Anatomy and lignin distribution of reaction wood in two Magnolia species

Summary Anatomical features of reaction wood formed in two Magnolia species, M. obovata Thunb. and M. kobus DC. which are considered to be among the primitive angiosperms, were observed. In addition, the distribution of guaiacyl and syringyl units of lignins in the cell walls of normal and reaction wood was examined using ultraviolet (UV)- and visible light (VL)- microspectrophotometry coupled with the Wiesner and M?ule reactions. The two Magnolia species formed a tension-like reaction wood without possessing the typical gelatinous layer (G-layer) on the upper side of the inclined stem or branch, in which a radial growth promotion occurred. Compared with the normal wood, the reaction wood had the following anatomical features: (1) the secondary walls of fiber tracheids lacked the S₃ layer, (2) the innermost layer of fiber-tracheid walls showed a small microfibril angle, a fact being similar to the orientation of the microfibril angle of the G-layer in tension wood, and (3) the amounts of lignin decreased in the cell walls of fiber tracheids, especially with great decrease in proportion of guaiacyl units in lignins. In addition, VL-microspectrophotometry coupled with the Wiesner and M?ule reactions adopted in the present study showed potential to estimate the lignin contents in the cell walls and the proportion of guaiacyl and syringyl units in lignins. Received: 15 July 1998     

Reaction wood anatomy and lignin distribution in <Emphasis Type="Italic">Gnetum gnemon</Emphasis> branches

This study investigated the anatomical and chemical characteristics of the reaction wood of a gymnpsperm species, Gnetum gnemon, and discussed on contributing factor for the type of reaction wood in this species. Cell morphology, microfibril angle (MFA) of the S₂ layer and lignin distribution in secondary walls of tracheary elements, and lignin content were examined on three branches. Observations included no G-layer formation, significant decreases in vessel frequency, and altered MFA, and visible-light absorbance after lignin colour reactions in tracheid and fiber tracheid walls on the upper side in almost all samples. These results suggest that reaction wood in G. gnemon was similar to that in ‘tension-wood-like-reaction wood’ in angiosperms. On the other hand, reaction wood showed decrease in the lignin concentration in the fiber tracheid walls compared to the tracheid walls. In addition, the lignin in the tracheid and fiber tracheid walls was originally rich in syringyl units, suggesting that changes in the anatomical and chemical characteristics of secondary xylem due to reaction wood formation might relate to the ratio of the syringyl to guaiacyl units in lignin in the cell walls which function for mechanical support.     

Condensation of lignin during heating of wood

Summary The structural change of lignin during heating of wood was investigated quantitatively by a method combining nucleus exchange and nitrobenzene oxidation. Lignin modification during heating was mainly a diphenylmethane type condensation. About 40 and 75% of noncondensed units in protolignins were converted to diphenylmethane type units by heating of dry and wet wood meals up to 220 °C, respectively. On the other hand, during heating of modified lignin (dioxane lignin) various types of modifications in addition to diphenylmethane type condensation occurred. Lignin modification via the diphenylmethane type condensation was proposed as a new route for its utilization.     

The application of ultraviolet microscopy to the distribution of lignin in wood description and validity of the technique

A method has been developed for the determination of lignin distribution in the wood cell wall by ultraviolet microscopy. The method incorporates some important advances on previus applications of UV microscopy to the study of lignin distribution. Ultrathin cross-sections of wood are obtained by the sample preparation and sectioning techniques of electron microscopy. The specimens are examined in monochromatic ultraviolet light using quartz reflection optics. The microscope image is photographically recorded and the negative is subsequently subjected to densitometric analysis. Each stage of the analytical procedure has been critically assessed to determine its validity and limitations. The method is ideally suited to the study of the removal of lignin from the wood cell wall during cooking and possesses other important applications in wood technology.     

