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Reaction wood is a wood defect arising during the growth of the tree in the part of the trunk that is under tension (hardwood tree species) or compression (coniferous tree species). Beech (Fagus sylvatica L.) tension wood has different anatomical and chemical characteristics than normal (opposite) wood. The difference in density is conditioned by the percentage of the gelatinous layer (G-layer). Fibre cells in reaction beech wood have a different cell wall structure and a different chemical composition. Tension wood cannot be detected by the naked eye. It is only possible to assume its occurrence based on the macroscopic characteristics of the logs, such as a woolly surface, taper or eccentric pith, and so forth. However, these are imprecise and unreliable methods that have minimal effectiveness, especially when shortening the length of the log for cut-outs. This study aimed to create a unique chemical reagent for the detection of tension wood in logs and timber and wood products immediately. The present research can contribute to the mitigation of flaws resulting from the reaction of wood in timber production while addressing noticeable constraints in manufacturing, such as energy resources and the availability of wood raw materials. This can be achieved through the efficient identification of reaction wood in materials. The colour change is only temporary and will fade over time. After the chemical reagent has dried on the surface, the surface can be milled. The colour change extends to a depth of approx. of 3 to 5 mm. [ABSTRACT FROM AUTHOR]

Reaction wood is a wood defect arising during the growth of the tree in the part of the trunk that is under tension (hardwood tree species) or compression (coniferous tree species). Beech (Fagus sylvatica L.) tension wood has different anatomical and chemical characteristics than normal (opposite) wood. The difference in density is conditioned by the percentage of the gelatinous layer (G-layer). Fibre cells in reaction beech wood have a different cell wall structure and a different chemical composition. Tension wood cannot be detected by the naked eye. It is only possible to assume its occurrence based on the macroscopic characteristics of the logs, such as a woolly surface, taper or eccentric pith, and so forth. However, these are imprecise and unreliable methods that have minimal effectiveness, especially when shortening the length of the log for cut-outs. This study aimed to create a unique chemical reagent for the detection of tension wood in logs and timber and wood products immediately. The present research can contribute to the mitigation of flaws resulting from the reaction of wood in timber production while addressing noticeable constraints in manufacturing, such as energy resources and the availability of wood raw materials. This can be achieved through the efficient identification of reaction wood in materials. The colour change is only temporary and will fade over time. After the chemical reagent has dried on the surface, the surface can be milled. The colour change extends to a depth of approx. of 3 to 5 mm.

Summary: Studying biochemical variation in reaction wood is needed to understand the nature of reaction wood formation and the physiology of trees. The activity of antioxidant enzymes (superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and guaiacol peroxidase (GPX)) was studied in normal wood (NW) and reaction wood (RW) (opposite wood (OW) and tension wood (TW)) in the bole of poplar trees (Populus alba L.). Four-year-old trees were induced to produce reaction wood by forced bending. The activity of antioxidant enzymes was studied with repeated sampling during a growing season. The results showed that total protein content and antioxidant enzymes are sensitive to mechanically induced stress. TW and OW showed higher activity of enzymes than NW as well as higher amounts of total protein. OW showed a higher concentration of CAT and SOD than TW. APX was more active than GPX in both TW and OW. In addition to mechanical stress, the effect of other climate factors was observed in the reactive oxygen species accumulation. [ABSTRACT FROM AUTHOR]

While numerous studies have examined microbial attacks on waterlogged archaeological wood, limited information is available regarding microbial attacks in waterlogged tropical hardwoods submerged in marine environments. In this context, we explored microbial attacks in waterlogged archaeological rosewood (Dalbergia species), a tropical hardwood species that was submerged in the Yellow Sea for approximately 700 years, using various microscopic techniques and next-generation sequencing (NGS) methods. Based on morphological features, Type-I soft rot decay was identified as the main decay type. Most fibers in waterlogged archaeological rosewood studied were gelatinous (G) fibers of tension wood and the mode of soft rot decay differed from fibers without the G-layer. Differences in decay resistance between vessel/axial parenchyma cells and fibers were not obvious. Vestured- and simple pit membranes showed higher decay resistance than vessel and axial parenchyma cell walls, respectively. Microbial community analysis by NGS revealed the dominance of Ascomycota and Basidiomycota in the fungal community. Various bacterial communities were also identified, although no prominent signs of bacterial decay were noted. The identified bacterial communities markedly differed from those reported previously in terms of their composition and abundance. Together, our results offer detailed insights into the microbial types and communities responsible for degrading waterlogged archaeological rosewood, contributing to a better understanding of microbial attacks in tropical hardwoods exposed to marine environments. [ABSTRACT FROM AUTHOR]

Abstract Background To gain a better understanding of bark layer structure and function, especially of the phloem fibres and their contribution to the posture control of trees, it is important to map the structural properties of these cells. The role of bark can also be linked to the reaction wood formation and properties which are essential when it comes to studying the questions related to tree growth. To offer new insights into the role of bark in the postural control of trees, we studied the micro- and nanoscale structures of the phloem and its nearest layers. This study is the first time, in which phloem fibres in trees have been extensively examined using X-ray diffraction (XRD). We determined the orientation of cellulose microfibrils in phloem fibres of Silver birch saplings by using scanning synchrotron nanodiffraction. The samples consisted of phloem fibres extracted from tension, opposite and normal wood (TW, OW, NW). Results Using scanning XRD, we were able to obtain new information about the mean microfibril angle (MFA) in cellulose microfibrils in phloem fibres connected to reaction wood. A slight but consistent difference was detected in the average MFA values of phloem fibres between the TW and OW sides of the stem. Using scanning XRD, different contrast agents (intensity of the main cellulose reflection or calcium oxalate reflection, mean MFA value) were used to produce 2D images with 200 nm spatial resolution. Conclusions Based on our results, the tension wood formation in the stem might be related to the structure and properties of phloem fibres. Thus, our results suggest that the nanostructure of phloem fibres is involved in the postural control of trees containing tension and opposite wood.

