The nature of reaction wood. IV. Variations in cell wall organization of tension wood fibres

1955 ◽  
Vol 3 (2) ◽  
pp. 177 ◽  
Author(s):  
AB Wardrop ◽  
HE Dadswell
Keyword(s):  
Cell Wall ◽  
Tension Wood ◽  
Secondary Wall ◽  
Degree Of Crystallinity ◽  
Tropical Species ◽  
Normal Wood ◽  
Reaction Wood ◽  
Gelatinous Layer ◽  
Wood Fibres ◽  
Cell Wall Organization

The cell wall organization, the cell wall texture, and the degree of lignification of tension wood fibres have been investigated in a wide variety of temperate and tropical species. Following earlier work describing the cell wall structure of tension wood fibres, two additional types of cell wall organization have been observed. In one of these, the inner thick "gelatinous" layer which is typical of tension wood fibres exists in addition to the normal three-layered structure of the secondary wall; in the other only the outer layer of the secondary wall and the thick gelatinous layer are present. In all the tension wood examined the micellar orientation in the inner gelatinous layer has been shown to be nearly axial and the cellulose of this layer found to be in a highly crystalline state. A general argument is presented as to the meaning of differences in the degree, of crystallinity of cellulose. The high degree of crystallinity of cellulose in tension wood as compared with normal wood is attributed to a greater degree of lateral order in the crystalline regions of tension wood, whereas the paracrystalline phase is similar in both cases. The degree of lignification in tension wood fibres has been shown to be extremely variable. However, where the degree of tension wood development is marked as revealed by the thickness of the gelatinous layer the lack of lignification is also most marked. Severity of tension wood formation and lack of lignification have also been correlated with the incidence of irreversible collapse in tension wood. Such collapse can occur even when no whole fibres are present, e.g. in thin cross sections. Microscopic examination of collapsed samples of tension wood has led to the conclusion that the appearance of collapse in specimens containing tendon wood can often be attributed in part to excessive shrinkage associated with the development of fissures between cells, although true collapse does also occur. Possible explanations of the irreversible shrinkage and collapse of tension wood fibres are advanced.

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.

scholarly journals SHRINKAGE OF THE GELATINOUS LAYER OF POPLAR AND BEECH TENSION WOOD

IAWA Journal ◽  
2001 ◽  
Vol 22 (2) ◽  
pp. 121-131 ◽  
Author(s):  
Bruno Clair ◽  
Bernard Thibaut
Keyword(s):  
Cell Wall ◽  
Tension Wood ◽  
Normal Wood ◽  
Longitudinal Shrinkage ◽  
Sem Observation ◽  
Gelatinous Layer ◽  
Dry Conditions

Macroscopic longitudinal shrinkage of beech and poplar tension wood is higher than in normal wood. This shrinkage is the result of mechanical interactions of cell wall layers. SEM observation of cut, dried surfaces showed that longitudinal shrinkage is much greater in the gelatinous layer than in other layers. AFM topographic images of the same cells, both in water and in air-dry conditions, confirm this result. Measurements on sections indicate around 4.7% longitudinal shrinkage for the G layer.

scholarly journals Diversity in the organisation and lignification of tension wood fibre walls – A review

IAWA Journal ◽  
2017 ◽  
Vol 38 (2) ◽  
pp. 245-265 ◽  
Author(s):  
Barbara Ghislain ◽  
Bruno Clair
Keyword(s):  
Cell Wall ◽  
Tensile Stress ◽  
Cell Walls ◽  
Tension Wood ◽  
Anatomical Structure ◽  
Wood Fibre ◽  
Tropical Species ◽  
Wood Fibres ◽  
Staining Techniques

Tension wood, a tissue developed by angiosperm trees to actively recover their verticality, has long been defined by the presence of an unlignified cellulosic inner layer in the cell wall of fibres, called the G-layer. Although it was known that some species have no G-layer, the definition was appropriate since it enabled easy detection of tension wood zones using various staining techniques for either cellulose or lignin. For several years now, irrespective of its anatomical structure, tension wood has been defined by its high mechanical internal tensile stress. This definition enables screening of the diversity of cell walls in tension wood fibres. Recent results obtained in tropical species with tension wood with a delay in the lignification of the G-layer opened our eyes to the effective presence of large amounts of lignin in the G-layer of some species. This led us to review older literature mentioning the presence of lignin deposits in the G-layer and give them credit. Advances in the knowledge of tension wood fibres allow us to reconsider some previous classifications of the diversity in the organisation of the fibre walls of the tension wood.

