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Quantitative anatomical characteristics and their radial variation in compression (COW), lateral (LAW), and opposite (OPW) woods of Agathis loranthifolia and Pinus merkusii stem woods growing in Indonesia were observed and compared to understand wood quality. The length, diameter, wall thickness, and lumen diameter of tracheids and ray height and numbers were observed using optical microscopy. In both species, COW had the shortest tracheid length, smallest tracheid and lumen diameter, thickest cell wall, and highest ray numbers among the parts, while LAW and OPW showed comparable or variable values in quantitative characteristics. In A. loranthifolia, COW had the highest ray height, whereas, in P. merkusii, it had the lowest uniseriate and fusiform ray heights. No significant difference was observed in ray numbers and heights between LAW and OPW. In both species, the tracheid length and lumen-to-diameter ratio in COW, LAW, and OPW tended to increase from near the pith to near the bark while the wall-to-diameter ratio decreased. The ray heights of all parts increased with increasing distance from the pith, whereas the ray number decreased.

This research investigated and compared the pitting type, pit number (PN), and pit diameter (PD) in the cross-field of compression wood (CWD), lateral wood (LWD), and opposite wood (OWD) in stem wood of Pinus merkusii and Agathis loranthifolia growing in Indonesia. Identification and quality evaluation were done using optical and scanning electron microscopy. A piceoid pit type was observed in the CWD of both species. The LWD and OWD of P. merkusii showed window-like and pinoid pits, whereas those of A. loranthifolia showed taxodioid and cupressoid pits. The PN and PD were the smallest in CWD of both species. In P. merkusii, LWD and OWD showed similar PN values, and PN in all parts increased from the pith to the bark. In A. loranthifolia, LWD had higher PN than in OWD, and PN in CWD and LWD decreased from near the pith to the bark, whereas in OWD, it increased. All parts of P. merkusii and CWD and OWD of A. loranthifolia showed a positive correlation between PN and radial tracheid diameter, whereas LWD showed a negative correlation. In P. merkusii, the PD of LWD approximated that of OWD, whereas, in A. loranthifolia, LWD had a larger PD than that exhibited by OWD.

This research investigated and compared the pitting type, pit number (PN), and pit diameter (PD) in the cross-field of compression wood (CWD), lateral wood (LWD), and opposite wood (OWD) in stem wood of Pinus merkusii and Agathis loranthifolia growing in Indonesia. Identification and quality evaluation were done using optical and scanning electron microscopy. A piceoid pit type was observed in the CWD of both species. The LWD and OWD of P. merkusii showed window-like and pinoid pits, whereas those of A. loranthifolia showed taxodioid and cupressoid pits. The PN and PD were the smallest in CWD of both species. In P. merkusii, LWD and OWD showed similar PN values, and PN in all parts increased from the pith to the bark. In A. loranthifolia, LWD had higher PN than in OWD, and PN in CWD and LWD decreased from near the pith to the bark, whereas in OWD, it increased. All parts of P. merkusii and CWD and OWD of A. loranthifolia showed a positive correlation between PN and radial tracheid diameter, whereas LWD showed a negative correlation. In P. merkusii, the PD of LWD approximated that of OWD, whereas, in A. loranthifolia, LWD had a larger PD than that exhibited by OWD. [ABSTRACT FROM AUTHOR]

Quantitative anatomical characteristics and their radial variation in compression (COW), lateral (LAW), and opposite (OPW) woods of Agathis loranthifolia and Pinus merkusii stem woods growing in Indonesia were observed and compared to understand wood quality. The length, diameter, wall thickness, and lumen diameter of tracheids and ray height and numbers were observed using optical microscopy. In both species, COW had the shortest tracheid length, smallest tracheid and lumen diameter, thickest cell wall, and highest ray numbers among the parts, while LAW and OPW showed comparable or variable values in quantitative characteristics. In A. loranthifolia, COW had the highest ray height, whereas, in P. merkusii, it had the lowest uniseriate and fusiform ray heights. No significant difference was observed in ray numbers and heights between LAW and OPW. In both species, the tracheid length and lumen-to-diameter ratio in COW, LAW, and OPW tended to increase from near the pith to near the bark while the wall-to-diameter ratio decreased. The ray heights of all parts increased with increasing distance from the pith, whereas the ray number decreased.

