Simulation of effects of wood microstructure on water transport

A tracheid-level model was used to quantify the effects of differences in wood microstructure between coastal and interior Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii and var. glauca) wood on larger scale properties like hydraulic conductivity. The model showed that tracheid length, the ease of flow through a bordered pit and effective tracheid diameter can all limit maximum hydraulic conductivity. Among the model parameters tested, increasing bordered pit conductivity and tracheid length resulted in the greatest increase in maximum conductivity in both the inland and coastal ecotypes. A sensitivity analysis of the uncertainty between parameters governing flow through the bordered pit and air-seeding potential showed that, although decreased pit flow resistance increased maximum hydraulic conductivity, increased cavitation led to lower conductivity over time. The benefits of increasing the number of bordered pits depended on the intensity of the meteorological driving function: in drier environmental conditions, wood with fewer pits was more conductive over time than wood with more pits. Switching the bordered pit characteristics between coastal and interior wood indicated that the conductivity time course of coastal and interior wood was primarily governed by differences in the number of bordered pits and not differences in tracheid dimensions. The rate at which tracheids refilled had little effect on the conductivity time course of either coastal or interior wood during the first two summers when the wood was highly saturated, but had a marked influence in subsequent years once the cavitation profile stabilized. Our work highlights the need for more empirical work on bordered pits to determine whether variation in their number and properties is related to changing environmental conditions. In addition, a detailed simulation model of a bordered pit is needed to understand how variation in pit properties affects the relationship between ease of flow through a bordered pit and its potential for facilitating air-seeding.     

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.     

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。     

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.     

Distribution of lignin in normal and compression wood of Pinus taeda L.

Summary The distribution of lignin in normal and compression wood of loblolly pine (Pinus taeda L.) has been studied by the technique of lignin skeletonizing. Hydrolysis of the wood carbohydrates with hydrofluoric acid left normal wood tracheids with a uniform distribution of lignin in the S1 and S2 cell wall layers. However, the S3 region of both earlywood and latewood tracheids consistently retained a dense network of unhydrolyzable material throughout, perhaps lignin.Lignin content in compression wood averaged about 7% more than in normal wood and appears to be concentrated in the outer zone of the S2 layer. The inner S2 region, despite helical checking, is also heavily lignified. The S1 layer, although thicker than normal in compression wood tracheids, contains relatively little lignin.Ray cells, at least in normal wood, appear to be lignified to the same extent, if not more so in certain cases, than the longitudinal tracheids. Other locations where lignin may be concentrated include initial pit border regions and the membranes of bordered pits.This report is a detailed excerpt from the Ph. D. dissertation of R. A. P. Financial support provided by the College of Forestry at Syracuse University and the National Defense Education Act is hereby gratefully acknowledged.     

Behavior of the cellulose microfibril in shrinking woods

We measured the longitudinal and tangential shrinking processes in wood specimens from Chamaecyparis obtuse Endl. with different microfibril angles (MFAs). The shape of the shrinking curve was compared with the MFA. Only the longitudinal shrinking process of specimens with a small MFA clearly showed nonlinearity, and the degree of nonlinearity increased as the MFA decreased. In contrast, the tangential shrinking process and the longitudinal shrinking process of compression wood with a large MFA were linear. The nonlinearity is probably caused by the longitudinal shrinkage of the noncrystalline region of the cellulose microfibril (CMF) in regions of low moisture content during water desorption. When the moisture content is high, the matrix substance in the cell wall begins to dry; however, the shrinkage in the chain direction is restrained by the rigid CMF. As the wood dries further, the noncrystalline region of the CMF embedded in the matrix substance begins to shrink. Because the longitudinal mechanical behavior of wood with a small MFA is greatly affected by a rigid CMF, longitudinal shrinkage increases suddenly at about 10% moisture content; as a result, the shrinking process shows nonlinearity.     

