Changes in Nanostructure of Wood Cell Wall during Deformation

2008 ◽  
Vol 599 ◽  
pp. 126-136 ◽  
Author(s):  
Marko Peura ◽  
Seppo Andersson ◽  
Ari Salmi ◽  
Timo Karppinen ◽  
Mika Torkkeli ◽  
...  

The excellent mechanical properties of wood arise from its cellular and cell wall structure. X-ray scattering, ultrasound, and mechanical testing is combined to study the effects of strain on crystalline cellulose in wood. Results for dry and re-moistened softwood samples are reviewed and new results are presented for native, never-dried samples of Silver birch. When softwood is stretched parallel to the cell axis, the mean microfibril angle diminishes significantly in compression wood, but only slightly in clear wood. The cellulose chains in the crystallites elongate and their distance diminishes. In the never-dried Silver birch samples, axial strain caused the mode of the microfibril angle distribution to slightly decrease from the initial value of 14 degrees to 12 degrees. Unlike in softwood, in never-dried birch crystalline cellulose showed auxetic tensile behaviour. The distance of the chains increased and the X-ray Poisson ratio νca was negative, -0.3 ± 0.2. Dehydration of never-dried Silver birch caused no difference to the microfibril angle distribution.

IAWA Journal ◽  
2010 ◽  
Vol 31 (1) ◽  
pp. 3-11 ◽  
Author(s):  
Clemens M. Altaner ◽  
Michael C. Jarvis ◽  
Jack B. Fisher ◽  
Thomas E. Marler

The molecular structure of tracheid walls of an inclined eccentrically grown stem of Cycas micronesica K.D. Hill did not differ between the upper and lower side. The absence the typical molecular features of compression wood tracheids, i.e. an increased galactose and lignin content as well as an increased microfibril angle, indicated that cycads do not have the ability to form even very mild forms of compression wood, which lacks anatomical features commonly observed in compression wood. Analysis of the sugar monomers in Cycas micronesica tracheids did reveal a rather unique composition of the non-cellulosic polysaccharides for a gymnosperm. The low mannose and high xylose content resembled a cell wall matrix common in angiosperms. The crystalline cellulose structure in Cycas micronesica tracheids closely resembled those of secondary cell walls in Picea sitchensis (Bong.) Carr. tracheids. However, the spacing between the sheets of cellulose chains was wider and the cellulose fibrils appeared to form larger aggregates than in Sitka spruce tracheids.


Holzforschung ◽  
2005 ◽  
Vol 59 (4) ◽  
pp. 422-427 ◽  
Author(s):  
Seppo Andersson ◽  
Ritva Serimaa ◽  
Tiina Väänänen ◽  
Timo Paakkari ◽  
Saila Jämsä ◽  
...  

AbstractWood is thermally modified by heating and steaming in order to change its properties, e.g., to improve the biological resistance and to increase the hardness of wood. The structure of thermally modified Scots pine (Pinus sylvestris) was studied using wide-angle, small-angle and ultra-small-angle X-ray scattering methods. Modification temperatures varied from 100 to 240°C. No marked changes in the microfibril angle distribution were observed. The mass fraction of crystalline cellulose in wood (the crystallinity of wood) and the size of cellulose crystallites increased above 150°C. After modification at 230°C for 4 h the thickness of the cellulose crystallites increased from 3.1 to 3.4 nm. Thermal modification had no effect on the orientation of the voids, but an increase in the porosity of the cell wall was observed. The distance between cellulose crystallites was approximately 4.7 nm in hydrated wood and a decrease in order between microfibrils was observed at 160–200°C.


The cell-wall structure of the red alga Rhodymenia palmata has been examined by the methods of X -ray diffraction analysis and electron microscopy, including ultra-thin sectioning. The cell wall is shown to consist of numerous lamellae each of which is made up of unoriented, crystalline microfibrils embedded in an amorphous matrix of other cell-wall constituents. The material can be stretched reversibly up to 100% when wet, and the stretching induces orientation of the microfibrils. The ‘∝ cellulose' fraction, which accounts for only 2 to 7 % of the original dry weight, was isolated chemically and was analyzed by means of hydrolysis and paper chromatographic separation of the resulting sugars, and it was found to be composed of approximately equal quantities of glucose and xylose residues. Chemical treatment of the cell wall was found to cause considerable variations in the X -ray diagrams, which are discussed. It is concluded that the microfibrils contain both glucose and xylose residues in approximately equal proportions and that chemical treatment in this case causes changes in crystallinity of the structural component of the wall. The importance of these findings for the meaning of the term cellulose is discussed. The X -ray diagram of older fronds was found to be complicated by the occurrence of extra rings due to the presence of floridean starch, and the highly elastic properties of the thallus enabled the diagrams of the starch and the cell wall to be separated.