Quantitative anatomical characteristics and their radial variation in compression (COW), lateral (LAW), and opposite (OPW) woods of Agathis loranthifolia and Pinus merkusii stem woods growing in Indonesia were observed and compared to understand wood quality. The length, diameter, wall thickness, and lumen diameter of tracheids and ray height and numbers were observed using optical microscopy. In both species, COW had the shortest tracheid length, smallest tracheid and lumen diameter, thickest cell wall, and highest ray numbers among the parts, while LAW and OPW showed comparable or variable values in quantitative characteristics. In A. loranthifolia, COW had the highest ray height, whereas, in P. merkusii, it had the lowest uniseriate and fusiform ray heights. No significant difference was observed in ray numbers and heights between LAW and OPW. In both species, the tracheid length and lumen-to-diameter ratio in COW, LAW, and OPW tended to increase from near the pith to near the bark while the wall-to-diameter ratio decreased. The ray heights of all parts increased with increasing distance from the pith, whereas the ray number decreased. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Radial growth of trees can result in opposite wood (OW) and compression wood (CW) due to the varying impact of stem mechanical stress, such as that caused by gravity or wind. Previous research has identified higher xylem production in CW compared to OW. Yet, it remains unclear whether the difference in the number of xylem cells between OW and CW results from differences in growth rate or the duration of xylem cells. In this study, we collected wood microcores on a weekly basis from March 2019 to January 2020 in Pinus massoniana Lamb. located on a steep slope. Our objective was to compare the dynamic of cambial activity and resulting cellular anatomical parameters between OW and CW in a humid subtropical environment. Our results showed that the xylem phenology of OW and CW was generally consistent with the xylem cell division process beginning in early March and ceasing in November. The last latewood cell completed its differentiation at the end of December. The response of wood formation dynamics to climate was consistent in both OW and CW. Moreover, both wood types exhibited a limited development of the enlargement phase due to the heat and drought during the summer. The rate of cell division was responsible for 90.7% of the variability in the number of xylem cells. The CW xylem obtained a larger number of cells by increasing the rate of cell division and displayed thinner earlywood cells with larger lumens than OW cells. Our findings showed that the xylem of conifer species responds to mechanical stress by accelerating the cell division rate. As a result, we suggest calculating the ratio between OW and CW widths to reconstruct wind stress changes rather than calculating the residuals used in the current study.

The increasing demand on the wood market makes it necessary to use all secondary resources, including the wood branches and spindles. The properties of deciduous branches and softwood spindles were evaluated in order to create wood panels, highlighting the transverse texture with high value added. The research analyzed the properties of the wood from deciduous (beech, maple) branches and from conifer (spruce, fir) spindles. The methodology of obtaining panels with transversal structure was also presented. The production of minor wood (branches or spindles) was 254 m3/ha/year in the case of beech (branches) and 109 m3/ha/year in the case of spruce (spindles). The proportion of compression wood for spruce had an average value of 40.6%. A similar value of tension wood for beech was obtained. The modulus of elasticity for pine spindle wood was 71.7% lower than the modulus of elasticity for bending strength in the trunk. The quality index was 36.7% lower in the case of spindles than in the case of pine trunk, and the value of the quality ratio in branch maple wood was 2.61% lower than in trunk wood. Thus, even if it is not the only efficient solution, wood in the minor portions can be used to make high value-added panels such as those that highlight the transverse structure.

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]

Summary: Spiral grain refers to the helical patterns formed by the wood grain in the trunks of many tree species. In most gymnosperms, grain near the pith is vertical but wood formed after several years of growth has a slight to pronounced left-handed twist. Grain changes presumably involve the slow rotation of cells within the vascular cambium, but the mechanisms that allow this reorientation to occur remain unclear. Understanding this process is, however, important as the presence of strong spiral grain within the corewood of gymnosperms is a major wood quality issue devaluing cut timber. In this study, we measured wood grain in stems of Pinus radiata (radiata pine) saplings through reconstructions of resin canals that follow the grain, visualised by serial sectioning and scanning with circularly polarised light, and through X-ray computed microtomography (μ CT) and image analysis in ImageJ. Vertical trees retained a symmetrical grain pattern that was weakly right-handed near the pith, but which became progressively more left-handed during the first eight months of growth. In tilted trees, however, the development of left-handed grain was inhibited by the formation of compression wood on the lower side of the tree whereas the wood on the upper side of the tree developed increasingly more left-handed grain as in the vertical controls. These results demonstrate that a previously unidentified link exists between compression wood formation and the inhibition of grain development. [ABSTRACT FROM AUTHOR]