Chemical composition of wood and its connection with wood anatomy in Betula pubescens

ABSTRACT

The connection between chemical composition and anatomy of wood is poorly studied. Abiotic and biotic growth conditions affect the synthesis of structural compounds and the anatomy of wood at the same time as they affect growth. In this study, the wood chemical composition, and connections between wood chemistry and anatomy were studied in downy birch (Betula pubescens Ehrh.) grown on four growth sites possessing mineral and peat soils. Lignin, holocellulose, extractive and ash contents, and effective heating value were analysed and compared with libriform fibre double wall thickness and lumen diameter, vessel size and number, and ray and axial parenchyma numbers. Measured lignin and extractive contents were exceptionally high and holocellulose content low. Correlations between wood anatomy and effective heating value were partly different on different soils. Correlations between holocellulose and anatomy were the opposite of the correlations between other chemical compounds and anatomy. The significant correlations between chemical composition and anatomy were commonly opposite in trees grown on peat than in trees grown on mineral soil. Particularly, vessel characteristics and fibre wall thickness correlated significantly with wood chemistry in trees grown on mineral soil. The connections between wood anatomy and chemistry proved to be soil-dependent.     

Analysis of birch wood lignin by oxidative degradation

Summary The lignin in birch wood has been subjected to analysis by oxidative degradation after milling of wood samples in a Wiley mill and in a rotatory ball mill. For comparison milled wood lignin from birch has also been analysed. It was found that the type of milling affects both the total yield of degradation acids and their relative frequency of occurrence. These findings are interpreted as being due to a cleavage of aryl ether linkages during the milling procedure. The oxidative degradation analysis gave rise to a total of 30 different acids which were tentatively identified on the basis of their mass spectral fragmentation patterns. Among the acids some were found having either a phenoxyacetic acid or a phenoxypropionic acid structure. The possible origin of these is discussed. A grant to one of us (R.A.N.) from Jacob Wallenbergs Forskningsstiftelse is gratefully acknowledged. Thanks are due to Kristina Gustafsson for skilful experimental assistance     

Chemical structural elucidation of total lignins in woods I: fractionation of the lignin in residual wood meal after extraction of milled wood lignin

Residual wood meal after extraction of milled wood lignin (WMEM) ofEucalyptus globulus was extracted with alkali and LiCl/N,N-Dimethylacetamide (DMAc). These agents dissolve mainly hemicellulose and cellulose, respectively. The extractability of WMEM in alkali solutions was influenced by the degree of swelling of the cellulose. Under good swelling conditions, considerable amounts of cellulose and lignin were extracted with the hemicellulose. Maximum extractability was about 60% of the WMEM under optimum conditions (3 M or 5 M LiOH or 3M NaOH solution). Some portion of cellulose was extracted with LiCl/DMAc at room temperature. Thus, lignin inE. globulus WMEM was divided into three fractions: hemicellulose-lignin fraction, cellulose-lignin fraction, and insoluble-lignin fraction.Part of this work was presented at the 49th annual meeting of the Japan Wood Research Society, Tokyo, April 1999; and at the 50th annual meeting of the Japan Wood Research Society, Kyoto, April 2000     

Notes on the spectrophotometric determination of lignin in wood samples

A method developed by Johnson et al. whereby lignin is determined spectrophotometrically was found to yield lignin values which showed variations of up to 10%. The presence of water in, and the freshness of the digestive mixture together with the grade of acetic acid used and the time between dilution and measurement of absorbance were found to affect the reproducibility of the method.     

Fracturing of wood under superimposed tension and torsion

A testing method using circumferentially notched round bars for investigating mixed mode behaviour under loading in tension and torsion is applied to wood. The applicability of the method to anisotropic materials is investigated for two types of wood, beech and spruce, considering the longitudinal and radial orientation with respect to the stem axis of the tree. The strong anisotropy of wood requires different evaluation procedures for radial and longitudinal sample orientation. The K-concept of linear elastic fracture mechanics (LEFM) and concepts of non-linear elastic fracture mechanics (NLEFM) were used for the evaluation of radial and longitudinal samples, respectively. Differences between the investigated wood types under radial orientation, in their durability to withstand torsional loads, could be observed by examining ratios of the values of the fracture toughness in mode III against mode I. Micrographs of the fracture surfaces support the assumption that the higher amount of wood rays in beech is responsible for the higher toughness under torsion. In case of longitudinal specimen geometry it was found that at very high levels of torsional deformation beech and spruce reach similar values in their specific fracture energy in mode I.     