1952 ◽  
Vol 5 (2) ◽  
pp. 223 ◽  
Author(s):  
AB Wardrop ◽  
HE Dadswell

The fine structure of the cell wall of both ray and vertical parenchyma has been investigated. In all species examined secondary thickening had occurred. In the primary cell wall the micellar orientation was approximately trans"erse to the longitudiJ)aI cell axis. Using optical and X-ray methods the secondary cell wall was shown to possess a helical micellar organization, the micelles being inclined between 30� and 60� to the longitudinal cell axis.


A general survey of cell-wall structure in the red algae has been carried out using the methods of X -ray diffraction analysis and electron microscopy. The fifteen species all show a similar wall structure consisting of numerous lamellae each of which is made up of random micro-fibrils embedded in an amorphous matrix. The X -ray diagrams obtained from several species are complicated by the existence of crystalline floridean starch, but nevertheless reveal the absence of cellulose I.


IAWA Journal ◽  
2008 ◽  
Vol 29 (4) ◽  
pp. 345-386 ◽  
Author(s):  
Lloyd Donaldson

Microfibril angle (MFA) is perhaps the easiest ultrastructural variable to measure for wood cell walls, and certainly the only such variable that has been measured on a large scale. Because cellulose is crystalline, the MFA of the S2 layer can be measured by X-ray diffraction. Automated X-ray scanning devices such as SilviScan have produced large datasets for a range of timber species using increment core samples. In conifers, microfibril angles are large in the juvenile wood and small in the mature wood. MFA is larger at the base of the tree for a given ring number from the pith, and decreases with height, increasing slightly at the top tree. In hardwoods, similar patterns occur, but with much less variation and much smaller microfibril angles in juvenile wood. MFA has significant heritability, but is also influenced by environmental factors as shown by its increased values in compression wood, decreased values in tension wood and, often, increased values following nutrient or water supplementation. Adjacent individual tracheids can show moderate differences in MFA that may be related to tracheid length, but not to lumen diameter or cell wall thickness. While there has been strong interest in the MFA of the S2 layer, which dominates the axial stiffness properties of tracheids and fibres, there has been little attention given to the microfibril angles of S1 and S3 layers, which may influence collapse resistance and other lateral properties. Such investigations have been limited by the much greater difficulty of measuring angles for these wall layers. MFA, in combination with basic density, shows a strong relationship to longitudinal modulus of elasticity, and to longitudinal shrinkage, which are the main reasons for interest in this cell wall property in conifers. In hardwoods, MFA is of more interest in relation to growth stress and shrinkage behaviour.


2005 ◽  
Vol 38 (3) ◽  
pp. 505-511 ◽  
Author(s):  
Kenneth M. Entwistle ◽  
Stephen J. Eichhorn ◽  
Namasivayam Navaranjan

A method is presented for the measurement, using small-angle X-ray scattering (SAXS), of the microfibril angle and the associated standard deviation for the cellulose microfibrils in the S2 layer of the cell walls of softwood specimens. The length and orientation of over 1000 cell walls in the irradiated volume of the specimen are measured using quantitative image analysis. From these data are calculated the azimuthal variation of the scattered intensity. The calculated values are compared with the measured values. The undetermined parameters in the analysis are the microfibril angle (M) and the standard deviation (σΦ) of the intensity distribution arising from the wandering of the fibril orientation about the mean value. The two parameters are varied to give the best fit between the calculated and the measured values. Six separate pairs of values are determined for six different values of the angle of incidence of the X-ray beam relative to the normal to the radial direction in the specimen. The results show good agreement. The azimuthal distribution of scattered intensity for the real cell-wall structure is compared with that calculated for an assembly of rectangular cells with the same ratio of transverse to radial cell-wall lengths. Despite the existence of marked differences in the intensity distributions around the zero azimuth angle, the position of the extreme flanks of the distribution is very close for the real and the rectangular cells. This means that useful values of the microfibril angle can be obtained from the curve for the real cells using the Meylan parameter T derived by drawing tangents to the flanks of the intensity distribution and using M = kT. The value of k is M/(M + 2σΦ). Since both of these parameters are determined in the work now described, k is also determined. It is also demonstrated that for β = 45° (where β is the angle between the plane face of the wood specimens and the radial direction) the peaks in the azimuthal intensity distribution for the real and the rectangular cells coincide. If this peak position is Φ45, then the microfibril angle can be determined from the relation M = tan−1(tanΦ45/cos45°), which is precise for rectangular cells.