Summary: The quantification of the intensities of tree growth responses to the impact of geomorphic processes is a modern research trend in dendrogeomorphology. It enables a more sensitive assessment of the activity of the studied geomorphic process compared to the traditional use of growth disturbances. The advanced definitions of individual intensity classes of growth disturbances are based exclusively on macroscopic observations. This study evaluates the possibility of anatomical quantification of compression wood (CW) intensity in the case of common spruce (Picea abies (L.) Karst.) and compares it with subjective macroscopic evidence of CW with respect to stem tilting intensity. In total, 25 disturbed (tilted) individuals of P. abies occupying a landslide active during July 1997 were sampled, and 21 of them were analysed. The intensity of external disturbance (stem tilting) was compared against the macroscopic (intensity and duration) and microscopic (quantitative change of the tracheid lumen area and the cell wall proportion) parameters of compression wood suitable for practical application in common dendrogeomorphic analysis. Generally, the macroscopic indices of CW were strongly correlated with stem tilting. The intensity of the anatomical growth response was stronger in the earlywood zone than in the latewood zone. Nevertheless, their dependence on stem tilting was not detected. Results suggest that CW classification based on the quantification of anatomical changes is not possible for dendrogeomorphic purposes. Nevertheless, based on the obtained results, the present study suggests preferring the most intensively tilted trees during future dendrogeomorphic research to obtain the most intensive macroscopic and subjective observable anatomical evidence for studying compression wood. [ABSTRACT FROM AUTHOR]

A single Norway spruce tree (Picea abies) was manually fractionated into heartwood, sapwood, juvenile wood and branches. These fractions were chemically pulped, individually, in laboratory scale. The pulps were characterized and investigated in relation to dewatering behavior and sheet strength properties. An unbleached and unbeaten commercial kraft pulp from softwood fibers was used as a reference, and the fractionated pulps were within the same range in all tested properties. The fractionated pulps were then compared with each other, and fiber characteristics were used to explain differences in dewatering and strength. Heartwood pulp results in stronger and stiffer papers that are harder to dewater. Sapwood pulp gives more open network structures resulting in easy dewatering and high air permeance, although with lower strength properties compared to heartwood. Pulp from Juvenile wood gives s quite strong but brittle sheets, with efficient dewatering. Pulp from branches gives paper sheets with efficient dewatering, air permeance and relatively high elongation of break but lower strength. The results show that there is definitely potential for utilizing more parts of the trees for pulp and paper making, especially when tailoring the raw material origins after preferred paper properties. [ABSTRACT FROM AUTHOR]

The Taxodium hybrid Zhongshanshan fast-growing species is susceptible to environment and gravity to form reaction wood. In this study, individual growth rings of reaction wood are used as subjects, and an individual growth ring is divided into three zones: compression zone (CZ), lateral zone (LZ), and opposite zone (OZ). The microanatomical structure and chemical properties of the tracheids in CZ, LZ, and OZ forms by the inclined or bent growth of T. Zhongshanshan are comparatively analyzed by using optical microscopy, scanning electron microscope, laser confocal microscopy, and Raman imaging techniques. In CZ, the length and diameter of compression wood (CW) tracheids decreased, and the shape of cross-sections became rounded as compared to the OZ and LZ tracheids. More notably, threaded fissures appeared on the cell wall of tracheids, and the thickness of the cell wall increased in CW. The analysis of tracheids' cell wall structure showed that CW tracheids had a complete outer secondary wall middle (S2L) layer, but had no secondary wall inner (S3) layer. In the transition zone (TA) between CW and normal early wood, tracheids were divided into compressed and normal tracheids. Despite the compressed tracheids having a similar cell morphology to normal tracheids, they had a thin secondary wall S2L layer. Tracheids in LZ had a thin S2L layer only at the angle of the cell. No S2L layer was seen in the cell wall of OZ and CZ late wood tracheids. It can be concluded that the response of lignin deposition location to external stress was faster than the change in cell morphology. The above results help provide the theoretical basis for the response mechanism of T. Zhongshanshan reaction wood anatomical structures to the external environment and has important theoretical value for understanding its characteristics and its rational and efficient usage. [ABSTRACT FROM AUTHOR]

