On the origin of phenolic compounds in the wood rays of Abies alba

Summary The development and localization of phenolic compounds in sapwood and heartwood of Abies alba Mill. have been studied using cytological and UV-microspectrophotometric methods. The synthesis of phenolic substances was apparently initiated in vesicles developing from rough ER of the ray parenchyma cells in sapwood. They are different from the phenolic material in the pit membrane and cell wall of heartwood. There was no indication to suggest that the polyphenols in the cell wall of heartwood were either derived from or identical with the phenolic substances in the lumina of ray cells.Dedicated to Prof. Dr. H. von Pechmann on the occasion of his 70th birthdayWe are thankful to Professor Dr. Walter Liese for his support and to Mrs. R. Schultze and Miss R. Endeward for technical assistance.     

Seasonal changes in gene expression at the sapwood-heartwood transition zone of black locust (Robinia pseudoacacia) revealed by cDNA microarray analysis

Heartwood is a determining factor of wood quality and understanding the biology of heartwood may allow us to control its formation. Heartwood formation is a form of senescence that is accompanied by a variety of metabolic alterations in ray parenchyma cells at the sapwood-heartwood transition zone. Although senescence has been studied at the molecular level with respect to primary growth, the cell maturation and death events occurring during heartwood formation have been difficult to study because of their location and timing. Analysis of global gene expression patterns during the transition from sapwood to heartwood may offer a powerful means of identifying the mechanisms controlling heartwood formation. Previously, we developed cDNA microarrays carrying 2567 unigenes derived from the bark/cambium region, sapwood and transition zone of a mature black locust tree. Here, we describe the use of these microarrays to characterize seasonal changes in the expression patterns of 1873 genes from the transition zone of mature black locust trees. When samples collected in summer and fall were compared, 569 genes showed differential expression patterns: 293 genes were up-regulated (> twofold) in summer (July 5) and 276 genes were up-regulated in fall (November 27). More than 50% of the secondary and hormone metabolism-related genes on the microarrays were up-regulated in summer. Twenty-nine out of 55 genes involved in signal transduction were differentially regulated, suggesting that the ray parenchyma cells located in the innermost part of the trunk wood react to seasonal changes. We established the expression patterns of 349 novel genes (previously unknown or no-hit), of which 154 were up-regulated in summer and 195 were up-regulated in the fall.     

Variability of the chemical composition of pit membranes in bordered pits of gymnosperms

Summary The pit membranes of the bordered pits in about one hundred coniferous wood species were investigated in regard to their chemical composition, in particular to the aromatic compounds. In many species pit membranes, even in the sapwood, contain phenolic substances. In heartwood, normally lignification takes place besides the development of other polyphenols. The variability of these compounds, both in sapwood an in heartwood, may be considerable not only between species but also within the same species and even between neighbouring tracheids and pits respectively.We are thankful to Prof. Dr. W. Liese for his support and his frequent discussions.     

西加云杉木材中单宁的提取、分布及组分

研究西加云杉木材中植物单宁的提取工艺、分布和组分构成,为木材单宁变色的防治提供参考。采用有机溶剂萃取法,通过单因素试验并结合L9(34)正交试验,优化西加云杉木材中植物单宁的提取工艺;采用香草醛-紫外法测定西加云杉木材内植物单宁在径向(由髓心至树皮方向)和轴向的含量分布;通过定性鉴定试验确定单宁类型,并采用高效液相色谱法测定西加云杉木材中单宁的成分构成及其相对含量。结果表明:西加云杉木材中单宁最佳提取条件为70%(体积分数)乙醇作为提取溶剂,提取温度70℃,提取时间2 h,料液比1∶18(g∶m L),提取次数2次;单宁含量在径向上分布情况为内部心材单宁含量5.440 mg/g,近心边材转换区的心材部分单宁含量最高,平均达10.015 mg/g,心边材转换区部位单宁含量介于心材和边材之间,平均为6.363 mg/g,靠近转换区的边材部位单宁含量最低,平均为4.821 mg/g,近树皮边材含量为6.997 mg/g。西加云杉木材内5种原花青素成分含量存在明显差异,其中儿茶素含量最高,占5种原花青素总量的83.21%,原花青素B2次之。由此可知,有机溶剂法适用于西加云杉木材中单宁的提取;木材中单宁含量径向分布自髓心至树皮整体呈现出先增后减再增、心材大于边材的规律,轴向分布规律表现为下部含量略高于上部,但差异不显著;西加云杉木材内植物单宁类型为缩合单宁,儿茶素为西加云杉木材单宁的主要成分。     

Chemical composition of earlywood and latewood in Norway spruce heartwood,sapwood and transition zone wood

