Characterizing compression wood formed in radiata pine branches

IAWA Journal ◽  
2014 ◽  
Vol 35 (4) ◽  
pp. 385-394
Author(s):  
Xinguo Li ◽  
Robert Evans ◽  
Washington Gapare ◽  
Xiaohui Yang ◽  
Harry X. Wu
Keyword(s):  
Compression Wood ◽  
Microfibril Angle ◽  
Marked Change ◽  
Radiata Pine ◽  
Wood Formation ◽  
Reaction Wood ◽  
Opposite Wood ◽  
Cambial Age ◽  
Cellulose Microfibril Orientation ◽  
Secondary Wall Formation

The formation of reaction wood is an adaptive feature of trees in response to various mechanical forces. In gymnosperms, reaction wood consists of compression wood (CW) and opposite wood (OW) that are formed on the underside and upperside of bent trunks and branches. Although reaction wood formed in bent trunks has been extensively investigated, relatively little has been reported from conifer branches. In this study SilviScan® technology was used to characterize radiata pine branches at high resolution. Compared to OW formed in the branches, CW showed greater growth, darker colour, thicker tracheid walls, higher coarseness, larger microfibril angle (MFA), higher wood density, lower extensional stiffness and smaller internal specific surface area. However, tracheids of CW were similar to those of OW in their radial and tangential diameters. These results indicated that gravity influenced tracheid cell division and secondary wall formation but had limited impact on primary wall expansion. Furthermore, seasonal patterns of CW formation were not observed in the branches from cambial age 4 while earlywood and latewood were clearly separated in all rings of OW. The marked change of MFA during reaction wood formation suggested that branches could be ideal materials for further study of cellulose microfibril orientation.

WITHIN-TREE VARIATION IN ANATOMICAL PROPERTIES OF COMPRESSION WOOD IN RADIATA PINE

IAWA Journal ◽  
2004 ◽  
Vol 25 (3) ◽  
pp. 253-271 ◽  
Author(s):  
Lloyd A. Donaldson ◽  
Jenny Grace ◽  
Geoff M. Downes
Keyword(s):  
Wall Thickness ◽  
Compression Wood ◽  
Microfibril Angle ◽  
Basic Density ◽  
Radiata Pine ◽  
Lumen Diameter ◽  
Normal Wood ◽  
Lignin Distribution ◽  
Opposite Wood ◽  
Anatomical Properties

Two trees of radiata pine, one showing severe lean, the other growing almost vertically, were assessed for the presence and anatomical properties of compression wood, including anatomy, lignin distribution, microfibril angle, basic density, radial and tangential lumen diameter and cell wall thickness. Both trees contained significant amounts of compression wood although the severity and amount of compression wood was greater in the leaning tree. Changes in lignin distribution seem to be characteristic of the mildest forms of compression wood with reduced lignification of the middle lamella representing the earliest change observed from normal wood. An increase in microfibril angle was associated with both mild and severe compression wood although examples of severe compression wood with the same or smaller microfibril angles than opposite wood, or with very small microfibril angles, were found. When segregated into mild and severe compression wood the average difference in microfibril angle was 4° and 8° respectively compared with opposite wood. Within-ring distribution of microfibril angle was different in severe compression wood compared to opposite wood with higher angles in the latewood.Severe compression wood showed a 22% increase in basic density compared to mild compression wood and opposite wood. The increased density was accounted for in terms of a 26% increase in tracheid wall thickness throughout the growth ring, offset by a 9% increase in radial lumen diameter, slightly greater in the latewood. There were no significant changes in density or cell dimensions in mild compression wood compared with opposite wood.

Compression Wood does not Form in the Roots of Pinus Radiata

IAWA Journal ◽  
2006 ◽  
Vol 27 (1) ◽  
pp. 45-54 ◽  
Author(s):  
Linda C.Y. Hsu ◽  
John C.F. Walker ◽  
Brian G. Butterfield ◽  
Sandra L. Jackson
Keyword(s):  
Mechanical Stress ◽  
Pinus Radiata ◽  
Compression Wood ◽  
Lateral Roots ◽  
Compressive Load ◽  
Radiata Pine ◽  
Wood Formation ◽  
Wall Layer ◽  
Reaction Wood ◽  
Wood Compression

We investigated the potential for the roots of Pinus radiata D. Don to form compression wood. Compression wood was not observed in either the tap or any lateral roots further than 300 mm from the base of the stem. This suggests that either the roots do not experience the stresses required to induce compression wood formation, or that they lack the ability to form it. Roots artificially subjected to mechanical stress also failed to develop compression wood. It is therefore unlikely that an absence of a compressive load on buried roots can account for the lack of compression wood. Application of auxin to the cambia of lateral roots was similarly ineffective at inducing the formation of compression wood. These observations suggest that the buried roots of radiata pine lack the ability to develop compression wood. We also report the formation of an atypical S3 wall layer in the mechanically-stressed and auxin-treated tracheids and suggest that a reaction wood that is different to compression wood may well form in roots.

