depends on their juvenile wood contents, and can thus be manipulated effectively through rotation age. In general, the longer the rotation age, the lower the juvenile wood content, and the stronger the mechanical properties of the plantation-grown woods. However, the differences between juvenile wood and mature wood vary with wood properties and species. In general, juvenile wood and mature wood have less difference in chemical composition than in anatomical and physico-mechanical properties. Compared to the softwoods studied, the hardwoods appear to have less difference between juvenile wood and mature wood. Received 25 June 1999     

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.     

Response of mature stands of Norway spruce (Picea abies) to boron fertilization

The effects of boron (B) fertilizer applied 10 growing seasons earlier were studied in mature Norway spruce (Picea abies (L.) Karst.) trees in long-term factorial fertilization experiments at two field sites. Needle nutrient status, above-ground and below-ground growth and δ¹³C and carbon concentrations in the annual rings were measured. Needle B concentrations varied between 4 and 19 mg kg⁻¹ on the plots that had not received B fertilization. On the B-fertilized plots they varied between 15 and 39 mg kg⁻¹. The lowest B concentrations were on the plots that had received N or NCa fertilization. Needle Mn and Zn concentrations were lower on the B plots than on the plots that had not received B fertilization, although not significantly. Mean annual volume growth was slightly higher on the B plots at the more fertile site, but not at the less fertile one. The living:dead fine root mass ratio and living fine root length were also higher on the B-fertilized plots than on the unfertilized plots, but δ¹³C was not significantly affected, suggesting that the water status of the trees was not markedly altered by the increase in root growth. The carbon concentration in the annual rings was higher in the B-fertilized trees than in the unfertilized ones, suggesting the importance of B for wood formation.     

Three mechanisms affecting the mechanical properties of spruce wood dried at high temperatures

Wood drying experiments are conducted in which the temperature and the drying rate are controlled independently. In relationship to drying processes, at least three mechanisms are believed to contribute to the properties of dried wood. However, only two of these are found to affect the properties of macroscopic specimens, the third mechanism being observable in microtomed earlywood sections, and possibly in specimens loaded in the radial direction. Degradation of structural components and irreversible hydrogen bonding (hornification) are found to contribute to both the hygroscopicity and the mechanical properties of macroscopic wood specimens. Mass loss from thermal degradation occurs predominantly in slow high-temperature drying processes. Irreversible hydrogen bonding takes place in high-temperature drying, in particular with high ultimate dryness. Regarding the effect on strength and stiffness, mass loss and hornification appear to compete. The third identified mechanism, microscopic cell wall damage caused by incompatible drying shrinkage of cell wall elements, does not seem to affect the mechanical properties of macroscopic wood specimens. Consequently, slow high-temperature drying processes do not provide much benefit regarding the mechanical behavior of dried wood. The reasons for this are discussed.     

Relationships between respiratory activity and water-soluble sulfhydryl content of Norway spruce needles

The rate of respiration of spruce needles was affected by the level of foliar glutathione. At 37 degrees C the rate of respiration increased with increasing water-soluble sulfhydryl content up to 1.3 micromol g(-1) fresh weight. Higher thiol concentrations inhibited the rate of respiration. At 25 degrees C no relationship was observed between glutathione content and rate of respiration. It was concluded that the mechanism by which SO(2) affects plants is temperature dependent.     

Aging law of spruce wood

Abstract

Accelerated aging of spruce wood samples have been carried out by thermo-hydro (TH) treatments. These treatments were applied to accelerate the chemical reactions that take place during the natural aging of wood. In order to avoid dissimilar chemical reactions between the TH treatments and the natural aging, mild temperatures (between 100 and 150°C) have been selected at low relative humidity (RH). The mechanical properties of non-aged, natural aged and accelerated aged spruce wood have been compared. It is apparent that longitudinal Young's modulus of accelerated aged wood increase slightly at the beginning of the treatment and is followed by a reduction. Along the radial direction, Young's modulus remains almost unchanged. On the other hand the radial strength is severely reduced. From these results, the relative radial strength has been fitted on the chemical kinetic law. The rate constant of this law has been calculated and the treatment temperature and wood moisture content have been integrated. Finally this law has been extrapolated to standard climatic conditions in order to predict the loss of strength of old wood by knowing its age and its mean climate history of temperature and RH (ambiance condition).