This study focused on the distribution of wood components along a cross section of a spruce stem. Thin samples of earlywood and latewood were analysed by special micro-scale analytical techniques. Heartwood contained significantly more lignin and less cellulose than sapwood. The total content of hemicelluloses was the same along the radial direction, but the distribution of sugar units differed. The amounts of arabinoglucuronoxylan and pectins were larger in the heartwood. The transition zone between heartwood and sapwood had a specific composition, with less lignin and lipophilic extractives than heartwood and sapwood. For earlywood and latewood, significant differences were found in the distribution of sugar units in hemicelluloses. Latewood contained clearly more galactoglucomannan than earlywood, and conversely less pectins. The lipophilic extractives were also less concentrated in the latewood.Abbreviations EW or E earlywood - LW or L latewood - HW heartwood - SW sapwood - TZ transition zone wood - A.R. annual ring - AcBr Acetyl bromide - Ara arabinose - Xyl xylose - Gal galactose - Glc glucose - Man mannose - Rha rhamnose - GlcA glucuronic acid - MGlcA 4-O-methyl-glucuronic acid - GalA galacturonic acid - o.d. oven dry     

Soft X-ray observation of water distribution in the stem ofCryptomeria japonica D. Don I: General description of water distribution

Water distribution in green stems ofCryptomeria japonica D. Don was observed by soft X-ray photography. In the sapwood, much water was present and evenly distributed. In the intermediate wood (the white zone), little water was present. The intermediate wood appeared in all cross sections of the stem and separated the heartwood from the sapwood in the intertracheid water connection. Maldistribution of water was generally observed in the heartwood, and three types of water presence were distinguishable: a “wet area” with accumulated water, a “dry area” with little water, and a “moderate moisture area” with intermediate accumulation. The distribution pattern and amount of water in the heartwood varied dramatically among and even within trees. Separation of the heartwood from the sapwood in the intertracheid water connection suggested that the presence of water in the heartwood was caused by rewetting of the tracheid lamina that occurred after heartwood formation. The maldistribution of water in the heartwood suggested that a difference in the process of rewetting causes both uneven distribution and the various types of water presence.     

Ultrastructural changes during aging of wood cells

Although most of the wood cells which are produced in the cambium die after the deposition of secondary wall thickening and lignification, the parenchymous cells contain living substances throughout the sapwood and are used for transportation and storage of metabolic materials. In the electron microscope dark globular particles of different sizes were traced from cambium to heartwood border in the parenchymous cells of pine. In the sapwood-heartwood transition zone the dark particles loose their globular shape and the dark content is deposited upon the parenchymous cell wall and therewith upon the membranes of the window-like pits. In this state the membranes are darkly colored so that it is assumed that the dark materials diffuse into the membranes. Dark deposits were observed within the compound middle lamella, in the chambers of bordered pits, and within the fiber lumina. From these observations it is concluded that the dark substances migrate through the middle lamella into the pit chambers and from there into the cell lumina. There is evidence that the decomposition of the globular particles as well as the diffusion of the dark substances into the cell walls begin in a region which is macroscopically considered to be part of the sapwood. Subsequent reactions in the deposition places result in the formation of typical heartwood substances.Presented at the International Wood Chemistry Symposium held at the University of Washington, Seattle, Wash., Aug. 31 to Sept. 4, 1969.     

“I-214杨”心材、边材木质素的红外光谱、质子和碳-13核磁共振波谱特征研究

对从杨树心、边材提取的磨木木质素进行了元素分析和红外光谱（FTIR）质子和碳－13核磁共振波谱（^1H,^13C NMR)等化学特征研究。研究结果表明：杨树心、边材木质素的经验式分别为C9H7．16O2．38（OCH3）1．99和C9H8．61O2．73（OCH3）1．33。心材木质素甲氧基含量28．16％,比边材高8．73％。两种木质素均具有典型阔叶材的特征,化学结构类型基本一致,碳骨架结构基本相同,但化学官能团和键型的组成上存在差异。     

柚木石油醚抽提物成分及其对心边材颜色的影响

为探究国产柚木抽提物对心材和边材颜色差异的影响,采用3种沸程的石油醚溶液分别对国产柚木的心材与边材进行抽提处理,利用气相色谱-质谱法分析抽提液的成分和含量,并以差异较大的抽提物溶液处理柚木边材,通过色度学参数表征柚木边材的变色情况,从而验证影响柚木心材和边材颜色的主要抽提物成分。结果表明:在3种沸程的石油醚抽提液中,柚木心材抽提物在成分种类数量和含量上均显著高于边材;不同沸程的石油醚对柚木心、边材中抽提物成分有一定的影响,其中30~60℃(低沸程)和60~90℃(中沸程)石油醚抽提物成分和含量的差异较小,而90~120℃(高沸程)石油醚抽提物的含量较高,约为中、低沸程抽提物含量的2倍;柚木心材抽提液中酚类、醌类、烯烃类物质的含量较多,但这些物质在边材中含量极低;4-叔丁基-2-苯基苯酚、2-甲基蒽醌(柚木醌)、全反式三十碳六烯(角鲨烯)是柚木心、边材抽提物含量差异最明显的物质。经角鲨烯溶液处理后的柚木边材表面颜色变黄,更接近心材颜色,因此,角鲨烯是导致柚木心边材颜色差异的重要抽提物成分。     