The increasing demand on the wood market makes it necessary to use all secondary resources, including the wood branches and spindles. The properties of deciduous branches and softwood spindles were evaluated in order to create wood panels, highlighting the transverse texture with high value added. The research analyzed the properties of the wood from deciduous (beech, maple) branches and from conifer (spruce, fir) spindles. The methodology of obtaining panels with transversal structure was also presented. The production of minor wood (branches or spindles) was 254 m3 /ha/year in the case of beech (branches) and 109 m3 /ha/year in the case of spruce (spindles). The proportion of compression wood for spruce had an average value of 40.6%. A similar value of tension wood for beech was obtained. The modulus of elasticity for pine spindle wood was 71.7% lower than the modulus of elasticity for bending strength in the trunk. The quality index was 36.7% lower in the case of spindles than in the case of pine trunk, and the value of the quality ratio in branch maple wood was 2.61% lower than in trunk wood. Thus, even if it is not the only efficient solution, wood in the minor portions can be used to make high value-added panels such as those that highlight the transverse structure [ABSTRACT FROM AUTHOR]

A new laboratory reactor to perform in situ studies of structural changes in wood during soda pulping using synchrotron X-ray tomography is presented. The reactor is of recirculation type to provide stable reaction conditions and mimic the industrial situation. Experiments have been performed using this reactor in situ at a synchrotron microtomography beamline to provide sequences of 3D images from which measurement of wood cell wall thickness have been possible for the first time. The results showed that the cell wall thickness increased significantly in the early stage of pulping (<10 min), which is due to the transportation of cooking chemicals through the tracheids, resin channels and pits into the cell wall, which is swollen with the increased pH. Subsequently, the cell wall thickness reduces over the processing time up to 60 min with a high rate, which is inferred to be due to the dissolution and transport of lignin and hemicellulose from the secondary walls, allowing for better transportation of active chemicals deep through the cell wall layers. After 60 min processing, the cell wall thickness reduction rate reduced, as dissolution of lignin and hemicelluloses from the cell walls ceased, while the remaining dissolution occurs mainly at the middle lamella. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract The microtensile properties of mechanically isolated compression wood (CW) and opposite wood (OW) tracheids of Chinese fir (Cunninghamia lanceolata) were investigated and discussed with respect to their structure. Major differences in the tensile modulus and ultimate tensile stress were found between CW and OW fibers. Compared to OW, CW showed a larger cellulose microfibril angle, less cellulose content and probably more pits, resulting in lower tensile properties. These findings contribute to a further understanding of the structural–mechanical relationships of Chinese fir wood at the cell and cell wall level, and provide a scientific basis for better utilization of plantation softwood.

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.

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. [ABSTRACT FROM AUTHOR]

Forested mountain slopes can be simultaneously affected by soil creep and landslide activity, both of which cause the tilting of tree stems, with the result that their dendrochronological record of tree-ring eccentricity and compression wood is potentially similar. There is a need to identify similarities and differences in these records and thus our research aimed to compare patterns of eccentricity and compression wood developed by trees under the impact of soil creep and landslides. We sampled trees growing on a landslide and creeping slopes in the Kamienne Mts., with 21 Norway spruce trees were sampled on each site. We found several main differences between the dendrochronological record of landslide activity and soil creep. On the landslide we found larger number of dendrochronological events, stronger and more variable eccentricity and a similar number of upslope and downslope events. On creeping slopes, upslope eccentricity events predominate, and the number of eccentricity events dated in all trees increases in time. We also compared the utility of eccentricity and compression wood for dating mass movements. They differ in their sensitivity to stem tilting. Thus, in analyses of landslide activity and soil creep activity, it is recommended to include both wood anatomy features. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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.

