人工兴安落叶松次生木质部的解剖学研究

运用木材解剖图像分析系统和显微照相的方法对人工兴安落叶松次生木质部的解剖结构进行研究,结果表明:落叶松具正常树脂道和受伤树脂道两种类型,前者常见于晚材。落叶松生长轮内的早晚材在干和枝内急变,在根内缓变。早材管胞呈六边形至多边形,胞壁常见单列具缘纹孔,偶见对列具缘纹孔;晚材管胞多呈矩形,胞壁鲜见具缘纹孔,通常为单列具缘纹孔。落叶松木射线同时具有单列木射线和纺锤形木射线两种类型,纺锤形木射线中仅含一枚纵行树脂道。纵行管胞与木射线交叉形成的纹孔场为云杉型。从根到干再到枝,管胞逐渐细化,管胞长度逐渐减小,木射线分布由密到疏。     

A model describing axial flow of liquids through conifer wood

Summary A mathematical model has been derived for the prediction of the resistance to viscous liquid flow generated by tracheid lumina and various parts of the bordered pit structure. The model also takes into account changes in pit geometry occurring as the pit membrane deflects when a pressure differential is applied across it. Methods for checking whether flow is truly viscous are presented.Data calculated for Pinus sylvestris suggest that the permeability of earlywood differs markedly from that of latewood; that in latewood the pit apertures contribute significantly to the total resistance to flow; and that kinetic energy corrections to the Poiseuille viscous flow equation may be of some importance at high flow rates.The authors wish to acknowledge the encouragement of Professors Matthews and Roche.     

Ultrastructure of pits inPinus banksiana lamb

Summary The cross-sectional view of pitting between various cell types inPinus banksiana Lamb. was studied at the ultrastructural level. Cell types inPinus banksiana include longitudinal tracheids, ray tracheids, ray parenchyma cells, buffer cells and epithelial cells. Two common characteristic features of bordered pit-pairs between longitudinal tracheids are an initial pit border and a thickened torus at the center of the pit membrane. The shape and size of the pit border and torus of bordered pit-pairs between two compression wood cells, and between the last-formed latewood longitudinal tracheid and first-formed earlywood longitudinal tracheid were different from those in the earlywood and latewood longitudinal tracheids. The pit border on the ray tracheid side varied in size and shape due to wall dentation. No initial pit border was found on the pit border of the ray tracheid side. The shape of bordered pit-pairs between two ray tracheids varied considerably due to irregularity of the dentate cell wall. The size of bordered pit-pairs in longitudinal tracheids was between 16 m to 20 m, which was twice the diameter of bordered pit-pairs in ray tracheids. Bordered pitpairs at the end wall of two ray tracheids appeared to be the smallest at 5 m, Pit aspiration occurred in the bordered pit-pairs with or without a torus. In the heartwood zone, some half-borders pit-pairs between tracheary and ray parenchyma cells showed an additional secondary wall on the ray parenchyma cell side. Plasmodesmata were found in the half-bordered pit-pairs as well in the simple pit-pairs. Blind pits were observed between a ray tracheid and a longitudinal tracheid. Bordered pit-pairs between two buffer cells were also observed. The possible functions of buffer cells were discussed.Use of transmission electron microscope provided by the Science Instrumentation Lab, Lakehead University and the technical assistance provided by Mr. A. MacKenzie, Director of Science Instrumentation Lab are gratefully appreciated     

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.     

Studies on opposite wood in conifers part II: Histology and ultrastructure

Summary A study has been made of the histology and ultrastructure of opposite wood in Larix laricina, Picea rubens, and Pinus resinosa. The width of the growth rings varied considerably, in one case from 0.1–1.0 mm, with the wide rings containing a much higher proportion of latewood than the narrow ones. The earlywood tracheids were square in outline and more regularly arranged than in normal wood. In the latewood they were sometimes irregular and distorted. The S₃ layer in the tracheids was 0.2 m thick in the earlywood and 0.4–0.8 m in the latewood, as compared to a thickness in normal wood of 0.1–0.2 m in both zones. The S₃ was often buckled in the latewood and was terminated towards the lumen by a spiral thickening. The cell wall structure of the tracheid pit border was described. Normal coniferous wood might be regarded as an intermediate between opposite wood and compression wood.This paper is dedicated to Dean Edwin C. Jahn in honor of his 70th birthday.     

