Role of the gelatinous layer (G-layer) on the origin of the physical properties of the tension wood of Acer sieboldianum

The tension wood (TW) properties of a 70-year-old specimen of Acer sieboldianum Miq. were analyzed by using the G-fiber model that was proposed in our previous report. The roles of the G-layer on the origins of (1) a high tensile growth stress, (2) a large longitudinal Young’s modulus, and (3) a high longitudinal drying shrinkage in the TW xylem are discussed on the basis of the simulations using the G-fiber model. The results suggest that the G-layer generates a high tensile stress in the longitudinal direction during xylem maturation; the longitudinal Young’s modulus of the green G-layer becomes significantly higher than that of the lignified layer; furthermore, the G-layer tends to shrink extraordinarily more than that of the lignified layer during moisture desorption.This report follows the previous report “Role of the gelatinous layer on the origin of the physical properties of the tension wood.” J Wood Sci (2004) 50:197–208. Part of this paper was presented at the 49th Annual Meeting of the Japan Wood Research Society, Tokyo, April 1999, and at the 2nd International Conference of the European Society for Wood Mechanics, Stockholm, May 2003This revised version was published online in July 2005. On pages 228–230 the character was replaced by a vertical line.     

On the detachment of the gelatinous layer in tension wood fiber

The detachment of the gelatinous layer (G-layer), often observed on microtome cross sections, has led some authors to believe that the G-layer cannot act as the driving force of longitudinal shrinkage in tension wood. The aim of this study was to observe the detachment of the G-layer along fibers. Green wood blocks were cut transversely into two samples. One sample was kept in water and the other was oven-dried. With one face being common to both samples, the detachment of the G-layer was studied on the same fibers. Observations were performed after blocking deformation by embedding. This revealed that the detachment of the G-layer is an effect produced by the act of cutting the transverse face of the wood block to be embedded. At distances greater than 100 µm from this primary surface of the sample, no detachment was observed. Drying shrinkage shows little or no effect on this detachment. The result seems to explain well why the detachment of the G-layer occurs during sectioning using conventional sliding microtomy. These observations prove the adhesion of the G-layer in massive wood and confirm the active role of the G-layer in tension wood properties.     

Lignification in poplar tension wood lignified cell wall layers

The lignification process in poplar tension wood lignified cell wall layers, specifically the S(1) and S(2) layers and the compound middle lamella (CML), was analysed using ultraviolet (UV) and transmission electron microscopy (TEM). Variations in the thickness of the gelatinous layer (G-layer) were also measured to clarify whether the lignified cell wall layers had completed their lignification before the deposition of G-layers, or, on the contrary, if lignification of these layers was still active during G-layer formation. Observations using UV microscopy and TEM indicated that both UV absorbance and the degree of potassium permanganate staining increased in the CML and S(1) and S(2) layers during G-layer formation, suggesting that the lignification of these lignified layers is still in progress during G-layer formation. In the context of the cell-autonomous monolignol synthesis hypothesis, our observations suggest that monolignols must go through the developing G-layer during the lignification of CML and the S(1) and S(2) layers. The alternative hypothesis of external synthesis (in the rays) does not require that monolignols go through the G-layer before being deposited in the CML, or the S(1) and S(2) layers. Interestingly, the previous observation of lignin in the poplar G-layer was not confirmed with the microscopy techniques used in the present study.     

