Localised laccase activity modulates distribution of lignin polymers in gymnosperm compression wood.

The woody stems of coniferous gymnosperms produce specialised compression wood to adjust the stem growth orientation in response to gravitropic stimulation. During this process, tracheids develop a compression-wood-specific S2 L cell wall layer with lignins highly enriched with p-hydroxyphenyl (H)-type units derived from H-type monolignol, whereas lignins produced in the cell walls of normal wood tracheids are exclusively composed of guaiacyl (G)-type units from G-type monolignol with a trace amount of H-type units. We show that laccases, a class of lignin polymerisation enzymes, play a crucial role in the spatially organised polymerisation of H-type and G-type monolignols during compression wood formation in Japanese cypress (Chamaecyparis obtusa). We performed a series of chemical-probe-aided imaging analysis on C. obtusa compression wood cell walls, together with gene expression, protein localisation and enzymatic assays of C. obtusa laccases. Our data indicated that CoLac1 and CoLac3 with differential oxidation activities towards H-type and G-type monolignols were precisely localised to distinct cell wall layers in which H-type and G-type lignin units were preferentially produced during the development of compression wood tracheids. We propose that, not only the spatial localisation of laccases, but also their biochemical characteristics dictate the spatial patterning of lignin polymerisation in gymnosperm compression wood.

Structural organization of the cell wall polymers in compression wood as revealed by FTIR microspectroscopy.

MAIN CONCLUSION: Glucomannan was more strongly oriented, in line with the orientation of cellulose, than the xylan in both compression wood and normal wood of Chinese fir. Lignin in compression wood was somewhat more oriented in the direction of the cellulose microfibrils than in normal wood. The structural organization in compression wood (CW) is quite different from that in normal wood (NW). To shed more light on the structural organization of the polymers in plant cell walls, Fourier Transform Infrared (FTIR) microscopy in transmission mode has been used to compare the S2-dominated mean orientation of wood polymers in CW with that in NW from Chinese fir (Cunninghamia lanceolata). Polarized FTIR measurements revealed that in both CW and NW samples, glucomannan and xylan showed a parallel orientation with respect to the cellulose microfibrils. In both wood samples, the glucomannan showed a much greater degree of orientation than the xylan, indicating that the glucomannan has established a stronger interaction with cellulose than xylan. For the lignin, the absorption peak also indicated an orientation along the direction of the cellulose microfibrils, but this orientation was more pronounced in CW than in NW, indicating that the lignin is affected by the orientation of the cellulose microfibrils more strongly in CW than it is in NW.

Tracheid cell-wall structures and locations of (1 → 4)-ß-D-galactans and (1 → 3)-ß-D-glucans in compression woods of radiata pine (Pinus radiata D. Don).

BACKGROUND: Compression wood (CW) forms on the underside of tilted stems of coniferous gymnosperms and opposite wood (OW) on the upperside. The tracheid walls of these wood types differ structurally and chemically. Although much is known about the most severe form of CW, severe CW (SCW), mild CWs (MCWs), also occur, but less is known about them. In this study, tracheid wall structures and compositions of two grades of MCWs (1 and 2) and SCW were investigated and compared with OW in slightly tilted radiata pine (Pinus radiata) stems. RESULTS: The four wood types were identified by the distribution of lignin in their tracheid walls. Only the tracheid walls of OW and MCW1 had a S3 layer and this was thin in MCW1. The tracheid walls of only SCW had a S2 layer with helical cavities in the inner region (S2i). Using immunomicroscopy, (1 → 4)-ß-D-galactans and (1 → 3)-ß-D-glucans were detected in the tracheid walls of all CWs, but in only trace amounts in OW. The (1 → 4)-ß-D-galactans were located in the outer region of the S2 layer, whereas the (1 → 3)-ß-D-glucans were in the inner S2i region. The areas and intensities of labelling increased with CW severity. The antibody for (1 → 4)-ß-D-galactans was also used to identify the locations and relative amounts of these galactans in whole stem cross sections based on the formation of an insoluble dye. Areas containing the four wood types were clearly differentiated depending on colour intensity. The neutral monosaccharide compositions of the non-cellulosic polysaccharides of these wood types were determined on small, well defined discs, and showed the proportion of galactose was higher for CWs and increased with severity. CONCLUSION: The presence of an S3 wall layer is a marker for very MCW and the presence of helical cavities in the S2 wall layer for SCW. The occurrence and proportions of (1 → 4)-ß-D-galactans and (1 → 3)-ß-D-glucans can be used as markers for CW and its severity. The proportions of galactose were consistent with the labelling results for (1 → 4)-ß-D-galactans.