A cell wall determines themechanical properties of a cell, serves as a barrier against plant stresses, and allows cell division and growth processes. The COBRA-Like (COBL) gene family encodes a putative glycosylphosphatidylinositol (GPI)- anchored protein that controls cellulose deposition and cell progression in plants by contributing to the microfibril orientation of a cell wall. Despite being studied in different plant species, there is a dearth of the comprehensive global analysis of COBL genes in poplar. Poplar is employed as a model woody plant to study abiotic stresses and biomass production in tree research. Improved genome resequencing has enabled the comprehensive exploration of the evolution and functional capacities of PtrCOBLs (Poplar COBRA-Like genes) in poplar. Phylogeny analysis has discerned and classified PtrCOBLs into two groups resembling the Arabidopsis COBL family, and group I genes possess longer proteins but have fewer exons than group II. Analysis of gene structure and motifs revealed PtrCOBLs maintained a rather stable motif and exon-intron pattern across members of the same group. Synteny and collinearity analyses exhibited that the evolution of the COBL gene family was heavily influenced by gene duplication events. PtrCOBL genes have undergone both segmental duplication and tandem duplication, followed by purifying selection. Promotor analysis flaunted various phytohormone-, growthand stress-related cis-elements (e.g., MYB, ABA, MeJA, SA, AuxR, and ATBP1). Likewise, 29 Ptr-miRNAs of 20 families were found targeting 11 PtrCOBL genes. PtrCOBLs were found localized at the plasma membrane and extracellular matrix, while gene ontology analysis showed their involvement in plant development, plant growth, stress response, cellulose biosynthesis, and cell wall biogenesis. RNA-seq datasets depicted the bulk of PtrCOBL genes expression being found in plant stem tissues and leaves, rendering mechanical strength and rejoinders to environmental cues. PtrCOBL2, 3, 10, and 11 manifested the highest expression in vasculature and abiotic stress, and resemblant expression trends were upheld by qRT-PCR. Coexpression network analysis identified PtrCOBL2 and PtrCOBL3 as hub genes across all abiotic stresses and wood developing tissues. The current study reports regulating roles of PtrCOBLs in xylem differentiating tissues, tension wood formation, and abiotic stress latency that lay the groundwork for future functional studies of the PtrCOBL genes in poplar breeding. [ABSTRACT FROM AUTHOR]

Understanding the maturation stress and wood properties of poplar tension wood is critical for improving lumber yields and utilization ratio. In this study, the released longitudinal maturation strains (RLMS), anatomical features, physical and mechanical properties, and nano-mechanical properties of the cell wall were analyzed at different peripheral positions and heights in nine artificially inclined, 12-year-old poplar (Populus × euramericana cv. 'Zhonglin46') trees. The correlations between the RLMS and the wood properties were determined. The results showed that there were mixed effects of inclination on wood quality and properties. The upper sides of inclined stems had higher RLMS, proportion of G-layer, bending modulus of elasticity, and indentation modulus of the cell wall but a lower microfibril angle than the lower sides. At heights between 0.7 m and 2.2 m, only the double-wall thickness increased with height; the RLMS and other wood properties such as fiber length and basic density fluctuated or changed little with height. The RLMS were good indicators of wood properties in the tension wood area and at heights between 0.7 m and 1.5 m. The results of this study present opportunities to better understand the interactions and effects of these two phenomena, which both occur quite frequently in poplar stands and can influence the wood quality of valuable assortments. [ABSTRACT FROM AUTHOR]

Background: To gain a better understanding of bark layer structure and function, especially of the phloem fibres and their contribution to the posture control of trees, it is important to map the structural properties of these cells. The role of bark can also be linked to the reaction wood formation and properties which are essential when it comes to studying the questions related to tree growth. To offer new insights into the role of bark in the postural control of trees, we studied the micro- and nanoscale structures of the phloem and its nearest layers. This study is the first time, in which phloem fibres in trees have been extensively examined using X-ray diffraction (XRD). We determined the orientation of cellulose microfibrils in phloem fibres of Silver birch saplings by using scanning synchrotron nanodiffraction. The samples consisted of phloem fibres extracted from tension, opposite and normal wood (TW, OW, NW). Results: Using scanning XRD, we were able to obtain new information about the mean microfibril angle (MFA) in cellulose microfibrils in phloem fibres connected to reaction wood. A slight but consistent difference was detected in the average MFA values of phloem fibres between the TW and OW sides of the stem. Using scanning XRD, different contrast agents (intensity of the main cellulose reflection or calcium oxalate reflection, mean MFA value) were used to produce 2D images with 200 nm spatial resolution. Conclusions: Based on our results, the tension wood formation in the stem might be related to the structure and properties of phloem fibres. Thus, our results suggest that the nanostructure of phloem fibres is involved in the postural control of trees containing tension and opposite wood. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

A cell wall determines the mechanical properties of a cell, serves as a barrier against plant stresses, and allows cell division and growth processes. The COBRA-Like (COBL) gene family encodes a putative glycosylphosphatidylinositol (GPI)-anchored protein that controls cellulose deposition and cell progression in plants by contributing to the microfibril orientation of a cell wall. Despite being studied in different plant species, there is a dearth of the comprehensive global analysis of COBL genes in poplar. Poplar is employed as a model woody plant to study abiotic stresses and biomass production in tree research. Improved genome resequencing has enabled the comprehensive exploration of the evolution and functional capacities of PtrCOBLs (Poplar COBRA-Like genes) in poplar. Phylogeny analysis has discerned and classified PtrCOBLs into two groups resembling the Arabidopsis COBL family, and group I genes possess longer proteins but have fewer exons than group II. Analysis of gene structure and motifs revealed PtrCOBLs maintained a rather stable motif and exon–intron pattern across members of the same group. Synteny and collinearity analyses exhibited that the evolution of the COBL gene family was heavily influenced by gene duplication events. PtrCOBL genes have undergone both segmental duplication and tandem duplication, followed by purifying selection. Promotor analysis flaunted various phytohormone-, growth- and stress-related cis-elements (e.g., MYB, ABA, MeJA, SA, AuxR, and ATBP1). Likewise, 29 Ptr-miRNAs of 20 families were found targeting 11 PtrCOBL genes. PtrCOBLs were found localized at the plasma membrane and extracellular matrix, while gene ontology analysis showed their involvement in plant development, plant growth, stress response, cellulose biosynthesis, and cell wall biogenesis. RNA-seq datasets depicted the bulk of PtrCOBL genes expression being found in plant stem tissues and leaves, rendering mechanical strength and rejoinders to environmental cues. PtrCOBL2, 3, 10, and 11 manifested the highest expression in vasculature and abiotic stress, and resemblant expression trends were upheld by qRT-PCR. Co-expression network analysis identified PtrCOBL2 and PtrCOBL3 as hub genes across all abiotic stresses and wood developing tissues. The current study reports regulating roles of PtrCOBLs in xylem differentiating tissues, tension wood formation, and abiotic stress latency that lay the groundwork for future functional studies of the PtrCOBL genes in poplar breeding.