The Taxodium hybrid Zhongshanshan fast-growing species is susceptible to environment and gravity to form reaction wood. In this study, individual growth rings of reaction wood are used as subjects, and an individual growth ring is divided into three zones: compression zone (CZ), lateral zone (LZ), and opposite zone (OZ). The microanatomical structure and chemical properties of the tracheids in CZ, LZ, and OZ forms by the inclined or bent growth of T. Zhongshanshan are comparatively analyzed by using optical microscopy, scanning electron microscope, laser confocal microscopy, and Raman imaging techniques. In CZ, the length and diameter of compression wood (CW) tracheids decreased, and the shape of cross-sections became rounded as compared to the OZ and LZ tracheids. More notably, threaded fissures appeared on the cell wall of tracheids, and the thickness of the cell wall increased in CW. The analysis of tracheids’ cell wall structure showed that CW tracheids had a complete outer secondary wall middle (S2L) layer, but had no secondary wall inner (S3) layer. In the transition zone (TA) between CW and normal early wood, tracheids were divided into compressed and normal tracheids. Despite the compressed tracheids having a similar cell morphology to normal tracheids, they had a thin secondary wall S2L layer. Tracheids in LZ had a thin S2L layer only at the angle of the cell. No S2L layer was seen in the cell wall of OZ and CZ late wood tracheids. It can be concluded that the response of lignin deposition location to external stress was faster than the change in cell morphology. The above results help provide the theoretical basis for the response mechanism of T. Zhongshanshan reaction wood anatomical structures to the external environment and has important theoretical value for understanding its characteristics and its rational and efficient usage.

Summary: 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 S2L 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. [ABSTRACT FROM AUTHOR]

Ensuring the quality and stability of trees as the most important components of the plant community has attracted the attention of foresters to identify and measure defects and ensure safety coefficients. Therefore, in the present study, the use of acoustic tomography to calculate the extent of decay, the area of the woody section and the safety factor of black pine was studied. Acoustic tomography was performed using 16 sensors at two elevations of 147 and 200 cm above the ground and on 8 tree in the Sopron region of Hungary. The results showed that the stress wave technique was an accurate method for determining safety coefficients and determining the amount of internal decay in the tree and also had the ability to identify compression areas in cross section. Resistography of the studied sections in different geographical directions assisted to locating the woody areas, which was compatible with the information of acoustic tomography. The findings of this study showed that the area of the canopy has a significant role in the safety of the tree and the possibility of tree fall, so that in some trees due to their condition, with doubling the area of the canopy, the torque increased from 80827 to 161655 N.m and the condition of the tree changed from low-risk to high-risk. [ABSTRACT FROM AUTHOR]

The microtensile properties of mechanically isolated compression wood (CW) and opposite wood (OW) tracheids of Chinese fir (Cunninghamia lanceolata) were investigated and discussed with respect to their structure. Major differences in the tensile modulus and ultimate tensile stress were found between CW and OW fibers. Compared to OW, CW showed a larger cellulose microfibril angle, less cellulose content and probably more pits, resulting in lower tensile properties. These findings contribute to a further understanding of the structural–mechanical relationships of Chinese fir wood at the cell and cell wall level, and provide a scientific basis for better utilization of plantation softwood. [ABSTRACT FROM AUTHOR]

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.

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.

To understand the irreversible dimensional changes caused by hygrothermal treatment of green wood, i.e. hygrothermal recovery (HTR), green hinoki compression wood (CW) and normal wood (NW) were hygrothermally (HT) treated in water at 100°C for 120 min and their HTR strains were determined. The specimens were then swollen using dimethyl sulfoxide (DMSO) and then completely dried after solvent exchange with water at room temperature. Their HTR strains were then compared with their DMSO swelling and drying shrinkage strains. The volumetric HTR strains in the CW were about twice as large as those in the NW. Moreover, the microfibril angle (MFA) was found to be an important factor for controlling the HTR intensity. A clear commonality between the HTR behavior and both DMSO swelling and drying shrinkage behavior was identified, which indicates that HTR is caused by volumetric changes in the matrix substances. HTR has been defined as a phenomenon due to the release of locked-in growth stress when a wood specimen is HT treated. To determine whether DMSO treatment has a similar effect as hygrothermal treatment, both HT-untreated and HT-treated specimens were swollen using DMSO, and their dimensional changes during and after DMSO treatment were compared. The results showed that DMSO treatment is a possible alternative for releasing the locked-in growth stress. [ABSTRACT FROM AUTHOR]

