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 and peroxidase in tension wood ofEucalyptus viminalis seedlings

Seedlings ofEucalyptus viminalis were grown for 50 days with their stems bent so tension wood would form. Every 10 days the lignin content, monomeric composition, and peroxidase activity in the tension wood were compared with those in the lower side (opposite wood) and in vertically grown controls. The lignin content in the developing tension wood started to decrease after 10 days of bending and kept decreasing for 50 days, whereas those in control plants and opposite wood remained almost unchanged. The yields of syringaldehyde from tension wood by nitrobenzene oxidation increased, and consequently the syringyl/ guaiacyl ratio of the lignin was higher in tension wood than in opposite wood and control plants. The peroxidase ionically bound to the cell walls (IPO) catalyzed oxidation of guaiacol and syringaldazine. The syringaldazineoxidizing activity of IPO from tension wood increased, whereas the activities of IPO from opposite wood and control plants did not show any marked change. In tension wood the increase in syringaldazine-oxidizing activity of IPO was consistent with an increase in the syringaldehyde yield. This suggests that IPO contributes to syringyl lignin deposition as other enzymes involved in the monolignol biosynthesis do in tension wood formation.This study was presented at the 50th Annual Meeting of the Japan Wood Research Society, Kyoto, April 2000     

Relation between developmental changes on anatomical structure and on protein pattern in differentiating xylem of tension wood

Tension wood was induced inEucalyptus camaldulensis L. by fixing the stem at an angle. Proteins in the differentiating tissue of tension wood were compared to those of normal wood on sodium dodecyl sulfate polyacrylamide gel electrophoresis. An obvious difference was found in the salt-soluble fraction of 14 days after inclination. At least five bands (19,22,37,41, and 55kDa) were specific in the differentiating tissue of tension wood. These proteins were bound to the cell wall, plasma membrane, or both by their electric charge; they were undetectable until 14 days after inclination. Mature tension wood was observed in the tissue at 14 days. Thus, all differentiating tissue at 14 days was produced after inclination. On the other hand, the differentiating zone at 7 days contained the same tissue, as in tension wood estimated by the vessel number and diameter during the early phase; and the tissue was indistinguishable from normal wood during the late phase. The proteins found here were related to the phenomenon occurring in the late stage of xylem differentiation.     

Stem tangential strain on the tension wood side of <Emphasis Type="Italic">Fagus crenata</Emphasis> saplings

The tangential strain on the inner bark surface of Fagus crenata sapling stems was continuously measured using strain gauges. The total strain increased daily, increasing at night and decreasing during the day. When tension wood was induced by artificial inclination, the strain increased more on the upper side than on the lower side; and the increment in the strain at night was larger on the upper side than on the lower. The change in tangential strain on the inner bark surface arose from changes in the water content and the volume of differentiating cells. Differentiating tension wood fibers appear to contain more water and to expand more at night than differentiating normal wood fibers. We can determine whether tension wood is formed from the tangential strain during growth.     

Role of the gelatinous layer on the origin of the physical properties of the tension wood

To discuss the role of the gelatinous layer (G-layer) on the origins of the physical properties peculiar to the tension wood fiber (TW fiber), the deformation process of an isolated TW fiber caused by a certain biomechanical state change was formulated mathematically. The mechanical model used in the present formulation is a four-layered hollow cylinder having the compound middle lamella (CML), the outer layer of the secondary wall (S1) and its middle layer (S2), and the G-layer (G) as an innermost layer. In the formulation, the reinforced matrix mechanism was applied to represent the mechanical interaction between the cellulose microfibril (CMF) as a framework bundle and the amorphous substance as a matrix skeleton in each layer. The model formulated in the present study is thought to be useful to investigate the origins of extensive longitudinal drying shrinkage, large tensile growth stress, and a high axial elastic modulus, which are rheological properties peculiar to the TW. In this article, the detailed process of the mathematical formulation is described. In a subsequent article, some TW properties from a 70-year-old Kohauchiwakaede (Acer sieboldianum Miq.) will be analyzed using the newly developed model.     