Gravitropisms and reaction woods of forest trees - evolution, functions and mechanisms.

Contents 790 I. 790 II. 792 III. 795 IV. 797 V. 798 VI. 800 VII. 800 800 References 800 SUMMARY: The woody stems of trees perceive gravity to determine their orientation, and can produce reaction woods to reinforce or change their position. Together, graviperception and reaction woods play fundamental roles in tree architecture, posture control, and reorientation of stems displaced by wind or other environmental forces. Angiosperms and gymnosperms have evolved strikingly different types of reaction wood. Tension wood of angiosperms creates strong tensile force to pull stems upward, while compression wood of gymnosperms creates compressive force to push stems upward. In this review, the general features and evolution of tension wood and compression wood are presented, along with descriptions of how gravitropisms and reaction woods contribute to the survival and morphology of trees. An overview is presented of the molecular and genetic mechanisms underlying graviperception, initial graviresponse and the regulation of tension wood development in the model angiosperm, Populus. Critical research questions and new approaches are discussed.

Cavitation of intercellular spaces is critical to establishment of hydraulic properties of compression wood of Chamaecyparis obtusa seedlings.

BACKGROUND AND AIMS: When the orientation of the stems of conifers departs from the vertical as a result of environmental influences, conifers form compression wood that results in restoration of verticality. It is well known that intercellular spaces are formed between tracheids in compression wood, but the function of these spaces remains to be clarified. In the present study, we evaluated the impact of these spaces in artificially induced compression wood in Chamaecyparis obtusa seedlings. METHODS: We monitored the presence or absence of liquid in the intercellular spaces of differentiating xylem by cryo-scanning electron microscopy. In addition, we analysed the relationship between intercellular spaces and the hydraulic properties of the compression wood. KEY RESULTS: Initially, we detected small intercellular spaces with liquid in regions in which the profiles of tracheids were not rounded in transverse surfaces, indicating that the intercellular spaces had originally contained no gases. In the regions where tracheids had formed secondary walls, we found that some intercellular spaces had lost their liquid. Cavitation of intercellular spaces would affect hydraulic conductivity as a consequence of the induction of cavitation in neighbouring tracheids. CONCLUSIONS: Our observations suggest that cavitation of intercellular spaces is the critical event that affects not only the functions of intercellular spaces but also the hydraulic properties of compression wood.

Fluorescence-Detected Linear Dichroism of Wood Cell Walls in Juvenile Serbian Spruce: Estimation of Compression Wood Severity.

Fluorescence-detected linear dichroism (FDLD) microscopy provides observation of structural order in a microscopic sample and its expression in numerical terms, enabling both quantitative and qualitative comparison among different samples. We applied FDLD microscopy to compare the distribution and alignment of cellulose fibrils in cell walls of compression wood (CW) and normal wood (NW) on stem cross-sections of juvenile Picea omorika trees. Our data indicate a decrease in cellulose fibril order in CW compared with NW. Radial and tangential walls differ considerably in both NW and CW. In radial walls, cellulose fibril order shows a gradual decrease from NW to severe CW, in line with the increase in CW severity. This indicates that FDLD analysis of cellulose fibril order in radial cell walls is a valuable method for estimation of CW severity.

Xylem parenchyma cell walls lack a gravitropic response in conifer compression wood.

MAIN CONCLUSION: Cell wall fluorescence and immunocytochemistry demonstrate that xylem parenchyma cell walls do not show changes in structure and composition related to gravitropic response comparable to those of tracheids, even when they have lignified secondary cell walls. Tracheid cell walls in compression wood have altered composition and structure which generates the strain responsible for correction of stem lean as part of the gravitropic response of woody plants. Xylem parenchyma cell walls vary among conifer species and can be lignified secondary walls (spruce) or unlignified primary walls (pine). It can be expected that xylem parenchyma with lignified secondary cell walls might show features of compression wood comparable to those of tracheids that have a similar type of cell wall. A comparison of xylem parenchyma cell walls in normal and compression wood in species with lignified and non-lignified parenchyma cell walls provides a unique opportunity to understand the process of reaction wood formation in conifers. Using both UV/visible fluorescence microscopy of cell wall fluorophores and immunocytochemistry of galactan and mannan epitopes, we demonstrate that xylem parenchyma cell walls do not show the changes in composition and structure typical of compression wood tracheids. Adjacent cells of different types but with similar cell wall structure can undergo cell wall developmental changes related to support or defence functions independent of their neighbours. Tracheids are sensitive to gravitropic signals while xylem parenchyma cells are not.