Growth strain in the trunk and branches of Chamaecyparis formosensis and its influence on tree form

Distributions of growth strains in branches, straight trunks and basal sweeping trunks of Chamaecyparis formosensis Matsum. trees were measured with strain gauges. Microfibril angles (MFAs) of the S2 layer of the cell wall were measured by the iodine deposition method and their relationships with growth strain examined. The magnitude of the compressive stress on the lower side of trunks with a basal sweep was greater than that of the tensile stress at the surface of straight trunks. However, transverse compressive stress was similar around the trunk regardless of whether normal wood or compression wood was present. The released surface growth strains varied with MFA. At MFAs of 20-25 degrees , growth stress changed from tension to compression, and compressive stress increased dramatically in the compression wood region. Branches suffer bending stress due to self-loading. This stress is superimposed on the growth stress. Growth strains on the upper or lower sides of branches were larger than those in the trunks, suggesting that generation of growth stress on the lower sides of branches with extensive compression wood is affected by the gravitational bending stress due to self-loading. We conclude that branch form is affected by the interaction between the bending moment due to self-loading and that due to the asymmetric distribution of growth stress. Growth strain distribution in a branch differed depending on whether the branch was horizontal, upward bending or downward bending.     

微生物对长白鱼鳞云杉木材渗透性的影响

本文利用水池贮存处理木材研究了细菌对改善长白鱼鳞云杉木材渗透性的有效性。渗透性的测定采用水上升置换气流法。试验结果表明,水池贮存处理长白鱼鳞云杉气干材10周,不仅可以非常显著地增加其边材的渗透性(渗透性平均增加29倍),而且其心材渗透性亦有明显改善(平均渗透性增加1.52倍)。但心材试样渗透性增加不均匀,仅有50％的心材试样在水池贮存处理后渗透性得到了增加。渗透性增加的原因主要是由于边材和部分心材试样受到短芽孢杆菌(Bacillus brevis)等细菌的侵蚀,据电镜观察,边材大多数具缘纹孔的纹孔塞被细菌降解后形成空洞,心材的管胞内壁及具缘纹孔的纹孔缘上有细菌存在。     

Structure and function of flexure wood in Abies fraseri

Wood produced during flexure in one-year-old leaders of Abies fraseri (Pursh) Poir. (Fraser fir) was analyzed anatomically and radio-densitometrically. More xylem cells were produced in stems subjected to flexing than in stems that were not flexed. The lumens of tracheids produced in response to flexure were smaller than the lumens of tracheids in normal wood. This was manifest as an increase in the cell wall area/cell lumen area ratio. Microfibril orientation in flexure-induced wood approached the less extreme values found in compression wood. The growth ring composed of flexure-induced wood also had a greater density than normal wood. Compression wood, as defined by cellular characteristics observed in transverse section, was absent in flexed stems. Detailed analysis of the anatomical structure, wood density and biomechanical properties of flexure-induced wood indicated that it has more in common with compression wood than with normal wood.     

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     

Automated quantification of the impact of the wood-decay fungus Physisporinus vitreus on the cell wall structure of Norway spruce by tomographic microscopy

The visualization and the quantification of microscopic decay patterns are important for the study of the impact of wood-decay fungi in general, as well as for wood-decay fungi and microorganisms with possible applications in biotechnology. In the present work, a method was developed for the automated localization and quantification of microscopic cell wall elements (CWE) of Norway spruce wood such as bordered pits, intrinsic defects, hyphae or alterations induced by white-rot fungus Physisporinus vitreus using high-resolution X-ray computed tomographic microscopy. In addition to classical destructive wood anatomical methods such as light or laser scanning microscopy, this method allows for the first time to compute the properties (e.g., area, orientation and size distribution) of CWE of the tracheids in a sample. This is essential for modeling the influence of microscopic CWE on macroscopic properties such as wood strength and permeability.     

The variation of microfibril angle in South African grown Pinus patula and its influence on the stiffness of structural lumber

Reduction in the rotation ages of softwood saw-log plantations in South Africa is causing increased proportions of low stiffness sawn lumber at final harvest. It has been shown for some species that the microfibril angle (MFA) of the S2 layer of tracheids is strongly related to the modulus of elasticity (MOE) of wood, even more so than wood density, especially in wood formed during juvenile growth. The objectives of this study were to describe the variation in MFA in young Pinus patula trees and to determine the relationship between MFA and the dynamic MOE of sawn P. patula lumber. Thirty 16- to 20-year-old trees from six compartments from the Mpumalanga escarpment were processed into discs and lumber. The MFA, density and ring width were measured at two height levels using Silviscan 3. The average annual ring MFA varied between 7° and 29°; the pattern of variation depended mainly on height level and the ring number from the pith. The MFA in P. patula followed the same within-tree variation trends as in New Zealand-grown Pinus radiata but the average MFA was lower in absolute terms and differences between height levels were less pronounced. The MFA and density exhibited highly significant Pearson correlations of 0.73 and 0.70, respectively, with board dynamic MOE. A multiple regression model, which included MFA, density and ring width, explained 71% of the variation in the dynamic MOE of boards. A sensitivity analysis on the model showed that MFA and density had approximately similar influences on predicting the dynamic MOE of Pinus patula boards.     