In a response to gravitropic stress, G-layers (gelatinous layers) were deposited in xylem cell walls of tilted flax plants. G-layers were produced in both tension wood (upper side) as expected but were also observed in opposite wood (lower side). Raman spectral profiles were acquired for xylem G-layers from the tension and opposite side as well as from the G-layer of bast fibers grown under non-tilted conditions. Statistical analysis by principal component analysis (PCA) and partial least square-discriminant analysis (PLS-DA) clearly distinguished bast fiber G-layers from xylem G-layers. Discriminating bands were observed for cellulose (380–1150–1376 cm–1), hemicelluloses (517–1094–1126–1452 cm–1) and aromatics (1270–1599–1658 cm–1). PCA did not allow separation of G-layers from tension/opposite-wood sides. In contrast, the two types of xylem G-layers could be incompletely discriminated through PLS-DA. Overall, the results suggested that while the architecture (polymer spatial distribution) of bast fibers G-layers and xylem G-layers are similar, they should be considered as belonging to a different cell wall layer category based upon ontogenetical and chemical composition parameters. [ABSTRACT FROM AUTHOR]

Betula platyphylla Suk (Asian white birch) is an economically important tree species in the paper-pulping and biofuel industries. To investigate the mechanism of wood formation at the metabolic level, we evaluated metabolic responses associated with tension-wood formation. Four-year-old trees were subjected to artificial bending treatment for 6 weeks. The xylem growth rate of tension wood (TW) was significantly faster than that of opposite wood (OW), and it exhibited a higher cellulose content. Metabolomics analysis was performed on metabolites of TW, OW and normal wood (NW), and 183 metabolites were identified, of which levels of 142 were altered between groups. Metabolites related to fatty-acid and amino-acid metabolism, the glycolytic pathway, and the metabolism of fructose, mannose and starch sucrose were abundant in TW. Glucose 1-phosphoric acid, fructose and mannose associated with tension-wood development were elevated. Levels of xylitol and ribosol (related to the conversion of glucose), coniferol (the main monomer of lignin) and shikimic acid (an intermediate in lignin synthesis) were decreased in TW. These metabolites are likely involved in xylem development. [ABSTRACT FROM AUTHOR]

Bioenergy will be one of the most important renewable energy sources in the conversion from fossil fuels to bio‐based products. Short rotation coppice Salix could be a key player in this conversion since Salix has rapid growth, positive energy balance, easy to manage cultivation system with vegetative propagation of plant material and multiple harvests from the same plantation. The aim of the present paper is to provide an overview of the main challenges and key issues in willow genetic improvement toward sustainable biofuel value chains. Primarily based on results from the research project "Optimized Utilization of Salix" (OPTUS), the influence of Salix wood quality on the potential for biofuel use is discussed, followed by issues related to the conversion of Salix biomass into liquid and gaseous transportation fuels. Thereafter, the studies address genotypic influence on soil carbon sequestration in Salix plantations, as well as on soil carbon dynamics and climate change impacts. Finally, the opportunities for plant breeding are discussed using willow as a resource for sustainable biofuel production. Substantial phenotypic and genotypic variation was reported for different wood quality traits important in biological (i.e., enzymatic and anaerobic) and thermochemical conversion processes, which is a prerequisite for plant breeding. Furthermore, different Salix genotypes can affect soil carbon sequestration variably, and life cycle assessment illustrates that these differences can result in different climate mitigation potential depending on genotype. Thus, the potential of Salix plantations for sustainable biomass production and its conversion into biofuels is shown. Large genetic variation in various wood and biomass traits, important for different conversion processes and carbon sequestration, provides opportunities to enhance the sustainability of the production system via plant breeding. This includes new breeding targets in addition to traditional targets for high yield to improve biomass quality and carbon sequestration potential. [ABSTRACT FROM AUTHOR]

Leaning stems of woody plants form reaction wood, in hardwood trees termed tension wood (TW). Typical TW fibers, gelatinous fibers (G-fibers), are characterized by an inner gelatinous cell wall layer (G-layer). Xyloglucan endotransglycosylases (XETs) was proposed as the essential enzyme in cell wall modifications in TW, by making xyloglucan (XG) cross-links between G- and S2-layers in G-fibers, and thus maintaining their contact. The determination of TW presence in a sample is of great importance for the forest products industry, biofuel production, and tree physiology studies. However, TW is not easy to detect visually. The colorimetric assay for XET activity determination as an indicator of TW presence in a sample was tested on powdered stem segments of juvenile Populus x euramericana trees exposed to severe long-term static bending. In parallel, histochemical and ultrastructural characterization of stem samples of bent and control trees was performed. The tested colorimetric assay for XET activity determination could be suggested as a useful and easily applicable tool for fast screening of powdered wood samples for the presence of TW. [ABSTRACT FROM AUTHOR]

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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.

The branchwood of fruit trees is being promoted to supplement the fiber material for paper manufacturing in China. This study was conducted to investigate the fiber morphology of walnut branchwood, and to highlight its potential utilizations in papermaking. The effects of the branch diameter, branching level, and tension wood on the fiber morphology were also investigated. The results showed that approximately 65% of the fibers were longer than 900 μm, 95% of the fibers had a slenderness ratio greater than 40, and 67% of the fibers had a Runkel ratio less than 1. It is evident from the results that the fiber morphology of walnut branchwood is reasonably good for paper manufacturing. However, the diameter of the branch does not provide reliable information to predict the fiber morphometrics. In addition, the branching level and tension wood are not very helpful in the screening of fiber raw materials. [ABSTRACT FROM AUTHOR]

Tree growth is influenced by various environmental factors that lead to anatomical, physical and chemical changes in their wood. Reaction wood is one of the tree's reactions that make many restrictions in wood usages. Reaction wood in broadleaf is called tension wood. This study was aimed to stimulate the formation of tension wood in two-year-old seedlings of Populus alba by using intermittent bending, nitrogen fertilization and gibberellin hormone. The application of different treatments increased the content of cellulose compared to the control sample. Meanwhile, the bent specimens had more increase while the straight specimens had no significant difference in the statistical grouping. The content of lignin decreased in all treatments compared to the control sample. The cellulose/lignin ratios obtained from ATR-FTIR (Attenuated Total Reflectance-Fourier Transform Infrared) analysis of wood sawdust and chemical composition measurements were almost close to each other and were higher in the treated samples than in the control. The degree of crystallinity obtained from XRD (X-ray Diffraction) measurements showed that all samples under intermittent bending had a significantly higher degree of crystallinity than the control sample, while this increase was not significant in all straight samples compared to the control sample. In general, it can be concluded that intermittent bending treatment had a greater effect on the stimulation and changes of chemical properties of tension wood in poplar and the application of nitrogen fertilization and gibberellin hormone increased this effect. The formation of gelatinous layer in the innermost part of the intermittent bent seedlings fiber cell wall was visible in light microscope images.