Cellulose structure and lignin distribution in normal and compression wood of the Maidenhair tree (Ginkgo biloba L.).

We studied in detail the mean microfibril angle and the width of cellulose crystals from the pith to the bark of a 15-year-old Maidenhair tree (Ginkgo biloba L.). The orientation of cellulose microfibrils with respect to the cell axis and the width and length of cellulose crystallites were determined using X-ray diffraction. Raman microscopy was used to compare the lignin distribution in the cell wall of normal/opposite and compression wood, which was found near the pith. Ginkgo biloba showed a relatively large mean microfibril angle, varying between 19° and 39° in the S2 layer, and the average width of cellulose crystallites was 3.1-3.2 nm. Mild compression wood without any intercellular spaces or helical cavities was observed near the pith. Slit-like bordered pit openings and a heavily lignified S2L layer confirmed the presence of compression wood. Ginkgo biloba showed typical features present in the juvenile wood of conifers. The microfibril angle remained large over the 14 annual rings. The entire stem disc, with a diameter of 18 cm, was considered to consist of juvenile wood. The properties of juvenile and compression wood as well as the cellulose orientation and crystalline width indicate that the wood formation of G. biloba is similar to that of modern conifers.

Modelling of the hygroelastic behaviour of normal and compression wood tracheids.

Compression wood conifer tracheids show different swelling and stiffness properties than those of usual normal wood, which has a practical function in the living plant: when a conifer shoot is moved from its vertical position, compression wood is formed in the under part of the shoot. The growth rate of the compression wood is faster than in the upper part resulting in a renewed horizontal growth. The actuating and load-carrying function of the compression wood is addressed, on the basis of its special ultrastructure and shape of the tracheids. As a first step, a quantitative model is developed to predict the difference of moisture-induced expansion and axial stiffness between normal wood and compression wood. The model is based on a state space approach using concentric cylinders with anisotropic helical structure for each cell-wall layer, whose hygroelastic properties are in turn determined by a self-consistent concentric cylinder assemblage of the constituent wood polymers. The predicted properties compare well with experimental results found in the literature. Significant differences in both stiffness and hygroexpansion are found for normal and compression wood, primarily due to the large difference in microfibril angle and lignin content. On the basis of these numerical results, some functional arguments for the reason of high microfibril angle, high lignin content and cylindrical structure of compression wood tracheids are supported.

Structural Characteristics of Reaction Tissue in Plants.

To maintain or adjust posture under the challenges of gravity and increased self-weight, or the effects of light, snow, and slope, plants have the ability to develop a special type of tissue called reaction tissue. The formation of reaction tissue is a result of plant evolution and adaptation. The identification and study of plant reaction tissue are of great significance for understanding the systematics and evolution of plants, the processing and utilization of plant-based materials, and the exploration of new biomimetic materials and biological templates. Trees' reaction tissues have been studied for many years, and recently, many new findings regarding these tissues have been reported. However, reaction tissue requires further detailed exploration, particularly due to their complex and diverse nature. Moreover, the reaction tissues in gymnosperms, vines, herbs, etc., which display unique biomechanical behavior, have also garnered the attention of research. After summarizing the existing literature, this paper provides an outline of the reaction tissues in woody plants and non-woody plants, and lays emphasis on alternations in the cell wall structure of the xylem in softwood and hardwood. The purpose of this paper is to provide a reference for the further exploration and study of reaction tissues with great diversity.

Seasonal Developing Xylem Transcriptome Analysis of Pinus densiflora Unveils Novel Insights for Compression Wood Formation.