Origin of the characteristic hygro-mechanical properties of the gelatinous layer in tension wood from Kunugi oak (Quercus acutissima)

The mechanism responsible for unusual hygro-mechanical properties of tension wood containing the gelatinous layer (G-layer) was investigated. Tension and normal wood specimens were sampled from the leaning stems of a 75- and a 40-year-old Kunugi oak (Quercus acutissima) tree, and the moisture dependencies of the longitudinal Young’s modulus and longitudinal dimensions were measured. The results, which were analyzed in relation to the anatomical properties of the specimens, revealed that the ratio of increase in the longitudinal Young’s modulus with drying was higher in the G-layer than in the lignified layer (L-layer); the longitudinal drying shrinkage displayed a similar pattern. It was found that the lattice distance of the [200] plane in the cellulose crystallite increased with drying, moreover, the half-width of the [200] diffraction peak increased with drying, which was remarkable in the tension wood. Those results suggest that in the green state, the polysaccharide matrix in the G-layer behaves like a water-swollen gel; however, it is transformed into a condensed and hard-packed structure by strong surface tension during moisture desorption, which is a form of xero-gelation. However, in the L-layer, condensation and subsequent xero-gelation of the polysaccharide matrix was prevented by the hydrophobic lignin that mechanically reinforces the matrix.     

Structural, histochemical and chemical characterization of normal, tension and opposite wood of Subabul (Leucaena leucocephala (lam.) De wit.)

Structure, histochemistry and chemical composition of tension (TW), opposite (OW) and normal wood (NW) of Leucaena leucocephala have been studied using histological, histochemical and biochemical methods. TW vessels are longer and have a larger diameter than NW vessels, and the rays are shorter and thinner. The G-layer replaced S3 and parts of S2 wall layers in TW fibres. TW further contained less lignin that also had a lower syringyl-to-guaiacyl ratio. The lignin was also more condensed. The content of α-cellulose was high in TW. TW contains hemicelluloses and pectins composed of more galactose, and less xylose, mannose, 4-O-methylglucuronic acid and galacturonic acid compared to that of OW. This study suggests that in the TW of Leucaena, the amount and composition of lignin, hemicelluloses and pectins differ considerably from that in OW and NW.     

Properties of cell wall constituents in relation to longitudinal elasticity of wood

 To predict the origin of longitudinal elasticity of the solid wood in relation to the composite structure of the wood cell wall, an analytical procedure was developed on the basis of the idea of “the reinforced-matrix hypothesis” originally introduced by Barber and Meylan (1964). A multi-layered circular cylinder, having the CML, the S1, and the S2 layers, was used as a model of the ligno-cellulosic (wood) fiber, and the elastic properties of an isolated wood fiber were formulated mathematically. In the formulation, not only the structural factors, such as the microfibril angle and the thickness of each layer, but also the environmental condition, e.g. the moisture content, were taken into consideration. The effects of the moisture content and the microfibril angle upon the longitudinal Young's modulus and the Poisson's ratio of the wood fiber were simulated by using the newly derived formulae. It is anticipated to give a start to estimate the fine structure and the internal properties of the cell wall constituents in relation to the macroscopic behaviors of the wood through simulating the mechanical behaviors of the wood fiber. Received 17 August 1999     

Proteomic analysis of the G-layer in poplar tension wood

Angiosperm trees bend their stems by forming tension wood at the upper side of leaning stems. Most tension wood has a cellulose-rich G-layer in the innermost surface of the fiber cell wall. Strong tensile stress is considered to occur in the G-layer. This study undertook to identify the proteins involved in G-layer formation and function through a proteomic analysis of G-layer-localized protein. G-layers of poplar were loosened by sonication and isolated as doughnut-shaped pieces of thinly sliced transverse sections. The proteins, once extracted with urea/detergent solution, were separated by two-dimensional polyacrylamide gel electrophoresis, and 110 spots were subjected to liquid chromatography tandem mass spectrometry (LC/MS/MS). A database search for these spots’ mass spectrum patterns identified 72 proteins. In addition, all peptide digestion mixtures of G-layer proteins were separated by strong cation exchange chromatography and 39 proteins were identified using LC/MS/MS analysis. Proteins involved in wall formation, such as lignin biosynthesis-related protein, xyloglucan endotransglucosylase, and fasciclin-like arabinogalactan protein, were notably detected in the G-layer.     