Abstract Background Phytohormones are the key factors regulating vascular development in plants, and they are also involved in tension wood (TW) formation. Although the theory of hormone distribution in TW formation is widely supported, the effects of endogenous hormones on TW formation have not yet been assessed. In this study, TW formation was induced in Catalpa bungei by artificial bending. The phytohormone content of TW, opposite wood (OW) and normal wood (NW) was determined using liquid chromatography-mass spectrometry (LC-MS), and transcriptome sequencing was performed. The hormone content and related gene expression data were comprehensively analyzed. Results The results of analyses of the plant hormone contents indicated significantly higher levels of cis-zeatin (cZ), indoleacetic acid (IAA) and abscisic acid (ABA) in TW than in OW. Genes involved in the IAA and ABA synthesis pathways, such as ALDH (evm.model.group5.1511) and UGT (evm.model.scaffold36.20), were significantly upregulated in TW. and the expression levels of ARF (evm.model.group5.1332), A-ARR (evm.model.group0.1600), and TCH4 (evm.model.group2.745), which participate in IAA, cZ and Brassinolide (BR) signal transduction, were significantly increased in TW. In particular, ARF expression may be regulated by long noncoding RNAs (lncRNAs) and the HD-ZIP transcription factor ATHB-15. Conclusions We constructed a multiple hormone-mediated network of C. bungei TW formation based on hormone levels and transcriptional expression profiles were identified during TW formation.

Lignin in Populus species is acylated with p -hydroxybenzoate. Monolignol p -hydroxybenzoyltransferase 1 (PHBMT1) mediates p -hydroxybenzoylation of sinapyl alcohol, eventually leading to the modification of syringyl lignin subunits. Angiosperm trees upon gravistimulation undergo the re-orientation of their growth along with the production of specialized secondary xylem, i.e., tension wood (TW), that generates tensile force to pull the inclined stem or leaning branch upward. Sporadic evidence suggests that angiosperm TW contains relatively a high percentage of syringyl lignin and lignin-bound p -hydroxybenzoate. However, whether such lignin modification plays a role in gravitropic response remains unclear. By imposing mechanical bending and/or gravitropic stimuli to the hybrid aspens in the wild type (WT), lignin p -hydroxybenzoate deficient, and p -hydroxybenzoate overproduction plants, we examined the responses of plants to gravitropic/mechanical stress and their cell wall composition changes. We revealed that mechanical bending or gravitropic stimulation not only induced the overproduction of crystalline cellulose fibers and increased the relative abundance of syringyl lignin, but also significantly induced the expression of PHBMT1 and the increased accumulation of p -hydroxybenzoates in TW. Furthermore, we found that although disturbing lignin-bound p -hydroxybenzoate accumulation in the PHBMT1 knockout and overexpression (OE) poplars did not affect the major chemical composition shifts of the cell walls in their TW as occurred in the WT plants, depletion of p -hydroxybenzoates intensified the gravitropic curving of the plantlets in response to gravistimulation, evident with the enhanced stem secant bending angle. By contrast, hyperaccumulation of p -hydroxybenzoates mitigated gravitropic response. These data suggest that PHBMT1-mediated lignin modification is involved in the regulation of poplar gravitropic response and, likely by compromising gravitropism and/or enhancing autotropism, negatively coordinates the action of TW cellulose fibers to control the poplar wood deformation and plant growth. [ABSTRACT FROM AUTHOR]

Lignin is a complex polymer in plant cell walls whose proportion is second only to that of cellulose and plays an important role in the mechanical properties of wood and stress resistance of plants. Here, we induced tension wood (TW) formation in Catalpa bungei by artificial bending and analyzed the lignin metabolism of the TW. LC-MS analysis showed that a significantly higher content of coniferyl aldehyde was observed in the TW cell wall than in the opposite wood (OW) and normal wood (NW) cell walls. TW had significantly lower contents of coniferyl alcohol than OW and NW. Raman spectroscopy results indicated that TW had lower total lignin than OW and NW. The transcription and translation levels of most of the differentially expressed genes (DEGs) involved in lignin monomer biosynthesis indicated upregulation in TW/OW and TW/NW. We found no significant difference in the transcription levels of three collision gases (CADs) between TW and OW or between NW, but their translation levels were significantly downregulated in TW, suggesting post-transcriptional control for CAD. We predicted and analyzed transcription factors that could target DEGs involved in lignin monomer biosynthesis in TW. Based on the analysis of the relationships of targeting and coexpression, we found that NAC (evm.model.group1.695) could potentially target 4CLs and CCoAOMT , that HD-Zip (evm.model.group7.1157) had potential targeting relationships with CCoAOMT , F5H , and CCR , and that their expression levels were significantly positive. It is speculated that the upregulation of NAC and HD-ZIP transcription factors activates the expression of downstream target genes, which leads to a significant increase in coniferyl aldehyde in TW. However, the decrease in total lignin in TW may be caused by the significant downregulation of CAD translation and the significant decrease in precursors (coniferyl alcohol). Whether the expression of CAD genes is regulated by post-transcriptional control and affects TW lignin metabolism needs further study. [ABSTRACT FROM AUTHOR]

SUMMARY: Cell walls, especially secondary cell walls (SCWs), maintain cell shape and reinforce wood, but their structure and shape can be altered in response to gravity. In hardwood trees, tension wood is formed along the upper side of a bending stem and contains wood fiber cells that have a gelatinous layer (G‐layer) inside the SCW. In a previous study, we generated nst/snd quadruple‐knockout aspens (Populus tremula × Populus tremuloides), in which SCW formation was impaired in 99% of the wood fiber cells. In the present study, we produced nst/snd triple‐knockout aspens, in which a large number of wood fibers had thinner SCWs than the wild type (WT) and some had no SCW. Because SCW layers are always formed prior to G‐layer deposition, the nst/snd mutants raise interesting questions of whether the mutants can form G‐layers without SCW and whether they can control their postures in response to changes in gravitational direction. The nst/snd mutants and the WT plants showed growth eccentricity and vessel frequency reduction when grown on an incline, but the triple mutants recovered their upright growth only slightly, and the quadruple mutants were unable to maintain their postures. The mutants clearly showed that the G‐layers were formed in SCW‐containing wood fibers but not in those lacking the SCW. Our results indicate that SCWs are essential for G‐layer formation and posture control. Furthermore, each wood fiber cell may be able to recognize its cell wall developmental stage to initiate the formation of the G‐layer as a response to gravistimulation. [ABSTRACT FROM AUTHOR]