2015 ◽  
Vol 22 (2) ◽  
pp. 267-272 ◽  
Author(s):  
Selina Storm ◽  
Malte Ogurreck ◽  
Daniel Laipple ◽  
Christina Krywka ◽  
Manfred Burghammer ◽  
...  

The high flux density encountered in scanning X-ray nanodiffraction experiments can lead to severe radiation damage to biological samples. However, this technique is a suitable tool for investigating samples to high spatial resolution. The layered cell wall structure of softwood tracheids is an interesting system which has been extensively studied using this method. The tracheid cell has a complex geometry, which requires the sample to be prepared by cutting it perpendicularly to the cell wall axis. Focused ion beam (FIB) milling in combination with scanning electron microscopy allows precise alignment and cutting without splintering. Here, results of a scanning X-ray diffraction experiment performed on a biological sample prepared with a focused ion beam of gallium atoms are reported for the first time. It is shown that samples prepared and measured in this way suffer from the incorporation of gallium atoms up to a surprisingly large depth of 1 µm.


Holzforschung ◽  
2013 ◽  
Vol 67 (3) ◽  
pp. 315-323 ◽  
Author(s):  
Keisuke Toba ◽  
Hiroyuki Yamamoto ◽  
Masato Yoshida

Abstract The present study is aimed at the detection of mechanical stresses generated in the cellulose microfibril (CMF) crystals situated in the secondary wall (S2) of living cells. Green wood specimens were boiled in water to release the internal stress in the CMF by the hygrothermal softening of the lignin-hemicellulose matrix (MT). Thereafter, the changes in d200 and d004 lattice spacings of crystalline cellulose were observed in boiled and nonboiled samples by wide-angle X-ray diffraction. The d200 lattice spacing increased, whereas d004 lattice spacing decreased. The results show that a mechanical stress still remained in the CMF and the MT region in the green cell wall, even after releasing the macroscopic surface growth stresses by removal of the wood block from the living stem. The interpretation is that CMF generates tensile stress in the longitudinal direction, and surrounding MT substances generate compressive stress in the living cell wall, which compresses the CMF in the lateral direction. The results confirm the “unified hypothesis” for explaining the mechanism of growth stress generation.


2010 ◽  
Vol 173 ◽  
pp. 72-77
Author(s):  
Tabet A. Tamer ◽  
Aziz Abdul Haji Fauziah ◽  
Radiman Shahidan

Partially crystalline cellulose microfibrils are wound helically around the longitudinal axis of the wood cell. A method is presented for the measurement, using small-angle X-ray scattering (SAXS), of the microfibril angle, (MFA) and the associated standard deviation for the cellulose microfibrils in the S2 layer of the cell walls of Acacia mangium wood. The length and orientation of the microfibrils of the cell walls in the irradiated volume of the thin samples are measured using SAXS and scanning electron microscope, (SEM). The undetermined parameters in the analysis are the MFA, (M) and the standard deviation (σФ) of the intensity distribution arising from the wandering of the fibril orientation about the mean value. Nine separate pairs of values are determined for nine different values of the angle of the incidence of the X-ray beam relative to the normal to the radial direction in the sample. The results show good agreement. The curve distribution of scattered intensity for the real cell wall structure is compared with that calculated with that assembly of rectangular cells with the same ratio of transverse to radial cell wall length. It is demonstrated that for β = 45°, the peaks in the curve intensity distribution for the real and the rectangular cells coincide. If this peak position is Ф45, Then the MFA can be determined from the relation M = tan-1 (tan Ф45 / cos 45°), which is precise for rectangular cells.


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