Lignification and peroxidase in tension wood ofEucalyptus viminalis seedlings

Seedlings ofEucalyptus viminalis were grown for 50 days with their stems bent so tension wood would form. Every 10 days the lignin content, monomeric composition, and peroxidase activity in the tension wood were compared with those in the lower side (opposite wood) and in vertically grown controls. The lignin content in the developing tension wood started to decrease after 10 days of bending and kept decreasing for 50 days, whereas those in control plants and opposite wood remained almost unchanged. The yields of syringaldehyde from tension wood by nitrobenzene oxidation increased, and consequently the syringyl/ guaiacyl ratio of the lignin was higher in tension wood than in opposite wood and control plants. The peroxidase ionically bound to the cell walls (IPO) catalyzed oxidation of guaiacol and syringaldazine. The syringaldazineoxidizing activity of IPO from tension wood increased, whereas the activities of IPO from opposite wood and control plants did not show any marked change. In tension wood the increase in syringaldazine-oxidizing activity of IPO was consistent with an increase in the syringaldehyde yield. This suggests that IPO contributes to syringyl lignin deposition as other enzymes involved in the monolignol biosynthesis do in tension wood formation.This study was presented at the 50th Annual Meeting of the Japan Wood Research Society, Kyoto, April 2000     

A comparison of static bending,compression and tension parallel to grain and toughness properties of compression wood and normal wood of a giant sequoia

Summary Compression wood (CW) of the giant sequoia studied had higher values than normal wood (NW) in crushing strength and ultimate stress in tension parallel to grain, in toughness, in modulus of rupture, and in work to maximum load and total work in static bending. In the green condition CW had higher values than NW in stress at the proportional limit and work to the proportional limit, and about the same modulus of elasticity in static bending. In the dry condition CW was about equivalent to NW in work to the proportional limit, but was slightly weaker in stress at proportional limit and modulus of elasticity in static bending. The compression wood of this giant sequoia, even though formed when the tree was suppressed and having relatively narrow rings, can therefore be said to be essentially equivalent to normal wood so far as the mechanical properties tested in this study are concerned.Given at FPRS meeting in Dallas, Texas, June 1972     

Comparative study on penetration characteristics of modern wood coatings

Summary The penetration characteristics of five modern wood coatings (three waterborne, one high solid and one solvent borne) into pine sapwood, spruce and dark red meranti have been systematically compared. The degree of coating penetration is mainly determined by the ability of the coating to flow into wood capillaries. Binder type, pigmentation, solid matter content and drying speed appeared to influence this ability. In softwoods the following different coating penetration routes are observed: the flow into open ends of longitudinal early-and latewood tracheids, the flow into ray cells and the transport from rays through the cross-field into longitudinal tracheids adjacent to rays. The possibility for the coating to follow the latter route is strongly influenced by the existing type of cross field pitting and to a lesser degree by the pigmentation of the paint. Clear differences between pine and spruce have been found with respect to the flow into ray parenchym and ray tracheids. The flow into open ends of longitudinal tracheids is strongly influenced by the grain angle of tracheids. Penetration into dark red meranti is mainly limited to vessels and rays. Tylose membranes can prevent the complete filling of vessels. The impact on penetration of the removal of extractives and of sanding of the surface has also been studied but appears to be of only minor importance.The authors want to thank Akzo Nobel Coatings, DSM Resins and Sigma Coatings for their technical support. This research was financed by the the Dutch Innovative Research Program on Coatings under contract number IVE 93-812. Authors also thank the University of Hamburg (BFH) for the use of their electron microscopy facilities.     