Lignin in Populus species is acylated with p-hydroxybenzoate. Monolignol p-hydroxybenzoyltransferase 1 (PHBMT1) mediates p-hydroxybenzoylation of sinapyl alcohol, eventually leading to the modification of syringyl lignin subunits. Angiosperm trees upon gravistimulation undergo the re-orientation of their growth along with the production of specialized secondary xylem, i.e., tension wood (TW), that generates tensile force to pull the inclined stem or leaning branch upward. Sporadic evidence suggests that angiosperm TW contains relatively a high percentage of syringyl lignin and lignin-bound p-hydroxybenzoate. However, whether such lignin modification plays a role in gravitropic response remains unclear. By imposing mechanical bending and/or gravitropic stimuli to the hybrid aspens in the wild type (WT), lignin p-hydroxybenzoate deficient, and p-hydroxybenzoate overproduction plants, we examined the responses of plants to gravitropic/mechanical stress and their cell wall composition changes. We revealed that mechanical bending or gravitropic stimulation not only induced the overproduction of crystalline cellulose fibers and increased the relative abundance of syringyl lignin, but also significantly induced the expression of PHBMT1 and the increased accumulation of p-hydroxybenzoates in TW. Furthermore, we found that although disturbing lignin-bound p-hydroxybenzoate accumulation in the PHBMT1 knockout and overexpression (OE) poplars did not affect the major chemical composition shifts of the cell walls in their TW as occurred in the WT plants, depletion of p-hydroxybenzoates intensified the gravitropic curving of the plantlets in response to gravistimulation, evident with the enhanced stem secant bending angle. By contrast, hyperaccumulation of p-hydroxybenzoates mitigated gravitropic response. These data suggest that PHBMT1-mediated lignin modification is involved in the regulation of poplar gravitropic response and, likely by compromising gravitropism and/or enhancing autotropism, negatively coordinates the action of TW cellulose fibers to control the poplar wood deformation and plant growth.

Lignin is a complex polymer in plant cell walls whose proportion is second only to that of cellulose and plays an important role in the mechanical properties of wood and stress resistance of plants. Here, we induced tension wood (TW) formation in Catalpa bungei by artificial bending and analyzed the lignin metabolism of the TW. LC-MS analysis showed that a significantly higher content of coniferyl aldehyde was observed in the TW cell wall than in the opposite wood (OW) and normal wood (NW) cell walls. TW had significantly lower contents of coniferyl alcohol than OW and NW. Raman spectroscopy results indicated that TW had lower total lignin than OW and NW. The transcription and translation levels of most of the differentially expressed genes (DEGs) involved in lignin monomer biosynthesis indicated upregulation in TW/OW and TW/NW. We found no significant difference in the transcription levels of three collision gases (CADs) between TW and OW or between NW, but their translation levels were significantly downregulated in TW, suggesting post-transcriptional control for CAD. We predicted and analyzed transcription factors that could target DEGs involved in lignin monomer biosynthesis in TW. Based on the analysis of the relationships of targeting and coexpression, we found that NAC (evm.model.group1.695) could potentially target 4CLs and CCoAOMT, that HD-Zip (evm.model.group7.1157) had potential targeting relationships with CCoAOMT, F5H, and CCR, and that their expression levels were significantly positive. It is speculated that the upregulation of NAC and HD-ZIP transcription factors activates the expression of downstream target genes, which leads to a significant increase in coniferyl aldehyde in TW. However, the decrease in total lignin in TW may be caused by the significant downregulation of CAD translation and the significant decrease in precursors (coniferyl alcohol). Whether the expression of CAD genes is regulated by post-transcriptional control and affects TW lignin metabolism needs further study.

Summary: Plant cellulose is synthesized by a large plasma membrane‐localized cellulose synthase (CesA) complex. However, an overall functional determination of secondary cell wall (SCW) CesAs is still lacking in trees, especially one based on gene knockouts.Here, the Cas9/gRNA‐induced knockouts of PtrCesA4, 7A, 7B, 8A and 8B genes were produced in Populus trichocarpa. Based on anatomical, immunohistochemical and wood composition evidence, we gained a comprehensive understanding of five SCW PtrCesAs at the genetic level.Complete loss of PtrCesA4, 7A/B or 8A/B led to similar morphological abnormalities, indicating similar and nonredundant genetic functions. The absence of the gelatinous (G) layer, one‐layer‐walled fibres and a 90% decrease in cellulose in these mutant woods revealed that the three classes of SCW PtrCesAs are essential for multilayered SCW structure and wood G‐fibre. In addition, the mutant primary and secondary phloem fibres lost the n(G + L)‐ and G‐layers and retained the thicker S‐layers (L, lignified; S, secondary). Together with polysaccharide immunolocalization data, these findings suggest differences in the role of SCW PtrCesAs‐synthesized cellulose in wood and phloem fibre wall structures.Overall, this functional understanding of the SCW PtrCesAs provides further insights into the impact of lacking cellulose biosynthesis on growth, SCW, wood G‐fibre and phloem fibre wall structures in the tree. [ABSTRACT FROM AUTHOR]

In this function, "counterparts" of a particular I Arabidopsis i gene means flax and poplar genes that best-hit to the particular I Arabidopsis i gene in a Blast search. In this case, I Arabidopsis i genes whose counterparts in flax and poplar meet each search condition are extracted and shown with the counterparts in flax and poplar. Each gene in the network is labeled by its symbol or the gene symbol of the corresponding I Arabidopsis i gene if the flax/poplar gene does not have a symbol. Keywords: database; flax; plant fibers; poplar; tension wood; tertiary cell wall; transcriptome EN database flax plant fibers poplar tension wood tertiary cell wall transcriptome 512 515 4 06/18/21 20210715 NES 210715 Acknowledgements The authors thank Prof. M. Deyholos (University of British Columbia, Canada) who provided seeds of the I rdf i mutant and corresponding background cultivar that were used to analyze transcriptomes in a specific sample set from an unpublished study. [Extracted from the article]