Wood is the most important renewable resource not only for numerous practical utilizations but also for mitigating the global climate crisis by sequestering atmospheric carbon dioxide. The compressed wood (CW) of gymnosperms, such as conifers, plays a pivotal role in determining the structure of the tree through the reorientation of stems displaced by environmental forces and is characterized by a high content of lignin. Despite extensive studies on many genes involved in wood formation, the molecular mechanisms underlying seasonal and, particularly, CW formation remain unclear. This study examined the seasonal dynamics of two wood tissue types in Pinus densiflora: CW and opposite wood (OW). RNA sequencing of developing xylem for two consecutive years revealed comprehensive transcriptome changes and unique differences in CW and OW across seasons. During growth periods, such as spring and summer, we identified 2255 transcripts with differential expression in CW, with an upregulation in lignin biosynthesis genes and significant downregulation in stress response genes. Notably, among the laccases critical for monolignol polymerization, PdeLAC17 was found to be specifically expressed in CW, suggesting its vital role in CW formation. PdeERF4, an ERF transcription factor preferentially expressed in CW, seems to regulate PdeLAC17 activity. This research provides an initial insight into the transcriptional regulation of seasonal CW development in P. densiflora, forming a foundation for future studies to enhance our comprehension of wood formation in gymnosperms.

Differential Expression of Arabinogalactan in Response to Inclination in Stem of Pinus radiata Seedlings.

Arabinogalactan proteins (AGPs) are members of a family of proteins that play important roles in cell wall dynamics. AGPs from inclined pines were determined using JIM7, LM2, and LM6 antibodies, showing a higher concentration in one side of the stem. The accumulation of AGPs in xylem and cell wall tissues is enhanced in response to loss of tree stem verticality. The differential gene expression of AGPs indicates that these proteins could be involved in the early response to inclination and also trigger signals such as lignin accumulation, as well as thicken cell wall and lamella media to restore stem vertical growth. A subfamily member of AGPs, which is Fasciclin-like has been described in angiosperm species as inducing tension wood and in some gymnosperms. A search for gene sequences of this subfamily was performed on an RNA-seq library, where 12 sequences were identified containing one or two fasciclin I domains (FAS), named PrFLA1 to PrFLA12. Four of these sequences were phylogenetically classified in group A, where PrFLA1 and PrFLA4 are differentially expressed in tilted pine trees.

Mechanics of a Biomimetic Moisture Sensitive Actuator Based on Compression Wood.

Various mechanisms of plant organ movements have been reported, including the close association of two layers with expressed differences in hygroscopic properties. Following this principle, actuator beams composed of thin veneers out of normal and compression wood cut from Scots pine (Pinus sylvestris L.) were prepared by using two types of adhesives. The mismatch of the swelling properties of the two layers in tight combination resulted in an expressed bending deflection in response to set humidity changes. The resulting curvatures were measured and analyzed by the Timoshenko bi-metal-model, as well as with an enhanced three-layer model, with the latter also considering the mechanical influence of the glueline on the actuator bending. The thermally induced strain in the original model was replaced by another strain due to moisture changes. The strain was modelled as a function of wood density, along with changes in wood moisture. Experiments with free movement of the bilayer to measure curvature, and with constraints to determine forces, were performed as well. Deformation and magnitude of actuators movements were in close agreement with the enhanced bilayer-model for the phenol-resorcinol-formaldehyde adhesive, which deviated substantially from the casein adhesive glued actuators. The obtained results are seen as critical for wood-based actuator systems that are potentially used in buildings or other applications.

Comprehensive Transcriptome Analysis of Stem-Differentiating Xylem Upon Compression Stress in Cunninghamia Lanceolata.

Compression wood (CW) in gymnosperm brings great difficulties to wood industry using wood as raw materials since CW presents special wood structure and have different physical and chemical properties from those of normal wood (NW). Chinese fir (Cunninghamia lanceolata) is widely distributed in China. However, global transcriptome profiling of coding and long non-coding RNA in response to compression stress has not been reported in the gymnosperm species. In this study, we revealed that CW in Chinese fir exhibited distinct morphology and cytology properties compared with those of NW, including high lignin content, thick and round tracheid cells. Furthermore, we combined both PacBio long-read SMRT sequencing (Iso-Seq) and Illumina short-read RNA-Seq to reveal the transcriptome in stem-differentiating xylem (SDX) under different time points (2, 26, and 74 h) upon compression stress in NW, CW, and OW (opposite wood), respectively. Iso-Seq was successfully assembled into 41,253 de-novo full-length transcriptome reference (average length 2,245 bp). Moreover, there were striking differences in expression upon compression stress, which were involved 13 and 7 key enzyme genes in the lignin and cellulose synthesis, respectively. Especially, we revealed 11 secondary growth-related transcription factors show differential expression under compression stress, which was further validated by qRT-PCR. Finally, the correlation between 6,533 differentially expressed coding genes and 372 differentially expressed long non-coding RNAs (lncRNAs) indicates that these lncRNAs may affect cell wall biogenesis and xyloglucan metabolism. In conclusion, our results provided comprehensive cytology properties and full-length transcriptome profiling of wood species upon compression stress. Especially we explored candidate genes, including both coding and long non-coding genes, and provided a theoretical basis for further research on the formation mechanism of CW in gymnosperm Chinese fir.