scholarly journals Developmental Trends of Black Spruce Fibre Attributes in Maturing Plantations

2016 ◽  
Vol 2016 ◽  
pp. 1-12 ◽  
Author(s):  
Peter F. Newton
Keyword(s):  
Black Spruce ◽  
Microfibril Angle ◽  
Basal Area ◽  
Temporal Correlation ◽  
Wood Formation ◽  
Cross Sectional ◽  
Forest Region ◽  
Cambial Age ◽  
Canadian Boreal Forest ◽  
Crown Closure

This study assessed the temporal developmental patterns of commercially relevant fibre attributes (tracheid length and diameters, wall thickness, specific surface area, wood density, microfibril angle, fibre coarseness, and modulus of elasticity) and their interrelationships within maturing black spruce (Picea mariana (Mill.) B.S.P.) plantations. A size-based stratified random sample procedure within 5 semimature plantations located in the Canadian Boreal Forest Region was used to select 50 trees from which radial cross-sectional xylem sequences at breast-height (1.3 m) were cut and analyzed. Statistically, the graphical and linear correlation analyses indicated that the attributes exhibited significant (p≤0.05) relationships among themselves and with morphological tree characteristics. Relative variation of each annually measured attribute declined with increasing size class (basal area quintile). The transitional shifts in temporal correlation patterns occurring at the time of approximate crown closure where suggestive of intrinsic differences in juvenile and mature wood formation processes. The temporal cumulative development patterns of all 8 of the annually measured attributes varied systematically with tree size and exhibited the most rapid rates of change before the trees reached a cambial age of 20 years. At approximately 50 years after establishment, plantation mean attribute values were not dissimilar from those reported for more mature natural-origin stands.

Cross-field pitting characteristics of compression, lateral, and opposite wood in the stem wood of Ginkgo biloba and Pinus densiflora

IAWA Journal ◽  
2020 ◽  
Vol 41 (1) ◽  
pp. 48-60
Author(s):  
Byantara Darsan Purusatama ◽  
Nam Hun Kim
Keyword(s):  
Electron Microscopy ◽  
Scanning Electron Microscopy ◽  
Ginkgo Biloba ◽  
Compression Wood ◽  
Pinus Densiflora ◽  
Reaction Wood ◽  
Stem Wood ◽  
Opposite Wood ◽  
The Cross ◽  
Scanning Electron

Abstract 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.

Strength and stiffness of the reaction wood in five Eucalyptus species

Holzforschung ◽  
2019 ◽  
Vol 73 (2) ◽  
pp. 219-222
Author(s):  
Bruno Charles Dias Soares ◽  
José Tarcísio Lima ◽  
Selma Lopes Goulart ◽  
Claudineia Olímpia de Assis
Keyword(s):  
Mechanical Behavior ◽  
Tension Wood ◽  
Compression Wood ◽  
Vertical Position ◽  
Eucalyptus Species ◽  
Reaction Wood ◽  
Opposite Wood ◽  
Average Slope ◽  
Strength And Stiffness

AbstractTree 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-oldEucalyptusspecies, 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.

The nature of reaction wood. VIII. The structure and differentiation of compression wood

1964 ◽  
Vol 12 (1) ◽  
pp. 24 ◽  
Author(s):  
AB Wardrop ◽  
GW Davies
Keyword(s):  
Compression Wood ◽  
Early Stage ◽  
Wood Formation ◽  
Ground Substance ◽  
Reaction Wood ◽  
Intercellular Spaces ◽  
Chemically Induced ◽  
Cell Wall Organization ◽  
Cytoplasmic Organization ◽  
Cytoplasmic Ground Substance

The cell wall organization of tracheids of natural and chemically induced compression wood of Pinus radiata and Actinostrobus pyramidalis has been shown to be the same, and is similar to that established in previous studies of natural compression wood. In the secondary wall only two layers were present. In the second of these there was a well-developed system of helical cavities, separating ribs of cellulose. The ribs of cellulose were parallel to the direction of microfibril orientation; were complex in form; and the cellulose lamellae lay parallel with the wall surface. A well-developed wart structure was present. During the differentiation of compression wood tracheids, the intercellular spaces were formed during the phase of surface enlargement of the differentiating tracheids. At an early stage the intercellular spaces appeared to contain cytoplasmic ground substance. During the development of the layer S1 the cytoplasmic organization was similar to that of normal tracheids, the cells containing a large vacuole with a well-developed tonoplast and plasmalemma. During the development of the layer S2 the cytoplasm contained numerous small vesicles with no large vacuoles, and in many instances the plasmalemma was absent. At the conclusion of the differentiation of the cell the plasmalemma was again present and penetrated the helical cavities of the wall. Compression wood induced by 3-indoleacetic acid (IAA) alone, gibberellic acid (GA) alone, or IAA and GA in combination was identical with that formed under natural conditions. The localized lateral application of IAA to vertical stems caused conspicuous bending of the stem as well as compression wood formation.

scholarly journals The Nature of Reaction Wood III. Cell Division and Cell Wall Formation in Conifer Stems