Hygrothermal treatment induces irreversible dimensional changes of green wood i.e. hygrothermal recovery (HTR). To understand what happened to cellulose-rich gelatinous (G-) layer in green tension wood during HTR, changes in vibrational properties of konara oak (Quercus serrata Thunb. ex Murray) tension wood (TW) and normal wood (NW) collected from sapwood after hygrothermal treatment were tested regarding HTR. After this treatment, all specimens were air-dried, and their vibrational properties and dimensions were measured in this dried state. The hygrothermal treatment induced an increase in mechanical loss tangent (tanδ) and a decrease in specific dynamic Young's modulus (E'/ρ). Changes in vibrational properties due to hygrothermal treatment appeared to depend on treatment time and temperature with higher temperatures and longer treatment durations producing larger increases in tanδ and larger decreases in E'/ρ. In TW with a G-layer, a clear correlation between changes in vibrational properties and HTR strains was identified. Tanδ increased and E'/ρ decreased corresponding to HTR strains. Contraction of the G-layer in TW cell walls due to release of locked-in growth stress by hygrothermal treatment seems to be the most plausible mechanism underlying changes in vibrational properties and generation of HTR. In NW without a G-layer, HTR strain was below the limit of detection, which obscures potential correlations. Differences in the intensities of changes in vibrational properties after 120 min hygrothermal treatments at 60, 80, and 100 °C were not significant after drying; however, the difference in the intensities of HTR strains apparently remained after drying. [ABSTRACT FROM AUTHOR]

Tension wood is a type of defect of wood, however, it has some especial character and structure. In this study, cellulase lignin structures in normal and tension wood of Poplar 107 (Populus × euramericana cv. '74/76') were compared using ultraviolet (UV), Fourier transform infrared (FT-IR), Raman and magnetic resonance (NMR) spectroscopy, gel permeation chromatography (GPC) and elemental analysis. The results showed that the lignins in both normal and tension wood were dominated by syringyl units, followed by guaiacyl and p -hydroxyphenyl units. The FT-IR result presented that the relative intensity of syringyl units in normal and tension wood were 0.87 and 0.90. The contents of aliphatic hydroxyl, methoxyl, condensed phenolic hydroxyl and carboxyl of lignin in tension wood were higher than those of phenolic hydroxyl groups and β-5. The contents of β-O-4, β-β and β-1 in tension wood were similar with those in normal wood. The average per-C 9 -unit formulae of the lignin in normal and tension wood of Poplar 107 were C 9 H 7.21 O 1.79 (OH) 1.17_x0001_ (OCH 3) 1.55 and C 9 H 7.23 O 1.76 (OH) 1.25_x0001_ (OCH 3) 1.62 , respectively. [ABSTRACT FROM AUTHOR]

In response to gravistimulation under anisotropic light, tree stems showing an active cambium produce reaction wood that redirects the axis of the trees. Several studies have described transcriptomic or proteomic models of reaction wood relative to the opposite wood. However, the mechanisms leading to the formation of reaction wood are difficult to decipher because so many environmental factors can induce various signalling pathways leading to this developmental reprogramming. Using an innovative isotropic device where the phototropic response does not interfere with gravistimulation we characterized the early molecular responses occurring in the stem of poplar after gravistimulation in an isotropic environment, and without deformation of the stem. After 30 min tilting at 35° under anisotropic light, we collected the upper and lower xylems from the inclined stems. Controls were collected from vertical stems. We used a microarray approach to identify differentially expressed transcripts. High-throughput real-time PCR allowed a kinetic experiment at 0, 30, 120 and 180 min after tilting at 35°, with candidate genes. We identified 668 differentially expressed transcripts, from which we selected 153 candidates for additional Fluidigm qPCR assessment. Five candidate co-expression gene clusters have been identified after the kinetic monitoring of the expression of candidate genes. Gene ontology analyses indicate that molecular reprogramming of processes such as 'wood cell expansion', 'cell wall reorganization' and 'programmed cell death' occur as early as 30 min after gravistimulation. Of note is that the change in the expression of different genes involves a fine regulation of gibberellin and brassinosteroid pathways as well as flavonoid and phosphoinositide pathways. Our experimental set-up allowed the identification of genes regulated in early gravitropic response without the bias introduced by phototropic and stem bending responses. [ABSTRACT FROM AUTHOR]

Cellulose nanofibrils (CNFs) are top-down nanomaterials obtainable from abundant lignocelluloses. Despite recent advances in processing technologies, the effects of variations in the lignocellulose structure and composition on CNF isolation and properties are poorly understood. In this study, we compared the isolation of CNFs from tension wood (TW) and normal wood (NW) from Populus tremula (aspen). The TW has a higher cellulose content, native cellulose fibrils with a larger crystalline diameter, and less lignin than the NW, making it an interesting material for CNF isolation. The wood powders were oxidized directly by 2,2,6,6-tetramethylpiperidin-1-oxyl, and the morphology and mechanical behaviors of the nanofibril suspensions and networks were characterized. The TW was more difficult to fibrillate by both chemical and mechanical means. Larger nanofibrils (5–10 nm) composed of 1.2 nm structures were present in the TW CNFs, whereas the NW samples contained more of thin (1.6 nm) structures, which also comprised 77% of the solid yield compared to the 33% for TW. This difference was reflected in the TW CNF networks as decreased transmittance (15% vs. 50%), higher degree of crystallinity (85.9% vs. 78.0%), doubled toughness (11 MJ/m3) and higher elongation at break (12%) compared to NW. The difference was ascribed to greater preservation of the hierarchical, more crystalline microfibril structure, combined with a more cellulose-rich network (84% vs. 70%). This knowledge of the processing, structure, and properties of CNFs can facilitate the breeding and design of wood feedstocks to meet the increasing demand for nanoscale renewable materials. [ABSTRACT FROM AUTHOR]

In angiosperm trees, the gelatinous layer (G-layer) takes a great part of the fiber cell wall in the tension wood (TW). However, the mechanism underlying G-layer formation in poplar is largely unknown. In this work, we demonstrate that G-layer formation in poplar TW cells is regulated by brassinosteroid (BR) and its signaling. PtiCYP85A3, a key BR biosynthesis gene, was predominantly expressed in the xylem of TW, accompanied with a relatively higher castasterone (CS) accumulation, than in the xylem of opposite wood (OW). A wider expression zone of BZR1, a key transcriptional factor in BR singling pathway, was also observed in G-fiber cells on TW side than in wood fiber cells on the OW side, as indicated by immunohistochemistry assays. Transgenic poplar plants overexpressing PtiCYP85A3 produced thicker G-layer with higher cellulose proportion, and accumulated more BZR1 protein in the xylem of TW than did the wild type (WT) plants. Expression of most TW-associated CesAs, which were induced by 2, 4-epibrassinolide, an active BR, and inhibited by brassinazole, a BR biosynthesis inhibitor, were also up-regulated in the xylem of TW in transgenic plants compared to that in WT plants. Further studies with dual-luciferase assays demonstrated that the promoters of PtiCesAs were activated by PtiMYB128, a TW specific transcription factor, which was then regulated by BZR1. All these results indicate that BR plays a crucial role in the G-layer formation of TW fiber cells by regulating the expression of BZR1, PtiMYB128, and PtiCesAs in poplar.