scholarly journals Cell wall thickening in developing tension wood of artificially bent poplar trees

IAWA Journal ◽  
2015 ◽  
Vol 36 (1) ◽  
pp. 44-57 ◽  
Author(s):  
Raoufeh Abedini ◽  
Bruno Clair ◽  
Kambiz Pourtahmasi ◽  
Françoise Laurans ◽  
Olivier Arnould
Keyword(s):  
Tensile Stress ◽  
Tension Wood ◽  
Secondary Wall ◽  
Low Cost ◽  
Growing Season ◽  
Progressive Increase ◽  
Wall Layer ◽  
Wall Thickening ◽  
Normal Wood ◽  
Wood Fibres

Trees can control their shape and resist gravity thanks to their ability to produce wood under tensile stress. This stress is known to be produced during the maturation of wood fibres but the mechanism of its generation remains unclear. This study focuses on the formation of the secondary wall in tension wood produced in artificially tilted poplar saplings. Thickness of secondary wall layer (SL) and gelatinous layer (GL) were measured from cambium to mature wood in several trees sampled at different times after tilting. Measurements on wood fibres produced before tilting show the progressive increase of secondary wall thickness during the growing season. After the tilting date, SL thickness decreased markedly from normal wood to tension wood while the total thickness increased compared to normal wood, with the development of a thick GL. However, even after GL formation, SL thickness continues to increase during the growing season. GL thickening was observed to be faster than SL thickening. The development of the unlignified GL is proposed to be a low cost, efficient strategy for a fast generation of tensile stress in broadleaved trees.

Degradation of the Gelatinous Layer in Aspen and Rubberwood by the Blue Stain Fungus Lasiodiplodia Theobromae

IAWA Journal ◽  
1997 ◽  
Vol 18 (2) ◽  
pp. 107-115 ◽  
Author(s):  
Osvaldo Encinas ◽  
Geoffrey Daniel
Keyword(s):  
Cell Wall ◽  
Cell Walls ◽  
Tension Wood ◽  
Soluble Carbohydrates ◽  
Tropical Species ◽  
Cell Wall Degradation ◽  
Lasiodiplodia Theobromae ◽  
Gelatinous Layer ◽  
Weight Losses ◽  
Blue Stain

Studies on the degradative ability of the blue stain fungus Lasiodiplodia theobromae (Pat.) Griffon ' Maublanc have shown several strains to cause significant weight losses (c. 20%) in wood of temperate and tropical species, aspen (Populus tremula) and rubberwood (Hevea brasiliensis), both species that commonly form tension wood. In addition to the consumption of soluble carbohydrates, major changes occurred in the ultrastructure of fibre cell walls, with a rapid attack of the G-layer of the gelatinous fibres. Following G-layer degradation, earlywood fibres of both species showed true cell wall degradation with pronounced erosion attack, suggesting that prior destruction of the G-layer afforded greater accessibility and ease of attack of the outer secondary cell wall layers.

ANATOMY AND CHEMICAL COMPOSITION OF LIRIODENDRON TULIPIFERA STEMS INCLINED AT DIFFERENT ANGLES

IAWA Journal ◽  
2014 ◽  
Vol 35 (4) ◽  
pp. 463-475 ◽  
Author(s):  
Tokiko Hiraiwa ◽  
Haruna Aiso ◽  
Futoshi Ishiguri ◽  
Yuya Takashima ◽  
Kazuya Iizuka ◽  
...  
Keyword(s):  
Lignin Content ◽  
Fibre Length ◽  
Liriodendron Tulipifera ◽  
Original Position ◽  
Chemical Characteristics ◽  
Normal Wood ◽  
Reaction Wood ◽  
Glucose Content ◽  
Wood Fibres ◽  
S2 Layer