Diffusion of non-swelling gases through dry conifer wood

Summary The conditions under which mutual, Knudsen and transition-range diffusion occur are described, and the corresponding gaseous diffusion coefficients are stated. The coefficient of hindered diffusion through pit membrane pores is shown to be about one third of the mutual diffusion coefficient and not about one thirtieth as postulated by earlier workers. A new longitudinal diffusion equation is developed, and results of a new gas permeability technique are used to derive coefficients for Picea sitchensis and Abies grandis. Only a small proportion of the tracheids is assumed to be conducting, owing to pit aspiration. About 98% of the resistance to longitudinal diffusion is in the tracheid lumina, 1% in the pit apertures and 0.2% in the pit membrane pores. From similar considerations a new tangential diffusion equation is developed, and coefficients are calculated. Roughly 90% of the resistance to tangential diffusion occurs in the pit apertures, 10% in the pit membrane pores and only 0.3% in the tracheid lumina. The agreement between the theoretical and published experimental values is satisfactory. For tangential diffusion the agreement is closer by an order of magnitude than that obtained by earlier workers. The experimentally determined temperature dependence of the diffusion coefficients is shown to be consistent with the above results. A hindered diffusion coefficient of one thirtieth of the mutual coefficient would lead to a temperature dependence not observed experimentally.     

Relationships of climate and cell features in stems and roots of black spruce and balsam fir

? The anatomical differences of mature black spruces and balsam firs were examined at stem and root level in order to characterize their wood properties at cellular level and link these differences to climate.

? Anatomical variability of these species was evaluated in relation to climate data gathered from 2001 to 2004 during the cell enlargement (CE) and wall thickening and lignification (WTL) phases. Lumen area, single cell wall thickness and total tracheid radial diameter were analyzed and regrouped into earlywood and latewood.

? Results from a principal component analysis (PCA) indicated that both first eigenvectors account for 82% and 90% of total variance for CE and WTL respectively. These component factors revealed that precipitation, humidity and number of days with precipitation significantly influence the lumen area (p = 0.0168) and radial cell diameter (p = 0.0222) in earlywood. Significant differences were registered between species and tree parts (stem and root) for the lumen area, radial cell diameter and cell wall thickness in both earlywood and latewood.

? In our study, black spruce exhibited smaller tracheid size in both stem and roots compared to balsam fir. Furthermore, the lower amount of tracheids produced during the growing season and higher proportion of latewood ensure a higher wood density of black spruce. The influence of temperature on earlywood formation is significant, whereas no influence was observed on latewood.

     

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.     

日本落叶松无性系木材性质的遗传变异

对10个10年生日本落叶松无性系的木材基本密度、管胞参数进行了测定.结果表明:木材基本密度,早、晚材管胞宽度和早材长宽比无性系间差异显著;木材基本密度,早、晚材管胞长度,早晚材管胞宽度和早晚材长宽比径向变异模式相似,即从髓心向外以曲线形式不断增加,有时亦有起伏;早材从髓心向外以近似直线的形式缓慢增加,晚材从髓心向外以曲线形式增加,初期增加幅度较大,到一定年龄后趋于水平变化并略有波动;材质性状与树木年轮间的关系以对数方程、幂函数方程、指数方程拟合效果较好;除了晚材壁腔比和早材壁厚外,其它木材性质的重复力均在0.5以上,受中度或中度以上的遗传制约,按照20%的选择率,长宽比和晚材管胞长能获得较高的遗传增益.     

Elevated temperature and CO(2) concentration effects on xylem anatomy of Scots pine