The influences of boiling and drying treatments on the behaviors of tension wood with gelatinous layers in <Emphasis Type="Italic">Zelkova serrata</Emphasis>

This study examined how boiling and drying treatments influenced various physical properties of the tension wood with gelatinous fibers (G-fibers) of a 29-yearold Zelkova branch. By boiling treatment, tension wood with numerous G-fibers contracted considerably in the longitudinal direction and the longitudinal Young’s modulus decreased in spite of the water-saturated condition. The drying treatment caused green tension wood and boiled tension wood with numerous G-fibers to shrink longitudinally and increased their longitudinal Young’s moduli. These specific behaviors in tension wood were highly correlated with the proportion of G-fibers in a specimen and were probably caused by the microscopic behavior of cellulose microfibril (CMF) in the gelatinous layers (G-layers). The longitudinal shrinkage of tension wood due to drying suggests the existence of a hygro-sensible, noncrystalline region in the CMF, which is abundant in the G-layer. Furthermore, the noncrystalline region in the CMF softens during boiling treatment, resulting in the reduction of the longitudinal Young’s modulus in tension wood. The longitudinal contraction of tension wood with G-fibers by boiling might be caused by the tensile growth stress remaining in green G-layers. However, no changes were detected in the 004 d-spacing of cellulose crystal in tension wood from the boiling and drying treatments, regardless of the proportion of G-fibers.     

Evaluation of field performance of poplar clones using selected competition indices

Use of competition indices in the analysis of forestry experiments may improve detection and understanding of treatment effects, and thereby improve the application of results. In this paper, we compared the performance of 8 indices in an analysis of a spacing trial of four Populus clones planted in pure and mixed clonal plots. Indices were included as covariates in analyses of variance and evaluated on their ability to decrease mean square error. Indices that were simple to calculate (i.e., required only diameter and spacing or distance information) decreased mean square error by as much as 32%. We then illustrate the use of a simple index to assess clonal response to intra- and inter-genotypic competition and to interpret treatment effects confounded by different levels of competition. In pure clonal plots (intra-genotypic competition), all the clones tested reacted similarly to competition, while the same clones tested in mixed plots (inter-genotypic competition) reacted differently to varying levels of competition. The use of a competition index to assess clone rankings within a mixed clonal plot can be an effective way to predict clonal performance within a mono-clonal planting.     

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.     

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。     

Investigation on transversal variation of poplar tension wood quality using ultrasound wave parameters

Transversal variations of some ultrasound wave parameters (phase velocity, group velocity, attenuation coefficient and acoustic radiation) were measured from the pith to the bark of poplar tension wood discs in different orthotropic directions. Over 80 cubic specimens of 2 × 2 × 10 cm³ (radial, tangential and longitudinal, respectively) were prepared and tested acoustically. Samples containing tension woods were separated from normal ones using anatomical experiments. Results showed acoustical behaviour of normal and tension wood improved in longitudinal direction while the distance between pith and bark increased; also, wave parameter variations were less important in radial and tangential directions. In addition, phase velocity and acoustic radiation – which significantly varied – were the best parameters for quality assessment of poplar wood compared with group velocity and attenuation coefficients. Since samples near the bark were acoustically better than counterparts near the pith, they could be used in products requiring more strength, like lumber. Finally, acoustic radiation and phase velocity were correlated more strongly with density than group velocity and attenuation.     

Effect of cutting position on rooting and shoot growth of two poplar clones

One-year-old shoots from stooling beds of Populus × deltoides Walker and Populus jackii Northwest were collected in late fall to determine the effect of cutting position on rooting and growth of hardwood poplar cuttings. Cuttings with a minimum diameter of 9.0 mm were obtained from basal, middle and distal locations on the poplar shoot. Rooting and shoot growth were assessed in the greenhouse and under field conditions. Walker poplar cuttings collected from basal portions of the shoot and planted in the field had 87.6% rooting and 103.8 cm growth compared to 78.8% and 103.4 cm and 67.6% and 88.8 cm for middle and distal locations respectively. For Northwest rooting and growth under field conditions was 84.4% and 94.7 cm for basal, 78.4% and 90.5 cm for mid and 64.4% and 84.3 cm for distal locations. Rooting in the greenhouse was superior to the field. Walker cuttings had fewer roots and buds per cutting than Northwest, however, growth of Walker in the field was superior.     