The anatomical and chemical characteristics of reaction wood (RW) were investigated in Liriodendron tulipifera Linn. Stems of seedlings were artificially inclined at angles of 30 (RW-30), 50 (RW-50) and 70° (RW-70) from the vertical, and compared with normal wood (NW) from a vertical seedling stem. The smallest values for the wood fibre length and vessel number were observed in RW-50. The pit aperture angle was less than 10° in RW-30 and RW-50, in which reduced lignin content was observed in the S2 layer of the wood fibres. An increase in the glucose content and a decrease in the lignin and xylose content was observed in RW-50. The stem inclination angle affected the degree of RW development with regard to anatomical and chemical characteristics: the severest RW was observed in RW-50, followed by RW-30. RW-70 was similar in anatomical and chemical characteristics to NW, apparently because the inclination was too strong to enable recovery of its original position. In this case a vertical sprouting stem was formed to replace the inclined stem.

Microfibril Angle Distribution of Poplar Tension Wood

IAWA Journal ◽  
2012 ◽  
Vol 33 (4) ◽  
pp. 431-439 ◽  
Author(s):  
Silke Lautner ◽  
Cordt Zollfrank ◽  
Jörg Fromm
Keyword(s):  
Electron Microscopy ◽  
Tension Wood ◽  
Microfibril Angle ◽  
Cellulose Microfibrils ◽  
Normal Wood ◽  
Angle Distribution ◽  
Transmission Electron ◽  
Wood Fibres ◽  
Characteristic Features ◽  
Fibril Angle

Tension wood of poplar (Populus nigra) branches was studied by lightand electron microscopy. The characteristic features of tension wood such as wider growth rings, reduced vessel density and higher gross density were confirmed by our results. Based on a novel combination of transmission electron microscopy (TEM) imaging and image analysis, involving Fourier transformation, the orientation of cellulose microfibrils in the S2- and G-layer was determined. Within the G-layer microfibril angle (MFA) was parallel to the growth axis (0°). However, in the S2 it was 13° in tension wood fibres and 4° in normal wood fibres. With the exception of the relatively low fibril angle in the S2 of tension wood fibres (13°) the results are in good agreement with those of the literature.

scholarly journals The Nature of Reaction Wood

1948 ◽  
Vol 1 (1) ◽  
pp. 3 ◽  
Author(s):  
AB Wardrop ◽  
HE Dads Well
Keyword(s):  
Tensile Strength ◽  
Tension Wood ◽  
Reaction Wood ◽  
Fibre Structure ◽  
Longitudinal Shrinkage ◽  
High Tensile Strength ◽  
Wood Fibres

The structure of tension wood fibres is ofconsiderahle academic and practicalinterest, both in relation to considerations of the stimuli which produce them,and to studies of the influence of fibre structure on the properties of the wood asa whole. As is well known, the chief abnormal properties of tension wood lie inits unusually high longitudinal shrinkage, its high tensile strength, and its lowcompressive strength .

scholarly journals Critical review on the mechanisms of maturation stress generation in trees

2016 ◽  
Vol 13 (122) ◽  
pp. 20160550 ◽  
Author(s):  
Tancrède Alméras ◽  
Bruno Clair
Keyword(s):  
Cell Wall ◽  
Tension Wood ◽  
Compression Wood ◽  
Current Knowledge ◽  
Stress Generation ◽  
Plant Science ◽  
Reaction Wood ◽  
Wood Structure ◽  
Structure State ◽  
Cell Organization

Trees control their posture by generating asymmetric mechanical stress around the periphery of the trunk or branches. This stress is produced in wood during the maturation of the cell wall. When the need for reaction is high, it is accompanied by strong changes in cell organization and composition called reaction wood, namely compression wood in gymnosperms and tension wood in angiosperms. The process by which stress is generated in the cell wall during its formation is not yet known, and various hypothetical mechanisms have been proposed in the literature. Here we aim at discriminating between these models. First, we summarize current knowledge about reaction wood structure, state and behaviour relevant to the understanding of maturation stress generation. Then, the mechanisms proposed in the literature are listed and discussed in order to identify which can be rejected based on their inconsistency with current knowledge at the frontier between plant science and mechanical engineering.

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