Tensile growth stress and lignin distribution in the cell walls of black locust (Robinia pseudoacacia)

Seven specimens that contained a continuous gradient of wood from normal to tension wood were collected from an inclined black locust (Robinia pseudoacacia), and the released strain of growth stress was quantified. Lignin distribution in the cell wall was investigated using ultraviolet (UV) microspectrophotometry to examine its relation to the intensity of growth stress. The UV absorption at cell corner middle lamella and in the compound middle lamella remained virtually constant, irrespective of the contractive released strain (i.e., tensile growth stress). The gelatinous (G)-layer began to differentiate, and the UV absorption decreased there in accordance with increases in the contractive released strain. The absorption maximum (max) remained virtually constant at the cell corner middle lamella and in the compound middle lamella at 277–280nm, irrespective of the released strain. The max for the secondary wall of normal wood was 272nm and shifted to 268nm in the G-layer of tension wood as the contractive released strain increased. The percentage of the cross-sectional area, consisting of the G-layer, with respect to the whole cross-sectional area increased with the contractive released strain.     

杨木应拉木微区结构可视化及化学成分分析

木材微区结构与木材宏观性质密切相关,杨木应拉木与对应木宏观性质存在较大差别,探究杨木应拉木和对应木微区结构和化学成分,可为了解杨木应力木的宏观性质提供理论根据。借助光学显微镜、荧光显微镜、显微拉曼成像光谱仪、透射电镜对杨木应拉木微区结构进行可视化研究,并借助X射线衍射技术和美国可再生能源实验室方法,分析杨木应拉木的微晶尺寸、结晶度以及化学成分。结果表明:杨木应拉木中应拉区和对应区纤维细胞微区结构差异显著。光学显微镜下显示应拉区木纤维中胶质层清晰可见,荧光显微镜和拉曼显微镜下显示胶质层的木质素浓度比对应区低。透射电镜下显示应拉区木纤维细胞壁结构由初生壁、次生壁和胶质层组成,未见次生壁外层,各层的平均厚度分别为0.61,1.22和2.53μm。对应区木纤维为典型的初生壁和次生壁结构,次生壁各层平均厚度分别为0.33,2.28和0.14μm。杨木应拉区纤维素含量(58.91%)比对应区(41.53%)高,木质素含量和半纤维素含量均比对应区的低,应拉区木质素和半纤维素含量分别为21.99%和12.01%,对应区分别为28.10%和17.08%。杨木应拉区结晶度(48.06%)比对应区(41.01%)高,应拉区晶区宽度为2.66 nm,长度为8.84 nm;对应区晶区宽度为2.65 nm,长度为9.87 nm。     

栽培措施对短周期工业材材性影响规律的研究

对4种不同栽培措施的日本落叶松、湿地松、马尾松和杉木等多种短周期工业材309株样木的材性进行了全面测试分析研究，结果表明．立地指数大，木材生长率和纤维宽度大．但纤维长度、纤维长宽比、木材基本密度、顺纹抗压强度和抗弯弹性模量减小；立地指数相差两级，木材的年龄宽度、纤维宽度、长度和微纤线角度的差异达到显著或极显著水平栽植密度小，木材的年轮宽度和干缩比增大但纤维长宽比及体积全干缩率减小；栽植密度增大，杉木和昭林6号杨木材的力学强度提高，但尾叶桉和I－69杨降低；不同栽植密度的木材pH值及酸碱总缓冲容量，都是心材大于边材，但木材的脲醛树脂胶凝速度，则是边材快于心材.与未间伐林分的木材比较，随着间伐强度的增大、年轮宽度增大．但纤维长宽比和微纤丝角度减小；间伐强度增大，针叶树村的力学强度降低，阔叶树树则提高．但将强度间伐材（间代40％）与未间伐材比较，结果完全相反；北京杨木材心材的pH值、酸碱缓冲容量及总容量，均随间伐强度的增大而增大，但边材却减小．施肥措施能够增大纤维直径，但将导致纤维长宽比、木材密度和力学强度的降低施肥可以使树木速生，但会降低木材密度和质量，在培育建筑结构材的林分应慎用．     