Micromechanics of wood subjected to axial tension

Summary The behaviour of a small group of wood fibers of Sitka spruce during tensile loading is investigated. The load-extension curves for both early and late wood fibers consist of three distinct segments. The first segment is almost a straight line, at some stage of loading a yield point is observed. Beyond this point the specimen becomes less stiff and undergoes a large, mainly irreversible deformation. As the load is increased further, the curve exhibits the third segment showed by a significant change in slope. These curves look different from those obtained on thick specimens. In this respect, the behaviour of a thin wood specimen subjected to cyclic type tensile loading along its longitudinal direction is also illustrated. Based on wood microstructure, a model is presented to interpret the evolution of the Young's modulus of a wood fiber during tensile loading. The model considers wood as an assembly of cylindrical fibers pasted together in a longitudinal direction. We have assumed the cell wall to comprise only an S₂ layer made of a composite material consisting of a lignin and hemicellulose matrix reinforced by helical microfibrils along the fiber. Furthermore, it is assumed that the microfibril angle a in the S₂ layer is not uniform along the fiber axis and matrix degradation occurs in the zones where the microfibril angles are bigger. The validity of this assumption is verified by using holographic interferometry to visualize the displacement field of the specimen's surface under tension.The work reported in this paper is supported by a grant from the Swiss National Science Foundation, and its support is gratefully acknowledged     

Studies on compression wood—part VII: Distribution of lignin in normal and compression wood of tamarack [Larix laricina (Du Roi) K. Koch]

Summary The distribution of lignin has been studied in tracheids and ray cells of normal and compression wood of tamarack [Larix laricina (Du Roi) K. Koch]. The three layers in the secondary wall of normal wood tracheids are lignified to approximately the same extent, and previous evidence that the S ₃ layer should contain a higher proportion of lignin than the other regions has not been confirmed. The lignin follows closely the orientation of the cellulose microfibrils in all three layers. Compared to the tracheids, the ray cells contain a denser network of lignin in their secondary wall.Only a small proportion of the total lignin in compression wood tracheids is present in the compound middle lamella. The thick S ₁ layer is only slightly lignified; the orientation of the lignin in this region is that of the transversely oriented, lamellated microfibrils. The outer portion of S ₂ consists largely of lignin but also contains lamellae of cellulose microfibrils which probably have the same helical orientation as the microfibrils in the inner part of S ₂. The latter region, which contains the helical cavities, consists of lamellae of cellulose microfibrils which are uniformly encrusted with lignin. The ray cells in compression wood appear to be lignified to the same extent as in normal wood. Transverse sections of the cells reveal a lateral orientation of the lignin. The orientation of the cellulose microfibrils in the S ₂ layer of the first-formed springwood tracheids of compression wood is the same as in the cells which are formed later. It is suggested that for ease of reference, the outer, lignin-rich layer in compression wood tracheids be referred to as the S ₂(L) layer.

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Zusammenfassung Im Druckholz und im normalen Holz von Tamarack (Larix laricina (Du Roi) K. Koch) wurde die Verteilung des Lignins in Tracheiden und Markstrahlzellen untersucht. Die drei Schichten der Sekundärwand in den Tracheiden normalen Holzes werden in nahezu demselben Umfange lignifiziert. Frühere Feststellungen, daß die S ₃-Schicht einen höheren Ligningehalt erreicht als andere Zellwandbereiche, konnten also nicht bestätigt werden. Das Lignin folgt sehr genau der Orientierung der Cellulose-Mikrofibrillen aller drei Schichten. Im Vergleich zu den Tracheiden erfahren die Sekundärwände der Markstrahlzellen eine stärkere Ligninauskleidung.Nur ein geringer Prozentsatz des gesamten Lignins der Druckholztracheiden befindet sich in der Mittellamelle. Die dicke S ₁-Schicht ist nur wenig lignifiziert. Die Orientierung des Lignins in diesem Bereich entspricht den transversal orientierten, lamellierten Mikrofibrillen. Der äußere Teil der S ₂-Schicht enthält sehr viel Lignin, daneben aber auch Lamellen von Cellulose-Mikrofibrillen, die wahrscheinlich dieselbe spiralige Orientierung besitzen wie die Mikrofibrillen des inneren Teiles der S ₂-Schicht. Der letzterwähnte Bereich, der spiralige Kavitäten enthält, weist Lamellen von Cellulose-Mikrofibrillen auf, in welche gleichmäßig Lignin eingelagert ist. Die Markstrahlzellen des Druckholzes erscheinen ebenso stark lignifiziert wie die Markstrahlzellen des Normalholzes. Querschnitte durch diese Zellen lassen die laterale Orientierung des Lignins erkennen. Die Orientierung der Cellulose-Mikrofibrillen in der S ₂-Schicht der zuerst gebildeten Frühholztracheiden des Druckholzes ist dieselbe wie in jenen Zellen, die später ausgeformt werden. Es wird vorgeschlagen, daß zur eindeutigeren Kennzeichnung die äußere ligninreiche Schicht der Druckholztracheiden als S ₂(L)-Schicht bezeichnet wird.