We studied the effects of elevated temperature and carbon dioxide concentration ([CO(2)]) alone and together on wood anatomy of 20-year-old Scots pine (Pinus sylvestris L.) trees. The study was conducted in 16 closed chambers, providing a factorial combination of two temperature regimes and two CO(2) concentrations (ambient and elevated), with four trees in each treatment. The climate scenario included a doubling of [CO(2)] and a corresponding increase of 2-6 degrees C in temperature at the site depending on the season. Anatomical characteristics analyzed were annual earlywood, latewood and ring widths, intra-ring wood densities (earlywood, latewood and mean wood density), tracheid width, length, wall thickness, lumen diameter, wall thickness:lumen diameter ratio and mass per unit length (coarseness), and numbers of rays, resin canals and tracheids per xylem cross-sectional area. Elevated [CO(2)] increased ring width in four of six treatment years; earlywood width increased in the first two years and latewood width in the third year. Tracheid walls in both the earlywood and latewood tended to become thicker over the 6-year treatment period when temperature or [CO(2)] was elevated alone, whereas in the combined treatment they tended to become thinner relative to the tracheids of trees grown under ambient conditions. Latewood tracheid lumen diameters were larger in all the treatments relative to ambient conditions over the 6-year period, whereas lumen diameters in earlywood increased only in response to elevated [CO(2)] and were 3-6% smaller in the treatments with elevated temperature than in ambient conditions. Tracheid width, length and coarseness were greater in trees grown in elevated than in ambient temperature. The number of resin canals per mm(2) decreased in the elevated [CO(2)] treatment and increased in the elevated temperature treatments relative to ambient conditions. The treatments decreased the number of rays and tracheids per mm(2) of cross-sectional area, the greatest decrease occurring in the elevated [CO(2)] treatment. It seemed that xylem anatomy was affected more by elevated temperature than by elevated [CO(2)] and that the effects of temperature were confined to the earlywood.     

Morphological changes in sugi (Cryptomeria japonica) wood after treatment with the ionic liquid, 1-ethyl-3-methylimidazolium chloride

We investigated morphological changes in wood tissues of sugi (Cryptomeria japonica) resulting from treatment with the ionic liquid 1-ethyl-3-methylimidazolium chloride ([C2mim][Cl]), which dissolves cellulose. Treatment with [C2mim][Cl] caused dissociation and distortion of tracheids in latewood, but not in earlywood. This difference was due to the difference in swelling behavior of the cell wall between earlywood and latewood. Many pit membranes in bordered pits were broken by treatment with [C2mim][Cl]. In addition, some chemical changes in wood components, such as cellulose and lignin, occurred before significant disruption or destruction of the cell wall. Our results show that the reaction of wood liquefaction by [C2mim][Cl] treatment is not homogeneous, both from chemical and morphological viewpoints.     

Tangential pitting in black spruce tracheids

Tangential pit features were studied in a 55-year old black spruce (Picea mariana (Mill) B.S.P.) tree by means of light and electron microscopy.It was found that tangential pitting is lacking from the greatest part of the growth ring, except for the last four tangential rows of latewood tracheids and the first row of early wood tracheids. The average number of pits per tangential wall of a 3.55-mm-long tracheid is 234, 144, 28, 4 and zero, respectively, in the last 5 tangential rows of latewood tracheids, starting at the growth-ring boundary.On the average, tangential pits measure 5.4 m in diameter, possess oval to elliptical apertures, and are randomly distributed uniformly over the tangential tracheid wall. All tangential intertracheid pits are bordered and in that respect are similar to those in the radial walls. Although most of the pits contain membranes with tori, some at the growth-ring boundary lack tori and exhibit randomly oriented microfibrillar structure.     

Fractography of shear failure surface of softwood decayed by brown-rot fungus

In order to investigate shear failure surface of decayed wood, wood pieces of ezomatsu (Picea jezoensis) were exposed to brown-rot fungus (Fomitopsis palustris), and standard shear test in radial plane was conducted. The failure surfaces were examined by scanning electron microscopy and surface roughness measurements were also conducted. Transwall failure that the crack elongated parallel to the tracheid axis was observed in the earlywood region through all phases of decay. Intrawall failure principally occurred in the latewood region on the early phase of decay. When decay progressed considerably, transwall failure that the crack elongated perpendicularly to the tracheid axis occurred. And transwall failure was also dominant failure morphology in the latewood region. Fragments of tracheids which were partly peeled out from S₂ layers were observed in some specimens. Size and appearance frequency of fragments of tracheid were smaller and lower when decay progressed. Arithmetic average roughness, which was the index of fragment size and appearance frequency, had positive correlation to shear strength ratio. Especially, line surface roughness of radial direction, which was measured across the radial files of tracheid, had the highest correlation to the shear strength ratio. The surface roughness would be a good indicator to evaluate the decay degree.     