To achieve efficient utilization of compression wood (CW), a deeper insight into the molecular interactions is necessary. In particular, the role of lignin in the wood needs to be better understood, especially concerning how lignin contributes to its mechanical properties. For this reason, the properties of CW and normal wood (NW) from Chinese fir (Cunninghamia lanceolata) have been studied on a molecular scale by means of polarized Fourier transform infrared (FTIR) spectroscopy, under both static and dynamic loading conditions. Under static tensile loading, only molecular deformations of cellulose were observed in both CW and NW. No participation of lignin could be detected. In relation to the macroscopic strain, the molecular deformation of the cellulose C-O-C bond was greater in NW than in CW as a reflection of the higher microfibril angle and the lower load taken up by CW. Under dynamic deformation, a larger contribution of the lignin to stress transfer was detected in CW; the molecular deformation of the lignin being highly related to the amplitude of the applied stress. Correlation analysis indicated that there was a direct coupling between lignin and cellulose in CW, but there was no evidence of such a direct coupling in NW. [ABSTRACT FROM AUTHOR]

The characteristics of cross-field pitting among compression wood, lateral wood, and opposite wood, in the stem woods of Ginkgo biloba and Pinus densiflora were investigated with optical and scanning electron microscopy. In Ginkgo biloba , compression wood exhibited piceoid pits, while lateral and opposite wood exhibited cupressoid pits. The compression wood of Pinus densiflora exhibited cupressoid pits and piceoid pits, while lateral wood and opposite wood exhibited pinoid and window-like pits in the cross-field. In both species, compression wood yielded the smallest pit number among each part, while opposite wood yielded the greatest pit number per cross-field. Cross-field pitting diameters of compression wood and opposite wood were significantly smaller than lateral wood in Ginkgo biloba , while the cross-field pitting of compression wood was the smallest in Pinus densiflora. Radial tracheid diameter of compression wood was slightly smaller than lateral and opposite wood in Ginkgo biloba and significantly smaller than lateral and opposite wood in Pinus densiflora. In conclusion, the cross-field pitting type, pit number, and cross-field pitting diameter could be used to identify reaction wood in the stem wood of Ginkgo biloba and Pinus densiflora. [ABSTRACT FROM AUTHOR]

The color changes of compression and opposite spruce (Picea abies L. Karst) wood were evaluated as a function of different drying conditions. Compression wood samples and their corresponding opposite wood samples from the opposite part of the log were compared after three different drying modes. The color of the samples before and after the drying was characterized using the color coordinates L*, a*, b*, C*ab, C, h*, Sab, and the color difference (ΔE*) in the standard color space according to the International Commission on Illumination (CIELAB). The drying temperature was the most remarkable factor for discoloration of the samples. The lightness (L*) of the samples decreased with increased temperature. However, darkening was more pronounced in the opposite wood samples. The overall color difference (ΔE*) was found to be higher in the opposite wood. The compression wood was more statured in color with a deeper hue angle due to the drying process. [ABSTRACT FROM AUTHOR]

Conifers, as a response to mechanical stress, such as wind and stem lean, form reaction wood called compression wood (CW). CW occurs in a range of gradations from near normal wood (NW) to severe CW (SCW). As the severity of CW affects the mechanical and chemical properties of wood, and as CW has limited value in the forest products industry, it is desirable to be able to measure CW severity. Picea omorika belong to slow-growing conifer species in which CW typically occurs in a severe form. We developed different morphometric and non-morphometric methods for estimation of CW severity tested on wood samples of P. omorika juvenile trees exposed to long term static bending. This specific review is aimed at presenting P. omorika as one of the most adaptable spruces, and as a good model for testing of methods for estimation of compression wood severity. First, we summarize main knowledge about P. omorika, features of CW, and methods for assessment of wood quality. Then, we present breifly our recently published methods for estimation of compression wood severity tested on P. omorika juvenile wood samples. [ABSTRACT FROM AUTHOR]