Effect of Wood Variables on the Properties of Wood Fiber-Polypropylene Composites

The effect of wood species (Chinese fir and Poplar), wood fiber content (10%, 25%, 40%) and wood fiber sizes (16 to 32 mesh, 32-65 mesh, above 65 mesh) on the properties of the wood fiber-Polypropylene composites were studied in this paper. The results indicate that the effect of wood fiber content and size in composite were more important than that of chosen wood species. Compared with polypropylene without wood fiber, the flexural strength of the composites increased when adding wood fiber into polypropylene, but the tensile and unnotched charpy impact strength decreased. And the above strength decreased with the wood fiber content increasing. When the wood fiber size becoming smaller (in higher mesh), the strength increased. In the comparison of wood species, the properties of composite using Chinese fir wood were better than that of Poplar, but not significant. The dynamic mechanical properties of the composites and PP were also tested and analyzed in this paper.     

Anatomy and lignin distribution of reaction wood in two Magnolia species

Summary Anatomical features of reaction wood formed in two Magnolia species, M. obovata Thunb. and M. kobus DC. which are considered to be among the primitive angiosperms, were observed. In addition, the distribution of guaiacyl and syringyl units of lignins in the cell walls of normal and reaction wood was examined using ultraviolet (UV)- and visible light (VL)- microspectrophotometry coupled with the Wiesner and M?ule reactions. The two Magnolia species formed a tension-like reaction wood without possessing the typical gelatinous layer (G-layer) on the upper side of the inclined stem or branch, in which a radial growth promotion occurred. Compared with the normal wood, the reaction wood had the following anatomical features: (1) the secondary walls of fiber tracheids lacked the S₃ layer, (2) the innermost layer of fiber-tracheid walls showed a small microfibril angle, a fact being similar to the orientation of the microfibril angle of the G-layer in tension wood, and (3) the amounts of lignin decreased in the cell walls of fiber tracheids, especially with great decrease in proportion of guaiacyl units in lignins. In addition, VL-microspectrophotometry coupled with the Wiesner and M?ule reactions adopted in the present study showed potential to estimate the lignin contents in the cell walls and the proportion of guaiacyl and syringyl units in lignins. Received: 15 July 1998     

The effect of sectioning and ultrasonication on the mesoporosity of poplar tension wood

Increasing interest in understanding tension stress generation in tension wood with fibres having a gelatinous layer (G-layer) has focused attention on the specific role of this layer. To distinguish its contribution from those of other wall layers, the G-layer of wood sections was isolated by ultrasonication. The aim of this study was to assess the effect of sectioning and of the ultrasonic treatment on the mesoporosity of tension wood using nitrogen adsorption–desorption analysis. The results showed that the process of isolating the G-layer using ultrasonication strongly affects its mesoporosity. Most damage was found to occur during sectioning rather than as a result of the 15-min ultrasonic treatment.     

木纤维PP/PE共混物复合材料的流变和力学性能(英文)

For evaluation of the rheological and mechanical properties of highly filled wood plastic composites （WPCs）, polypropylene/polyethylene （PP/PE） blends were grafted with maleic anhydride （MAH） to enhance the interfacial adhesion between wood fiber and matrix. WPCs were prepared from wood fiber up to 60 wt.% and modified PP/PE was blended by extrusion. The rheological properties were studied by using dynamic measurement. According to the strain sweep test, the linear viscoelastic region of composites in the melt was determined. The result showed that the storage modulus was independent of the strain at low strain region （〈0.1%）. The frequency sweep resuits indicated that all composites exhibited shear thinning behavior, and both the storage modulus and complex viscosity of MAH modified composites were decreased comparing to those unmodified. Flexural properties and impact strength of the prepared WPCs were measured according to the relevant standard specifications. The flexural and impact strength of the manufactured composites significantly increased and reached a maximum when MAH dosage was 1.0 wt%, whereas the flexural modulus after an initial decreased, also increased with MAH dosage. The increase in mechanical properties indicated that the presence of anhydride groups enhanced the interracial adhesion between wood fiber and PP/PE blends.     