Cavitation in dehydrating xylem of Picea abies: energy properties of ultrasonic emissions reflect tracheid dimensions

Ultrasonic emission (UE) testing is used to analyse the vulnerability of xylem to embolism, but the number of UEs often does not sufficiently reflect effects on hydraulic conductivity. We monitored the absolute energy of UE signals in dehydrating xylem samples hypothesizing that (i) conduit diameter is correlated with UE energy and (ii) monitoring of UE energy may enhance the utility of this technique for analysis of xylem vulnerability. Split xylem samples were prepared from trunk wood of Picea abies, and four categories of samples, derived from mature (I: earlywood, II: 30-50% latewood, III: >50% latewood) or juvenile wood (IV: earlywood) were used. Ultrasonic emissions during dehydration were registered and anatomical parameters (tracheid lumen area, number per area) were analysed from cross-sections. Attenuation of UE energy was measured on a dehydrating wood beam by repeated lead breaks. Vulnerability to drought-induced embolism was analysed on dehydrating branches by hydraulic, UE number or UE energy measurements. In split samples, the cumulative number of UEs increased linearly with the number of tracheids per cross-section, and UE energy was positively correlated with the mean lumen area. Ultrasonic emission energies of earlywood samples (I and IV), which showed normally distributed tracheid lumen areas, increased during dehydration, whereas samples with latewood (II and III) exhibited a right-skewed distribution of lumina and UE energies. Ultrasonic emission energy was hardly influenced by moisture content until ～40% moisture loss, and decreased exponentially thereafter. Dehydrating branches showed a 50% loss of conductivity at -3.6 MPa in hydraulic measurements and at -3.9 and -3.5 MPa in UE analysis based on cumulative number or energy of signals, respectively. Ultrasonic emission energy emitted by cavitating conduits is determined by the xylem water potential and by the size of element. Energy patterns during dehydration are thus influenced by the vulnerability to cavitation, conduit size distribution as well as attenuation properties. Measurements of UE energy may be used as an alternative to the number of UEs in vulnerability analysis.     

Fibril angle of loblolly pine wood as related to specific gravity,growth rate,and distance from pith

Summary Fibril angles were greater for earlywood (avg. 33.4°) than for latewood tracheids (avg. 26.9°). For earlywood, fibril angle did not differ between growth rates when the specific gravity was low (avg. 33.3°). When the specific gravity was high, wood of fast growth had a higher fibril angle (avg. 35.1°) than wood of slow growth (avg. 32.0°). No differences were detected between core, middle, and outer wood. In latewood tracheids, fibril angles were greater in corewood (avg. 28.0°) than in middle or outer wood (avg. 26.3°). For whole wood (a weighted average of earlywood and latewood), fibril angle averaged 30.7° and was greater in corewood (avg. 32.2°) than in middle or outer wood (avg. 29.9°).     

Microfibril angle non-uniformities within normal and compression wood tracheids

The pattern and extent of variation of microfibril angle (MFA) in normal and compression tracheids of softwood were investigated by using confocal laser scanning microscopy technique. All measurements support the idea that the orientation of microfibrils in single wood tracheids is not uniform. MFA of the radial wall of earlywood tracheids was highly non-uniform and had an approximately circular form of arrangement around the bordered pits (inside the border). Between the bordered pits the measured MFAs were less than the other parts of the tracheid. In the latewood tracheids MFA was less variable. The average orientation of simple pits in the crossfield region was consistent with the mean MFA of the tracheids; however some of the measurements showed a highly variable arrangement in the areas between the simple pits. In many cases the local measured MFAs of compression wood tracheids agreed with the orientation of natural helical cavities of compression wood. Comparing the measured results in different growth rings showed that MFAs in juvenile wood are generally larger than in perfect wood.     