Is the Responsiveness to Light Related to the Differences in Stem Straightness among Populations of Pinus pinaster?

Stem straightness is related to wood quality and yield. Although important genetic differences in stem straightness among the natural populations of Pinus pinaster are well established, the main drivers of these differences are not well known. Since the responses of trees to light are key ecological features that induce stem curvature, we hypothesized that populations with better straightness should exhibit lower photomorphogenetic and phototropic sensitivity. We compared three populations to identify the main processes driven by primary and secondary growth that explain their differences in response to light. One-year-old seedlings were grown under two treatments-direct sunlight and lateral light plus shade-for a period of 5 months. The length and the leaning of the stems were measured weekly. The asymmetry of radial growth and compression wood (CW) formation were analyzed in cross-sections. We found differences among the populations in photomorphogenetic and phototropic reactions. However, the population with straighter stems was not characterized by reduced sensitivity to light. Photo(gravi)tropic responses driven by primary growth and gravitropic responses driven by secondary growth explained the kinetics of the stem leaning and CW pattern. Asymmetric radial growth and CW formation did not contribute to the phototropic reactions.

Dimensional Changes of Tracheids during Drying of Radiata Pine (Pinus radiata D. Don) Compression Woods: A Study Using Variable-Pressure Scanning Electron Microscopy (VP-SEM).

Variable-pressure scanning electron microscopy was used to investigate the dimensional changes in longitudinal, tangential and radial directions, on wetting and drying, of tracheids of opposite wood (OW) and three grades of compression woods (CWs), including severe CW (SCW) and two grades of mild compression wood (MCW) (MCW1 and MCW2) in corewood of radiata pine (Pinus radiata) saplings. The CW was formed on the underside and OW on the upper side of slightly tilted stems. In the longitudinal direction, the shrinkage of SCW tracheids was ~300% greater than that of OW tracheids, with the shrinkage of the MCW1 and MCW2 tracheids being intermediate. Longitudinal swelling was also investigated and hysteresis was demonstrated for the tracheids of all corewood types, with the extent of hysteresis increasing with CW severity. A statistical association was found between longitudinal shrinkage and the content of lignin and galactosyl residues in the cell-wall matrix. The galactosyl residues are present mostly as (1→4)-ß-galactans, which are known to have a high capacity for binding water and swell on hydration. The small proportions of (1→3)-ß-glucans in the CWs have similar properties. These polysaccharides may play a functional role in the longitudinal shrinking and swelling of CW tracheids. Tangential shrinkage of tracheids was greater than radial shrinkage but both were greatest for OW and least for SCW, with the MCW1 and MCW2 being intermediate.

Using near infrared spectroscopy to predict the lignin content and monosaccharide compositions of Pinus radiata wood cell walls.

Near infrared (NIR) spectroscopy coupled with partial least squares (PLS-1) regression was used to predict the lignin contents and monosaccharide compositions of milled wood of Pinus radiata. The effects of particle size and moisture content were investigated by collecting NIR spectra of four sample types: large (<0.422mm) and small (<0.178mm) particles, in both ambient and dry conditions. PLS-1 models were constructed using mixtures of compression wood (CW) and opposite wood (OW) that provided a linear range of cell-wall compositions. Our results show that lignin contents and monosaccharide compositions of pure CWs and OWs can be successfully predicted using NIR spectra of all four sample types. However, large particles in ambient conditions have the most efficient preparation and the standard error (SE) values for lignin (2.10%), arabinose (0.34%), xylose (1.33%), galactose (2.54%), glucose (6.98%), mannose (1.48%), galacturonic acid (0.22%), glucuronic acid (0.06%), and 4-O-methylglucuronic acid (0.25%) were achieved.