Winter is a critical season for land‐surface feedbacks and ecosystem processes; however, most high‐latitude paleo‐environmental reconstructions are blind to cold season conditions. Here we introduce a winter‐sensitive, paleo‐proxy record that is based on the relative frequency of tangential rows of traumatic resin ducts (TRDs) in the annual growth rings of mountain hemlocks (Tsuga mertensiana) growing near treeline in Southeast Alaska. Hemlocks produce a row of TRDs in the earlywood portion of their annual rings in response to cambial damage incurred during winter. Multidecadal bouts of TRD production were followed by growth‐leader replacement, reaction wood formation, and divergence in radial growth between storm‐damaged trees and less exposed mountain hemlock forests. These patterns are consistent with TRDs being a response to tree damage caused by ice and snowstorms, a conclusion supported by the krummholz tree architecture at these sites. This relationship is further corroborated by significant correlations between our TRD record and the strength of the wintertime Aleutian Low (AL) pressure system that is linked to tree‐damaging agents like wind, precipitation, and ice storm strength in Southeast Alaska. The combined TRD/krummholz architecture record indicates that abrupt shifts between strong and weak AL phases occurred every several decades since CE 1700 and that the 1800s had relatively long AL phases with heavy snowpacks. In addition to describing the magnitude and tempo of wintertime climate change in Northwestern North America, these results suggest that North Pacific Decadal Variability underlies the long‐term dynamics of treeline ecosystems along the northeast Pacific coast. Key Points: Traumatic resin ducts in the cross‐dated tree rings of subalpine mountain hemlocks record tree damage during winter storms in Southeast AlaskaWarmer, windier winters that accompany strong Aleutian Low pressure systems result in more tree damageDecadal‐scale variability in North Pacific climate has impacted treeline ecosystems throughout this region [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

To clarify the mechanism of irreversible dimensional changes due to the hygrothermal treatment of green wood, i.e., the hygrothermal recovery (HTR), the changes in vibrational properties of compression wood and normal wood were measured after hygrothermal treatment at 60 °C, 80 °C, and 100 °C. In addition, the relationship between those changes and HTR strains were discussed. The hygrothermal treatment induced an increase in mechanical loss tangent (tanδ) and decrease in specific dynamic Young’s modulus (E′/ρ). It seems that the changes in vibrational properties due to hygrothermal treatment had a time-temperature dependency: Higher temperatures and longer treatment durations induce larger increases in tanδ and larger decreases in E′/ρ. In contrast to the quenching effect, tanδ and E′/ρ did not recover to their original state even after 60 days of conditioning in water at 20 °C. For compression wood with a large microfibril angle (MFA), there was a clear relationship between the changes in vibrational properties and HTR strains. The tanδ increased and E′/ρ decreased with hygrothermal treatment, corresponding to dimensional changes in the L-direction. This suggests that structural changes in wood components are responsible for HTR. The most likely mechanism for HTR is that the hygrothermal treatment softens the lignin to release locked-in growth stress. Subsequently, irreversible structural changes in lignin induce both the changes in vibrational properties and HTR. For normal wood, because of the small MFA, the structural changes in lignin in the L-direction are possibly restricted by crystalline cellulose. As a result, the relationship between the changes in vibrational properties and HTR is uncertain. [ABSTRACT FROM AUTHOR]

Tree stems deviating from the vertical position react by the formation of tension wood (TW) or compression wood (CW), which are called in general as reaction wood (RW), in which the cells are modified chemically and anatomically. The focus of the present work is the mechanical behavior of TW in five 37-year-old Eucalyptus species, which were grown on a planting area with an average slope of 28% leading to decentralized pith in the trees, which is an unequivocal indication of the presence of RW. TW and opposite wood (OW) samples were isolated and subjected to a compression-parallel-to-grain test. It was observed that TW is less resistant and less stiff than the OW. [ABSTRACT FROM AUTHOR]