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The authors wish to express their gratitude to Messrs. A. K. Bentum, D. C. Jones, and B. W. Simson for technical assistance. They are also thankful to Dr. D. A. I. Goring, McGill University, Montreal, Canada, for valuable discussions and for making available to them important, unpublished information. This investigation was supported by the United States Department of Agriculture, Forest Service, through Forest Service Research Grant No. 1, which is hereby gratefully acknowledged.     

杞柳初植密度对比和品种对比试验效应的研究

设置固定试验地,随机区组排列,经裂方差分析,检验初植密度间、品种间差异性情况,再进行单项对比统计分析,筛选出0.3m×0.1m为最佳初植密度;莒稀为优良品种,莒红为良好品种。从而为杞柳造林解决了初植密度和选定品种两大关键性技术。     

A comparative study of bamboo (Phyllostachys incarnata Wen) milled wood lignin and the successively alkali-fractionated lignins

The chemical composition and structural characterization of the milled wood lignin (MWL) and successively alkali-fractionated lignins from bamboo (Phyllostachys incarnata Wen) were comparatively investigated using Fourier transform infrared (FT-IR), gel permeation chromatography (GPC), 2D heteronuclear single quantum coherence (HSQC), and alkaline nitrobenzene oxidation. It was found that bamboo MWL (S/G?~?1.3) is of SGH type with a considerable amount of esterified p-coumaric acid and etherified ferulic acid. Molecular weights of 3,320?g/mol for MWL and varying weights between 970 and 3,430?g/mol for the alkali-fractionated lignins were obtained. Moreover, 2D HSQC of MWL showed a predominance of β-O-4′ linkages (65%), followed by β–β′ linkages (15%) and lower amounts of β-5′ (7%), β-1′ (4%), 5–5′ (4%) linkages, and cinnamyl acetate end groups (5%). In addition, a percentage (~45%) of the lignin side chain was found to be acylated at the γ-carbon. However, the alkali-fractionated lignins lost parts of these linkages during successive alkaline fractionation processes.     

An improved acetyl bromide procedure for determining lignin in woods and wood pulps

Summary The acetyl bromide procedure for spectrophotometrically determining lignin in wood and wood pulp samples has been modified by adding perchloric acid to the digestion medium. This enables faster dissolution of the materials and the use of coarser samples. Subsequent treatment of the digestion mixtures with larger amounts of sodium hydroxide than those used in the conventional procedure has obviated the need for treatment with hydroxylamine. The modified method has been successfully applied to woodmeals and pulps from Pinus radiata and Eucalyptus species.     

Active enzymes in wood chips and their action on lignin during outside storage

Summary Wood storage has a great influence on the efficiency of a pulpmill. Great problems arise when using outside chip storage, especially in processing hardwoods, in that hardwood chips discolorate after a few days, which is not the case with spruce chips. The causes of such a different behavior of beech and spruce were investigated. It could be shown that a sufficient amount of active peroxidase, malate dehydrogenase and oxygen are prerequisites for the formation of phenoxy radicals within the beech lignin. This causes formation of new chromophores that discolorize the wood. Sufficient amounts of such enzymes are present in beech wood after harvesting.