1952 ◽  
Vol 5 (4) ◽  
pp. 385 ◽  
Author(s):  
ABW Ardrop ◽  
HE Dadswell
Keyword(s):  
Cell Wall ◽  
Cell Division ◽  
Secondary Wall ◽  
Compression Wood ◽  
Normal Wood ◽  
Reaction Wood ◽  
Primary Wall ◽  
Wall Formation ◽  
Tracheid Length ◽  
Secondary Wall Formation

Cell division, the nature of extra-cambial readjustment, and the development of the secondary wall in the tracheids of conifer stems have been investigated in both compression wood and normal wood. It has been shown that the reduction in tracheid length, accompanying the development of compression wood and, in normal wood, increased radial growth after suppression, result from an increase in the number of anticlinal divisions in the cambium. From observations of bifurcated and otherwise distorted cell tips in mature tracheids, of small but distinct terminal canals connecting the lumen to the primary wall in the tips of mature tracheids, and of the presence of only primary wall at the tips of partly differentiated tracheids, and from the failure to observe remnants of the parent primary walls at the ends of differentiating tracheids, it has been concluded that extra-cambial readjustment of developing cells proceeds by tip or intrusive growth. It has been further concluded that the development of the secondary wall is progressive towards the cell tips, on the bases of direct observation of secondary wall formation in developing tracheids and of the increase found in the number of turns of the micellar helix per cell with increasing cell length. The significance of this in relation to the submicroscopic organization of the cell wall has been discussed. Results of X-ray examinations and of measurements of� tracheid length in successive narrow tangential zones from the cambium into the xylem have indicated that secondary wall formation begins before the dimensional changes of differentiation are complete.

Lignin and carbohydrate variation with earlywood, latewood, and compression wood content of bent and straight ramets of a radiata pine clone

Holzforschung ◽  
2010 ◽  
Vol 64 (1) ◽  
Author(s):  
R. Paul Kibblewhite ◽  
Ian D. Suckling ◽  
Robert Evans ◽  
Jennifer C. Grace ◽  
Mark J.C. Riddell
Keyword(s):  
Radial Direction ◽  
Compression Wood ◽  
Juvenile Wood ◽  
Radiata Pine ◽  
Carbohydrate Content ◽  
Strongly Correlated ◽  
Growth Layer ◽  
Opposite Wood ◽  
Radial Dimension ◽  
Wood Content

Abstract Changes in lignin and carbohydrate content with radial direction, growth-layer number, compression wood (CW) severity, and earlywood (EW) and latewood (LW) origin are described for one near-ground position in each of a severely bent and a nominally straight ramet (tree) of a clone (genotype) of Pinus radiata. Bark-to-bark strips were taken through the pith and the longest radial dimension of the CW side of the discs. Separate EW and LW samples were obtained for most growth layers, yielding a total of 95 samples. Differences in lignin and carbohydrate content between EW and LW were large where CW formation was moderate and small where it was severe. Mannose content was consistently different in the EW and LW of opposite wood (OW) and CW. Results suggested that the inner juvenile wood of OW rings might contain either a galactose-rich galactoglucomannan or a β-1,4-galactan. Consideration of all 95 samples showed that although the contents of galactose, lignin, glucose, and mannose were linearly and strongly correlated with one another, their relationship with xylose and arabinose content was non-linear.

scholarly journals The Cytoskeleton and Its Role in Determining Cellulose Microfibril Angle in Secondary Cell Walls of Woody Tree Species

Plants ◽  
2020 ◽  
Vol 9 (1) ◽  
pp. 90
Author(s):  
Tobias ◽  
Spokevicius ◽  
McFarlane ◽  
Bossinger
Keyword(s):  
Molecular Mechanisms ◽  
Cellulose Microfibril ◽  
Wood Quality ◽  
Microfibril Angle ◽  
Wood Formation ◽  
Cellulose Microfibrils ◽  
Reaction Wood ◽  
Molecular Control ◽  
Cellulose Deposition ◽  
Cellulose Microfibril Angle

Recent advances in our understanding of the molecular control of secondary cell wall (SCW) formation have shed light on molecular mechanisms that underpin domestication traits related to wood formation. One such trait is the cellulose microfibril angle (MFA), an important wood quality determinant that varies along tree developmental phases and in response to gravitational stimulus. The cytoskeleton, mainly composed of microtubules and actin filaments, collectively contribute to plant growth and development by participating in several cellular processes, including cellulose deposition. Studies in Arabidopsis have significantly aided our understanding of the roles of microtubules in xylem cell development during which correct SCW deposition and patterning are essential to provide structural support and allow for water transport. In contrast, studies relating to SCW formation in xylary elements performed in woody trees remain elusive. In combination, the data reviewed here suggest that the cytoskeleton plays important roles in determining the exact sites of cellulose deposition, overall SCW patterning and more specifically, the alignment and orientation of cellulose microfibrils. By relating the reviewed evidence to the process of wood formation, we present a model of microtubule participation in determining MFA in woody trees forming reaction wood (RW).

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