Effects of tension wood on kraft paper from a short-rotation hardwood (Populus “Tristis No. 1”)

The physical properties and morphology of kraft paper handsheets obtained from tension wood of intensively managed, 5-year-old trees of Populus Tristis No. 1 were compared to those produced from isolated normal wood of the same stems. Pulp yields of tension wood (TW) and normal wood (NW) were 60 and 53% respectively. Over a beating range of 0–45 minutes, strength properties of TW paper were in all cases noticeably inferior to those obtained from NW. During paper formation, the TW or gelatinous fibers resisted collapse, even upon extended refining, and produced thick, porous sheets of poorly bonded elements. It was concluded that the differential behavior of NW and TW pulps was in several respects analogous to those displayed by earlywood and latewood pulps, respectively, of softwood species as well as thin-vs. thick-walled hardwood fibers. Consequently, it appears that the inferior strength of TW paper is primarily a function of fiber morphology, and the difference in hemicellulose content between NW and TW (viz., lower pentosan content of TW) often cited in the literature as a potential major factor here probably contributes little if any significant effect on ultimate interfiber bonding and paper quality.The experimental phase of this investigation was carried out by K. W. Robinson in partial fulfillment of the requirements of The Institute of Paper Chemistry for the M.S. degree from Lawrence University, Appleton, Wisconsin.     

Performance of three-layer composites with densified surface layers of Nothofagus pumilio and N. antarctica from Southern Patagonian forests

Nothofagus pumilio and N. antarctica forests of Southern Patagonia, Argentina are currently being managed for production of saw logs with fast growing conditions. The result of these management strategies is faster growing forests, but also an increase in the percentage of low-density wood. The motivation for this study was to find a way to valorise this low-density wood. Surface and bulk densification treatments were applied these wood species and were then applied as face layers in three-layer composites. The mechanical properties of three-layer composites were studied. The modulus of elasticity, modulus of rupture, and modulus of hardness were improved compared to many types of structural composite lumber. This increase in mechanical properties of N. pumilio and N. antarctica wood opens the possibility for its use in structural composites and added value to these otherwise underutilised and undervalued species.     

Physical and mechanical properties of wood after moisture conditioning

Some properties of wood (hinoki:Chamaecyparis obtusa) moisture-conditioned by an adsorption process from a dry state and by two desorption processes (from a water-saturated state and from a state with a moisture content slightly below the fiber saturation point) were investigated. The moisture contents of wood conditioned by the adsorption process and by the desorption process continued to approach to one another for the moisture-conditioning period of over 50 weeks. Accordingly, sorption hysteresis should be regarded as a transitional phenomenon that occurs during the process of approaching the true equilibrium, which requires a long time. The wood conditioned by the desorption process beginning from a water-saturated state showed slightly smaller dimensions than those conditioned by the adsorption process with the same moisture content; however, the wood conditioned by the desorption process from a moisture content below the fiber saturation point showed slightly larger dimensions than those conditioned by the adsorption process. The wood conditioned by the adsorption process from a dry state showed a higher modulus of elasticity and modulus of rupture than did the wood conditioned from a water-saturated state with the same moisture content. The mechanical properties of the wood also varied based on the states at which the desorption process was started. This is a notable characteristic of the relation between the drying condition and the mechanical properties of wood.     