Abstract Background Tension wood is a type of reaction wood in response to bending or leaning stem as a corrective growth process. Tension wood is formed by both natural and man-made processes. Most attractively, tension wood contains higher glucan content and undergoes higher enzymatic conversion to fermentable sugars. Here, we have employed structural techniques, small-angle neutron scattering (SANS) and wide-angle X-ray diffraction (WAXD) to elucidate structural and morphological aspects of tension wood conducive to higher sugar yields. Results Small-angle neutron scattering data exhibited a tri-modal distribution of the fibril cross-sectional dimension. The smallest size, 22 Å observed in all samples concurred with the WAXD results of the control and opposite side samples. This smallest and the most abundant occurring size was interpreted as the cellulose elementary microfibril diameter. The intermediate size of 45 Å, which is most pronounced in the tension side sample and consistent with WAXD results for tension side sample, indicates association of neighboring elementary microfibrils to form larger crystallite bundles. The largest size 61 Å observed by SANS was however not observed by WAXD and therefore associated to mesopores. Conclusions Structure and morphology of tension wood is different from control wood. Cellulose crystallinity increases, lignin content is lower and the appearance of mesopores with 61 Å diameter is observed. Despite the presence of higher crystalline cellulose content in tension side, the lower lignin content and may be combined with the abundance of mesopores, substantially improves enzyme accessibility leading to higher yields in cellulose digestion.

The aim of the study is to characterise the cellular structures in tension wood and compare its effect on the selected anatomical parameters in two mangrove wood species viz. Excoecaria agallocha Linn. and Sonneratia caseolaris (L.) Engl. Wood blocks of 1 cm3 were prepared and thin sections of cross sections were cut using microtome and stained with Azure II to differentiate the structure of normal wood from tension wood fibres. The anatomical parameters analysed in this present study were vessel frequency, tangential vessel diameter and fibre wall thickness in tension wood as well as in normal wood. Vessel diameter and vessel frequency were significantly lower in tension wood compared to the normal wood in both the species. The reduction of tangential diameter in tension wood region is a way of protecting these elements from cavitation and providing mechanical support for the plant than liquid transport. A significant variation in fibre wall thickness was observed between tension wood and normal wood in both species. [ABSTRACT FROM AUTHOR]

Key message: The important anatomical changes in tension wood, e.g., the high fiber ratio and rich mesopores, did not significantly increase the air and nitrogen flow; thus the gas permeability in the longitudinal direction of poplar (Populus deltoidesCL.’55/65′) tension wood is actually affected by the cell tissue macroporous porosity. Context: Gas permeability is one of the most important physical properties of wood and is closely related to its internal microstructure, particularly porosity. Tension wood is widespread in woody plants and displays significant structural differences compared with opposite wood. Aims: The study was designed to clarify the relationship between pore structure and gas permeability in poplar tension wood. Methods: The gas permeability was measured using a self-made device. The meso- and macroporosity characteristics were measured by nitrogen adsorption–desorption and mercury intrusion porosimetry. The flow was simulated using ANSYS Fluent software to illustrate the role of pore structure on permeability. Results: The morphological features of vessels have an effect on wood permeability. Compared with tension wood, opposite wood, which has higher vessel ratio, larger cell lumen diameter, and more rich pits, shows stronger gas permeability. Increasing the airflow path will actually reduce the gas permeability. The simulation results are consistent with the experimental results. Conclusion: In hardwoods, the gas permeability in the longitudinal direction is mainly dictated by the vessels. The high fiber ratio and rich mesopore in tension wood do not significantly increase gas flow, suggesting the permeability of wood was actually determined by the cell tissue with macroporous porosity. Vessel tissue ratio, length and diameter, and intervessel pit size were found responsible for influencing the permeability in the longitudinal direction. [ABSTRACT FROM AUTHOR]

The structure of hardwoods representing eight tropical and five temperate species was characterized from the atomistic level up to the cellular level using X-ray scattering, X-ray microtomography and light microscopy. The species were chosen for this study based on their popularity as tonewoods. The ultrastructure of wood cell walls, including crystallite size, orientation and close-range order of cellulose microfibrils were determined by small- and wide-angle X-ray scattering (SAXS, WAXS). The SAXS patterns were interpreted by using an analytical model of cylinders packed in a hexagonal close-range order with paracrystalline distortion. The values for the cylinder diameters given by this model were compared to the average crystallite widths obtained by WAXS using the Scherrer equation. In six out of 26 samples, all of these representing tropical species used especially in fretboard parts of electric guitars, large differences between these two sizes were obtained. The WAXS and microscopy results of these samples corresponded to tension wood structures. These comparisons and interpretations of SAXS results have not been previously presented for any tropical hardwoods, especially related to those containing tension wood tissue. The importance of the ultrastructural characterization was highlighted in this study in the case of tropical hardwood samples. [ABSTRACT FROM AUTHOR]

In angiosperm trees, the gelatinous layer (G-layer) takes a great part of the fiber cell wall in the tension wood (TW). However, the mechanism underlying G-layer formation in poplar is largely unknown. In this work, we demonstrate that G-layer formation in poplar TW cells is regulated by brassinosteroid (BR) and its signaling. PtiCYP85A3 , a key BR biosynthesis gene, was predominantly expressed in the xylem of TW, accompanied with a relatively higher castasterone (CS) accumulation, than in the xylem of opposite wood (OW). A wider expression zone of BZR1, a key transcriptional factor in BR singling pathway, was also observed in G-fiber cells on TW side than in wood fiber cells on the OW side, as indicated by immunohistochemistry assays. Transgenic poplar plants overexpressing PtiCYP85A3 produced thicker G-layer with higher cellulose proportion, and accumulated more BZR1 protein in the xylem of TW than did the wild type (WT) plants. Expression of most TW-associated CesAs , which were induced by 2, 4-epibrassinolide, an active BR, and inhibited by brassinazole, a BR biosynthesis inhibitor, were also up-regulated in the xylem of TW in transgenic plants compared to that in WT plants. Further studies with dual-luciferase assays demonstrated that the promoters of PtiCesAs were activated by PtiMYB128, a TW specific transcription factor, which was then regulated by BZR1. All these results indicate that BR plays a crucial role in the G-layer formation of TW fiber cells by regulating the expression of BZR1 , PtiMYB128 , and PtiCesAs in poplar. [ABSTRACT FROM AUTHOR]