Location and characterization of lignin in tracheid cell walls of radiata pine (Pinus radiata D. Don) compression woods.

Tilted stems of softwoods form compression wood (CW) and opposite wood (OW) on their lower and upper sides, respectively. More is known about the most severe form of CW, severe CW (SCW), but mild CWs (MCWs) also occur widely. Two grades of MCWs, MCW1 and MCW2, as well as SCW and OW were identified in the stems of radiata pine (Pinus radiata) that had been slightly tilted. The four wood types were identified by the distribution of lignin in the tracheid walls determined by fluorescence microscopy. A solution of the fluorescent dye acridine orange (AO) (0.02% at pH 6 or 7) was shown to metachromatically stain the tracheid walls and can also be used to determine lignin distribution. The lignified walls fluoresced orange to yellow depending on the lignin concentration. Microscopically well-characterized discs (0.5 mm diameter) of the wood types were used to determine lignin concentrations and lignin monomer compositions using the acetyl bromide method and thioacidolysis, respectively. Lignin concentration and the proportion of p-hydroxyphenyl units (H-units) relative to guaiacyl (G-units) increased with CW severity, with <1% H-units in OW and up to 14% in SCW. Lignin H-units can be used as a marker for CW and CW severity. Similar discs were also examined by Raman and FTIR micro-spectroscopies coupled with principal component analysis (PCA) to determine if these techniques can be used to differentiate the four different wood types. Both techniques were able to do this, particularly Raman micro-spectroscopy.

Distribution of coniferin in differentiating normal and compression woods using MALDI mass spectrometric imaging coupled with osmium tetroxide vapor treatment.

Matrix-assisted laser desorption/ionization mass spectrometric imaging (MALDI-MSI) was employed to detect monolignol glucosides in differentiating normal and compression woods of two Japanese softwoods, Chamaecyparis obtusa and Cryptomeria japonica Comparison of matrix-assisted laser desorption/ionization time-of-flight mass spectrometry collision-induced dissociation fragmentation analysis and structural time-of-flight (MALDI-TOF CID-FAST) spectra between coniferin and differentiating xylem also confirmed the presence of coniferin in differentiating xylem. However, as matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS) and MALDI-TOF CID-FAST spectra of sucrose were similar to those of coniferin, it was difficult to distinguish the distribution of coniferin and sucrose using MALDI-MSI and collision-induced dissociation measurement only. To solve this problem, osmium tetroxide vapor was applied to sections of differentiating xylem. This vapor treatment caused peak shifts corresponding to the introduction of two hydroxyl groups to the C=C double bond in coniferin. The treatment did not cause a peak shift for sucrose, and therefore was effective in distinguishing coniferin and sucrose. Thus, it was found that MALDI-MSI combined with osmium tetroxide vapor treatment is a useful method to detect coniferin in differentiating xylem.

Quantification of (1→4)-ß-d-Galactans in Compression Wood Using an Immuno-Dot Assay.

Compression wood is a type of reaction wood formed on the underside of softwood stems when they are tilted from the vertical and on the underside of branches. Its quantification is still a matter of some scientific debate. We developed a new technique that has the potential to do this based on the higher proportions of (1→4)-ß-d-galactans that occur in tracheid cell walls of compression wood. Wood was milled, partially delignified, and the non-cellulosic polysaccharides, including the (1→4)-ß-d-galactans, extracted with 6 M sodium hydroxide. After neutralizing, the solution was serially diluted, and the (1→4)-ß-d-galactans determined by an immuno-dot assay using the monoclonal antibody LM5, which specifically recognizes this polysaccharide. Spots were quantified using a dilution series of a commercially available (1→4)-ß-d-galactan from lupin seeds. Using this method, compression and opposite woods from radiata pine (Pinus radiata) were easily distinguished based on the amounts of (1→4)-ß-d-galactans extracted. The non-cellulosic polysaccharides in the milled wood samples were also hydrolysed using 2 M trifluoroacetic acid followed by the separation and quantification of the released neutral monosaccharides by high performance anion exchange chromatography. This confirmed that the compression woods contained higher proportions of galactose-containing polysaccharides than the opposite woods.