Reaction wood anatomy and lignin distribution in <Emphasis Type="Italic">Gnetum gnemon</Emphasis> branches

This study investigated the anatomical and chemical characteristics of the reaction wood of a gymnpsperm species, Gnetum gnemon, and discussed on contributing factor for the type of reaction wood in this species. Cell morphology, microfibril angle (MFA) of the S₂ layer and lignin distribution in secondary walls of tracheary elements, and lignin content were examined on three branches. Observations included no G-layer formation, significant decreases in vessel frequency, and altered MFA, and visible-light absorbance after lignin colour reactions in tracheid and fiber tracheid walls on the upper side in almost all samples. These results suggest that reaction wood in G. gnemon was similar to that in ‘tension-wood-like-reaction wood’ in angiosperms. On the other hand, reaction wood showed decrease in the lignin concentration in the fiber tracheid walls compared to the tracheid walls. In addition, the lignin in the tracheid and fiber tracheid walls was originally rich in syringyl units, suggesting that changes in the anatomical and chemical characteristics of secondary xylem due to reaction wood formation might relate to the ratio of the syringyl to guaiacyl units in lignin in the cell walls which function for mechanical support.     

Effects of tension wood on specific conductivity and vulnerability to embolism of Quercus ilex seedlings grown at two atmospheric CO2 concentrations

To determine whether there are decreases in hydraulic function of a woody stem when it has increased mechanical loading, Quercus ilex L. seedlings were grown upright or inclined to force the production of large amounts of tension wood (TW). Seedlings were grown in ambient or elevated carbon dioxide concentrations ([CO2]) for 16-17 months to provide two sets of seedlings differing in growth rates and allocation patterns. In both CO2 environments, inclination caused formation of large amounts of TW at the base and mid-section of most stems, but not at the stem tips. Contrary to expectation, there were no significant effects of stem inclination or amount of TW on specific conductivity (k(s)) or vulnerability to embolism. Samples with high amounts of TW had higher vessel frequency, similar average vessel lumen area, similar vessel lumen fraction (6% of the transverse area), elevated frequency of vessels in the smallest diameter class, and higher wood density than samples with very little TW. Samples from seedlings in the elevated [CO2] treatment had similar vessel frequency, larger average vessel lumen area (caused by a higher frequency of large-diameter vessels), similar vessel lumen fraction, and similar wood density as samples from seedlings in the ambient [CO2] treatment. There was a strong position effect: the highest wood density and lowest ks were at the stem base, intermediate values were at the middle, and the lowest density and highest ks were at the stem tip. We conclude that, in a species that uses different cells for mechanical support and water transport, there can be large modifications in performance of the mechanical function through TW formation without impacting the water transport functions-ks and vulnerability to embolism.     

Fundamental studies on wood/cellulose-plastic composites: effects of composition and cellulose dimension on the properties of cellulose/PP composite

Although wood/cellulose-plastic composites (WPC) of low wood/cellulose content have been more accepted worldwide and are promoted as low-maintenance, high-durability building products, composites containing high wood/cellulose content are not yet developed on an industrial scale. In this study, flow properties, mechanical properties, and water absorption properties of the compounds of cellulose microfiber/polypropylene (PP) and maleic anhydride-grafted polypropylene (MAPP) were investigated to understand effects of the high cellulose content and the dimensions of the cellulose microfiber. The molding processes studied included compression, injection, and extrusion. It was found that fluidity is not only dependent on resin content but also on the dimension of the filler; fluidity of the compound declined with increased fiber length with the same resin content. Dispersion of the composite was monitored by charge-coupled device (CCD) microscope. Increasing the plastic content in the cellulose-plastic formulation improved the strength of mold in addition to the bond development between resin and filler, and the tangle of fibers. The processing mode affected the physicomechanical properties of the cellulosic plastic. Compression-molded samples exhibited the lowest modulus of rupture (MOR) and modulus of elasticity (MOE) and the highest water absorption, while samples that were injection-molded exhibited the highest MOR (70 MPa) and MOE (7 GPa) and low water absorption (2%).