Under the effect of disturbances, like unbalanced stem, but also during normal development, poplar trees can develop a specific secondary xylem, called "tension wood" (TW), which is easily identifiable by the presence of a gelatinous layer in the secondary cell walls (SCW) of the xylem fibers. Since TW formation was mainly performed on 2-year-old poplar models, an in vitro poplar that produces gelatinous fibers (G-fibers) while offering the same experimental advantages as herbaceous plants has been developed. Using specific cell wall staining techniques, wood structural features and lignin/cellulose distribution were both detailed in cross-sections obtained from the curved stem part of in vitro poplars. A supposed delay in the SCW lignification process in the G-fibers, along with the presence of a G-layer, could be observed in the juvenile plants. Moreover, in this G-layer, the immunolabeling of various polymers carried out in the SCW of TW has allowed detecting crystalline cellulose, arabinogalactans proteins, and rhamnogalacturonans I; however, homogalacturonans, xylans, and xyloglucans could not be found. Interestingly, extensins were detected in this typical adaptative or stress-induced structure. These observations were corroborated by a quantitation of the immunorecognized polymer distribution using gold particle labeling. In conclusion, the in vitro poplar model seems highly convenient for TW studies focusing on the implementation of wall polymers that provide the cell wall with greater plasticity in adapting to the environment. [ABSTRACT FROM AUTHOR]

Betula platyphylla Suk (birch) is an excellent short-term hardwood species with growth and wood characteristics well suited to wood industries. To investigate the molecular mechanism of wood development in birch, a tension wood (TW) induced system was used to explore the regulatory mechanism at the protein level and identify the key proteins involved in xylem development in birch. The results of dyeing with Safranin O-Fast Green indicated that the cellulose content of TW was significantly higher than that of opposite wood (OW) or normal wood (NW), and the lignin content in TW was significantly lower than that in OW and NW after artificial bending of birch stems. Protein profile analysis of TW, NW and OW by iTRAQ revealed that there were 639 and 460 differentially expressed proteins (DEPs) between TW/OW and TW/NW, respectively. The DEPs were mainly enriched in tyrosine metabolism, glycolysis/gluconeogenesis, phenylalanine and tyrosine metabolism, phenylpropanoid and pyruvate metabolism, the pentose phosphate pathway, the citrate cycle (TCA cycle), fructose and mannose metabolism, carbon fixation in photosynthetic organisms, fatty acid biosynthesis, photosynthesis proteins and other pathways. The proteins in the citrate cycle were upregulated. The expression levels of PGI, PGM and FRK proteins related to cellulose synthesis increased and the expression levels of PAL, 4CL and COMT related to lignin synthesis decreased, leading to an increase in cellulose content and decreased lignin levels in TW. PPI analysis revealed that key DEPs interact with each other, indicating that these proteins form complexes to implement this function, which may provide important insights for wood formation at the molecular level. • Identified 639 and 460 DEPs in TW/OW and TW/NW of birch by iTRAQ. • The expression levels of proteins related to cellulose synthesis increased, and related to lignin synthesis decreased. • In TW, the C flow towards cellulose synthesis increases, while the C flow towards lignin synthesis decreases. • PPI analysis revealed that key DEPs in cellulose synthesis and lignin synthesis interact with each other. [ABSTRACT FROM AUTHOR]

Tension wood (TW) in hybrid aspen trees forms on the upper side of displaced stems to generate a strain that leads to uplifting of the stem. TW is characterized by increased cambial growth, reduced vessel frequency and diameter, and the presence of gelatinous, cellulose-rich (G-)fibers with its microfibrils oriented parallel to the fiber cell axis. Knowledge remains limited about the molecular regulators required for the development of this special xylem tissue with its characteristic morphological, anatomical, and chemical features. In this study, we use transgenic, ethylene-insensitive (ETI) hybrid aspen trees together with time-lapse imaging to show that functional ethylene signaling is required for full uplifting of inclined stems. X-ray diffraction and Raman microspectroscopy of TW in ETI trees indicate that, although G-fibers form, the cellulose microfibril angle in the G-fiber S-layer is decreased, and the chemical composition of S- and G-layers is altered than in wild-type TW. The characteristic asymmetric growth and reduction of vessel density is suppressed during TW formation in ETI trees. A genome-wide transcriptome profiling reveals ethylene-dependent genes in TW, related to cell division, cell wall composition, vessel differentiation, microtubule orientation, and hormone crosstalk. Our results demonstrate that ethylene regulates transcriptional responses related to the amount of G-fiber formation and their properties (chemistry and cellulose microfibril angle) during TW formation. The quantitative and qualitative changes in G-fibers are likely to contribute to uplifting of stems that are displaced from their original position.

Caffeic acid O-methyltransferase (COMT) is an important protein that participates in lignin synthesis and is associated with the ratio of G-/S-type lignin in plants. COMTs are associated with the wood properties of forest trees; however, little known about the COMT family in Catalpa bungei, a valuable timber tree species in China . We performed a comprehensive analysis of COMT genes in the C. bungei genome by describing the gene structure and phylogenetic relationships of each family member using bioinformatics-based methods. A total of 23 putative COMT genes were identified using the conserved domain sequences and amino acid sequences of COMTs from Arabidopsis thaliana and Populus trichocarpa as probes. Phylogenetic analysis showed that 23 CbuCOMTs can be divided into three groups based on their structural characteristics; five conserved domains were found in the COMT family. Promoter analysis indicated that the CbuCOMT promoters included various cis-acting elements related to growth and development. Real-time quantitative polymerase chain reaction (PCR) analysis showed differential expression among CbuCOMTs. CbuCOMT2, 7, 8, 9, 10, 12, 13, 14, 21, and 23 were mainly expressed in xylem. Only CbuCOMT23 was significantly downregulated in tension wood and upregulated in opposite wood compared to normal wood. Our study provides new information about the CbuCOMT gene family and will facilitate functional characterisation in further research.