The effects of flooding on several hybrid poplar clones in Northern China

The ecophysiological, morphological, and growth characteristics of 14 poplar clones were studied during 37 days of flooding and a 13-day recovery period. Cuttings were subjected to three soil water regimes, viz. drained (control), shallow flooding to 10 cm above the soil, and deep flooding to a depth of 120 cm. All hybrids modified their ecophysiological and morphological patterns to decrease carbon loss and maintain water balance. In response to flooding, all 14 hybrids reduced their expansion and initiation of new leaves, reduced height and root collar growth, and reduced the number of leaves. For shallowly flooded plants, adventitious roots developed by day 14, and their number increased with flooding duration; net photosynthesis, stomatal conductance, and growth decreased significantly compared with the control; dry weights of roots, leaves, and total biomass decreased and the allocation of growth to shoots and roots changed. After flooding ended, net photosynthesis recovered, but stomatal conductance recovered before net CO₂ assimilation since photosynthesis was limited by stomatal factor at the initial stage of stress and it was limited by non-stomatal factors over relatively long periods of stress. Transpiration and the amount of water obtained from the roots both decreased. In the deeply flooded plants, similar but often more severe changes were observed. Based on our results, we classified the hybrids into three types using hierarchical cluster analysis. Clones 15-29, 196-522, 184-411, 306-45, 59-289, DN-2, DN-182, DN-17, DN-14274, NE-222, DTAC-7, and R-270 were flood-tolerant, clone NM-6 was flood-susceptible, and clone 328-162 was moderately flood-tolerant.     

Effect of thinning on the development of compression wood in stems of Corsican pine

This study considered the effects of thinning on the development of compression wood in stems of 35-year-old stand of Corsican pine (Pinus nigra L.). Part of the stand had been thinned at 5-yearly intervals and part left unthinned. Twenty trees each from the thinned and unthinned stands were randomly selected and felled. Measurements were made on tree height, stem diameter, stem slenderness and canopy depth. Wood samples were removed from the central part of the main log and cross-sectional measurements made on ring width, basic density and compression wood content. Cross-sectional area of compression wood was found to be three time higher in stems from the unthinned trees in comparison with those from the thinned trees. No significant differences in mean radial ring width or basic density were found between treatments. Correlations indicated that, with increasing in stem diameter, compression wood content increased in the unthinned trees, while a decline in compression was observed in the thinned trees. Tree height was also positively correlated with compression wood content in unthinned trees, while no equivalent relationship was observed in thinned trees. Observations from this study, while not conclusive, suggest that phototropic stimulus may be producing stem inclinations in the unthinned stand as trees compete for space in the canopy, whereas crown competition has been largely eliminated in the thinned stand; and that this is responsible for compression wood levels recorded in this study.     

Biomass production by two-year-old poplar clones on floodplain sites in the Lower Midwest, USA

Four Populus clones were grown for two years at 1×1 m spacing for study of total biomass production and carbon sequestration capacity on floodplain sites previously in forage grasses under climatic conditions of the lower Midwest, U.S.A. Total biomass (above-and below-ground) in the first year ranged from 3.9 Mg ha^–1 in a Populus deltoides x P. nigra clone (I45/51) to 1.9 Mg ha^–1 for a local-source Populus deltoides clone (2059). Second year total biomass production was substantially higher, ranging from 13.9 Mg ha^–1 in I45/51 to 7.4 Mg ha^–1 in P. deltoides clone 26C6R51. Second-year leaf area index (LAI) values for I45/51 plants reached 4 during mid-season, indicating essentially complete canopy closure in this clone by the second year after planting. In contrast, maximum mid-season, second-year LAI was significantly lower in P. deltoides clones ( 2.4). There was some evidence for differential allocation to roots and shoots among Populus clones, with 26C6R51 showing relatively more allocation to root biomass than other clones. Second-year growth in Populus deltoides clone 2059 accelerated substantially, and this genotype exhibited two-year biomass accumulation nearly equal to that of I45/51 despite having less leaf area. This result suggested a higher photosynthetic capacity or assimilation efficiency in the former. This revised version was published online in August 2006 with corrections to the Cover Date.     

Fallen wood decomposition ofPinus koraiensis andTilia amurensis

Fallen wood decomposition rate ofPinus koraiensis andTilia amurensis in broadleaved Korean pine forest was studied in this paper. The result showed that decomposition rate of fallen wood was different from that of little diameter wood and coarse woody debris for the same tree species. Fallen wood decomposition was generally rotten from outside to inside. And decomposition speed of fallen woods was different according to tree species and site, and it was also related to diameter of fallen woods. Decomposition depth ofTilia amurensis fallen wood for 17 years was 14 cm, but that ofPinus koraiensis in the same condition was less than 7 cm.Tilia amurensis was decomposed faster thanPinus koraiensis. For same tree species, if the diameter was small, the decomposition speed was quick. This project was supported by Natural Science Foundation of China (No. 39670144) and funded by the Opened Research Station of Changhai Mountain Forest Ecosystem, Chinese Academy of Sciences. (Responsible editor: Zhu Hong)     