Study of the transverse liquid flow paths in pine and spruce using scanning electron microscopy

Samples of pine (Pinus sylvestris) and spruce (Picea abies) were impregnated with a low-viscous epoxy resin using a vacuum process. The epoxy was cured in situ and the specimens sectioned. Deposits of the cured epoxy was then observed in the wood cavities using a scanning electron microscope. The investigation concentrated on tracing the transverse movements of a viscous liquid in the wood, and special attention was therefore given to the cross-field area between ray cells and longitudinal tracheids. A damage hypothesis is proposed based on the results obtained in the present investigation in combination with those from earlier studies on linseed oil-impregnated pine: In addition to the morphology of the bordered pits, viscous liquid flow in wood is dependent on damage that occurs during the impregnation procedure. For pine sapwood, liquid flow is enabled through disrupted window pit membranes, which divide the longitudinal tracheids and the ray parenchyma cells. A mechanism accounting for the reduced permeability of pine heartwood is believed to be deposits of higher-molecular-weight substances (extractives) in the ray parenchyma cells and on the cell walls. In spruce the thicker ray cells in combination with the smaller pits, which are connected to the longitudinal tracheids, reduce permeability considerably.     

Radial and between-family variations of the microfibril angle and the relationships with bending properties in Picea jezoensis families

With emphasis on tree breeding for wood quality in Picea jezoensis, we aimed to evaluate radial and between-family variations in the microfibril angle (MFA) of the S₂ layer in the latewood tracheids in 10 open-pollinated families of 43-year-old P. jezoensis trees. In addition, the relationships between MFA/wood density with the modulus of elasticity (MOE) or modulus of rupture (MOR) were investigated. Significant differences in MFA between families were found from the pith toward the bark. MFA showed higher values around the pith area, although some families showed relatively lower values than others around this area. In addition, due to a larger coefficient of variations of MFA near the pith, the potential for juvenile wood MFA improvement may be greater compared with mature wood. MOE was correlated with MFA in juvenile wood and with wood density in mature wood, whereas MOR was mainly correlated with wood density at radial positions in both woods. Therefore, to improve the MOE and MOR of P. jezoensis wood, both MFA and wood density would be factors to consider in both juvenile and mature woods. On the other hand, there are indications that, only wood density would be an important criterion for improving mature wood properties.     

Variability of the chemical composition of pit membranes in bordered pits of gymnosperms

Summary The pit membranes of the bordered pits in about one hundred coniferous wood species were investigated in regard to their chemical composition, in particular to the aromatic compounds. In many species pit membranes, even in the sapwood, contain phenolic substances. In heartwood, normally lignification takes place besides the development of other polyphenols. The variability of these compounds, both in sapwood an in heartwood, may be considerable not only between species but also within the same species and even between neighbouring tracheids and pits respectively.We are thankful to Prof. Dr. W. Liese for his support and his frequent discussions.     

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.     

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.     

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.     

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.     

Peroxidase activity, isoenzymes and histological localisation in sapwood and heartwood of Scots pine (Pinus sylvestris L.)

Peroxidase activity and isoenzymes of fresh wood samples of the third shoot of 12-year old trees and from the sapwood, transition zone and heartwood of c. 60-year old stems of Scots pine (Pinus sylvestris L.) were investigated. Wood samples were ground at −30°C, extracted, and the extracts concentrated c. 20-fold for peroxidase activity assays (guaiacol method) and for IEF-PAGE. At least 11 major isoenzymes could be found in the gels. Even the heartwood contained some peroxidase isoenzymes. Isoenzyme patterns of the juvenile wood did not change with the season. However, juvenile wood showed the highest peroxidase activity at the end of the growing season. Peroxidase activity decreased from the outer sapwood towards the heartwood. Thin sections of different wood zones stained for peroxidase revealed activity in ray parenchyma and resin canal epithelial cells. Intensive staining was localised in the bordered pits of vertical and ray tracheids, and in the end walls of ray parenchyma cells.