The cellular distribution of lignans in Tsuga heterophylla wood

Summary Western hemlock heartwood contains patches of tracheids with large amounts of cellular inclusions. Microscopic and chemical examination of the wood showed several types of deposits containing the lignans matairesinol, hydroxymatairesinol and conidendrin. The deposits, which were often relatively pure individual lignans, frequently assumed different physical forms and chemical composition. A check in the wood contained three distinct forms of deposits each of which was a different lignan. Rays contained deposits physically similar to those in adjacent tracheids but, while lignans were present in the tracheids, they were not detected in the rays. Lignans lined tracheid walls as surface films and often encrusted the bordered pits. The amount of lignans in the wood was not related to wet wood zones although surface films and pit encrustations should have an influence on physical properties. The location and physical nature of lignan deposits in western hemlock heartwood indicates their biosynthesis has probably taken place at the heartwood periphery in the vicinity of the half-bordered pit.Presented at the International Wood Chemistry Symposium, Seattle, U. S. A. September 1969.Requests for reprints should be sent to W. E. Hillis. We thank Miss J. Barratt for assistance with part of this work and Mr. C. J. Kozlik for collecting wood samples.     

Incorporation of transfer resistance between tracheary elements into hydraulic resistance models for tapered conduits

The model of West, Brown and Enquist (1999) shows that hydraulic resistance in trees can be independent of path length, provided that vascular conduits widen sufficiently from tree top to base. We demonstrate that this result does not depend theoretically on branching architecture or cross-sectional conductive area of the stem. Previous studies have shown that pit membrane resistance, encountered when water moves between either tracheids or vessels, accounts for up to 60% of the total resistance in stem segments. When pit membrane resistance, which is neglected by most whole-tree hydraulic models, was incorporated in hydraulic models in three different ways, the near invariance of hydraulic resistance was preserved. If relative pit resistance was independent of tracheid size or if tracheid dimensions were scaled to minimize wood resistivity, the minimum conduit taper required for path length independence equaled that in the original model of West et al. (1999). Under the most realistic model, in which relative pit resistance increased with tracheid radius, this value was doubled. Such taper is not possible within the typical size range of tracheids over the entire length of moderately tall trees, but it might be possible for vessel-bearing trees. Preliminary results indicated that although tracheid radius in the outer growth ring initially increased basipetally from the top of an 18-m tall Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), it stabilized at mid-trunk. Also, conduit taper was not constant in this species, violating a key assumption of the model of West et al. (1999), on which the invariance of hydraulic resistance depends.     

Relationships among selected wood properties of 20-year-old Taiwania (<Emphasis Type="Italic">Taiwania cryptomerioides</Emphasis>) trees

The relationships between bending properties, compressive strength, tracheid length, microfibril angle, and ring characteristics of 20-year-old Taiwania (Taiwania cryptomerioides Hay.) trees were examined. The trees came from different thinning and pruning treatments, but the practices showed no significant effect on the investigated properties. The results showed that based on comparison with the literature, plantation-grown immature Taiwania have noticeably lower average strength properties than mature trees of the same species. Wood density and bending and compressive strengths were not related to either tracheid length or microfibril angle in young Taiwania. There were positive relationships between bending strength and compressive strength. The wood density, ring width, earlywood width, earlywood density, and latewood percentage were the most important predictors of strength by simple linear regressions. The wood density and ring width/earlywood width may be considered as indicators for assessing the bending strength, while wood density and latewood percentage were the best predictors of compressive strength by multiple linear regressions.     

Effects of steam explosion on wood appearance and structure of sub-alpine fir

The technology of steam explosion was applied to pre-treat sub-alpine fir (Abies lasiocarpa (Hook.) Nutt.) lumber and to improve its drying characteristics. Effects of steam explosion on the appearance and structure of the lumber are discussed in this paper. The structure of the wood was examined using light microscope (LM) and scanning electron microscope (SEM). The following results were obtained. With increasing temperature, pressure, and explosion cycles the color of the lumber darkened gradually. No significant structural difference between treated and untreated samples was observed using LM when the treatments were carried out at temperatures below 130°C with ten explosion cycles (group A or B). Some fractures were observed in bordered pit pairs between tracheids after 20 explosion cycles at 130°C (group C). More fractures occurred in bordered pit pairs between earlywood tracheids at a temperature of 160°C (group D). More or less fractures in pits between ray parenchyma cells and earlywood tracheids were observed using SEM in all four cases of treatments. Although no change in bordered pits in the tracheid walls between group A and the control group was discovered, groups B, C, and D showed different extents of ruptures in bordered pits, which may lead to break aspirated pits and improve permeability. In these groups, wrinkles and separations in the inner tracheid walls and detachments in middle lamella also occurred and became more serious as temperature or cycles of the treatment increased.