Factors of affecting the spring back of compressedPaulownia wood

In order to increase its hardness and gravity as well as dimension stability, the technology of hot-compressing onPaulownia wood was studied. The main factors of affecting the spring back of the compressedPaulownia samples were discussed. It was discovered that every factor in the experiment had obvious effects on wood hardness and dimension stability of compressed wood. When the MC (Moisture Content) of experimental specimens was 13.89%, it was useful to spray water on the surface of samples before hot pressing. The best result was the recovery of compression set could decrease from 90.69% of untreated wood to 45.51% of soaking specimens into PF (Phenol Formaldehyde) water solution. The hot pressing time was 8 min at 190°C. Responsible editor: Zhu Hong     

Pathogenicity of the pine wood nematode at different developmental stages

The ability of the pine wood nematode,Bursaphelenchus xylophilus, a pathogen that causes pine wilt disease, to kill cortical cells of Japanese black pine,Pinus thunbergii, during early development of the disease was conjectured to be a function of nematode developmental stage. A tangential segment of bark was removed from a 2-cm-long current-year stem. The cortex-exposed segments with cut cortical resin canals were designated as + RC-segments and those without them as − RC-segments. When a nematode population containing many older juveniles and adults (NL) was inoculated onto the cut surface, the − RC-segments were still alive 4 d after inoculation, as were non-inoculated control segments. When cortex-exposed segments were inoculated with either a nematode population containing many younger nematodes (NS) or with nematodes isolated from inoculated pine cuttings that also contained many younger juveniles, most tissue cells in − RC-segments died 4 d after inoculation, suggesting that younger juveniles killed pine cells directly, in contrast with older juveniles and adults. When nematodes were inoculated onto + RC-segments in which they could easily enter resin canals, both NL and NS killed the segment tissues. This suggests that NL is pathogenic to pine cells while living in resin canals. Such differences in the pathogenicity of NL and NS to pine parenchymatous cells were also demonstrated in a pathogenicity assay system using bark peelings, which allowed an estimate of direct attack on the cambial cells by nematodes. Based on these results, we hypothesize that younger juveniles are pathogenic to pine parenchymatous cells, while adults and older juveniles are not pathogenic. This work was supported in part by Grants-in-Aid for Scientific Research (No.01440012 and 06454088) and for Young Scientists (to K.I.) from Ministry of Education, Science, Culture, and Sports of Japan, and by a grant from PROBRAIN.     

Effect of thinning and fertilizer on the cellulose crystallite width of Eucalyptus globulus

An analysis of cellulose crystallite width, microfibril angle and wood density after the time of thinning (at 8 years) in straight vertical trees was undertaken in a 13-year-old E. globulus trial designed to assess the effect of thinning on tension wood formation. The most important effect was on cellulose crystallite width, which increased with thinning intensity and this was mitigated where fertilizer was applied at the time of thinning. Given the relationship between high crystallite width and tension wood occurrence the results demonstrate that heavy thinning of E. globulus at this age can contribute to tension wood formation. However, tension wood production may be significantly reduced where fertilizer is applied. This is possibly because increased diameter growth as a response to fertilizer application stabilises the stems and this is the mechanism by which trees cope with internal stresses that are generated from wind in destabilsed stands following thinning. In contrast, trees that respond poorly to thinning produce tension wood.     

Effects of radial growth rate on selected indices for juvenile and mature wood of the Japanese larch

To examine the differences between juvenile and mature wood, 12 aged sample trees from two areas of Nagano Prefecture were harvested; and the radial development of tracheid length, the ring density, and the relation of the radial growth rate (observed by ring width) with some selected indices of ring structure were investigated. The results proved that the radial variation of tracheid length with ring number can be described by a logarithmic formula, and both plantations reached the demarcation of juvenile and mature wood at age 18. With the segmented regression method, we also analyzed radial variation of mean density and found that the demarcation of juvenile and mature wood was at age 15 for sample trees from Saku and at age 21 for those from Yabuhara. By using the results of estimates from juvenile and mature wood based on ring density, we found that high growth rates resulted when producing lower-density wood during the juvenile period, but these rates did not occur during the mature period. The basic reason for this phenomenon is the variation in patterns of earlywood and latewood in juvenile and mature wood, respectively. This result advised us that when managing plantations of Japanese larch it is necessary to take different measurements at different growth periods.