Some factors influencing susceptibility to discoloring fungi and water uptake of Scots pine (Pinus sylvestris), Norway spruce (Picea abies) and Oriental spruce (Picea orientalis)

Abstract

The heartwood and sapwood from Scots pine (PS), Norway spruce (PA), and Oriental spruce (PO) were tested for susceptibility to discoloring fungi and water uptake. In addition, annual ring width and density were measured. The methods used were Mycologg for testing growth of fungi and a modified version of EN 927-5 to investigate water uptake. For pine, the heartwood showed a lower water uptake and no discoloring fungi growing in the tests. The heartwood had a significantly higher density and smaller annual ring width than the sapwood. In PA the heartwood had significantly lower discoloration than sapwood. The total water uptake in g/m² was significantly higher in sapwood, but not the calculated moisture content. As for wood properties, the density was significantly higher in sapwood compared to heartwood, although there were no differences in annual ring width. Regarding PO, differences in water uptake could be seen between sapwood and heartwood although the densities were similar. These results show that susceptibility to discoloring fungi and water uptake is hard to correlate to a single inherent property when looking at different wood species.     

13种杨树木材物理力学性质的研究

采集1－69杨等10种黑杨的9年生试材，易县毛白杨等3种白杨的13年生试材，测试木材物理力学性能，分析心边材，不同树高位置，株间的变化规律，比较品种间木材性质的差异，试验结果表明，13种杨树全树木材心材率在14．6－34．1％，之间，生材含水率在86．2－148．5％之间，由生材干燥至含水率为12％所产生的干缩率在8．66％－11．96％这间，气干密度属低类或很低类，强度属弱类，硬度属软类或甚软类，冲击韧性中等；黑杨心材颜色深，心边材性质差异大，白杨心材颜色淡，心边材性质差异小，品种间木材物理力学性质差异较大，按其主要特点可将13个品种的木材分为4类。     

Collection of expressed genes from the transition zone of <Emphasis Type="Italic">Cryptomeria japonica</Emphasis> in the dormant season

Heartwood affects the utility of wood because it differs in some properties compared to sapwood. To regulate heartwood formation, its mechanism must be elucidated. However, the molecular basis underlying heartwood formation remains largely unknown. To obtain clues to understand the mechanism at a molecular level, we collected expressed sequence tags (ESTs) from the transition zone (TZ) of Cryptomeria japonica D. Don in November, in which heartwood formation is considered to proceed. A total of 1029 ESTs were assembled into 744 unique sequences (103 clusters and 641 singletons). Putative functions were assigned to 291 nuclear-encoded sequences, and they were grouped into 21 categories according to the eukaryotic orthologous groups functional classification. We selected 20 genes for enzymes or proteins, then examined their expression patterns among different organs. The expression levels of nine genes were higher in November than in June in the TZ. The genes encode two enzymes in glycolysis, invertase, methionine adenosyltransferase, glutathione transferase, the lipid transfer protein, Bet v 1 allergen, the dehydrin and the function-unknown protein. This study has provided the first large-scale EST information from the TZ of conifers, which will be useful for understanding the physiological processes in the TZ at a molecular level.     

Heartwood,its function and formation

Summary Some of the changes which occur in wood during the transition from sapwood to heartwood have been reviewed. The nature of these changes suggest that heartwood formation is a regulatory process serving to keep the amount of sapwood at an optimum level. The pattern formed by the transition of cells from sapwood to heartwood suggests that heartwood development is controlled by a centripetally-translocated growth-active substance. The nature of the heartwood transformation is indicative of a developmental process rather than a deterioration of cell function with age so that death of the parenchyma cells is the result and not the cause of heartwood formation.The assistance of Jeanette Gregory, transmission electron microscopy; Dianne Higginbotham, scanning electron microscopy; N. Omar, statistical analysis; C. Taylor, photography; F. R. Humphreys, D. Edwards, D. Adamson, R. C. Foster, and F. V. Mercer, critical comments, is acknowledged. The scanning electron microscopy was undertaken at the Electron Microscopy Unit, Sydney University with the kind co-operation of C. Nockolds. The permission of Marcia Lambert to use unpublished data in Table 2 is also acknowledged.     

Seasonal variation in ethylene concentration in the wood of Pinus sylvestris L

Ethylene concentrations were determined in gas samples extracted from sealed holes made in the sapwood and heartwood of stems of 70-100-year-old Scots pine (Pinus sylvestris L). Gas could be collected from the heartwood holes by lowering the pressure by means of a large syringe. However, attempts to extract gas from air spaces in the sapwood with the same technique failed, presumably because of lack of an interconnected system of gas-filled canals. High ethylene concentrations, usually in excess of 1 ppm, accumulated in the sapwood holes within one day after sealing. Ethylene concentrations in the sapwood rose to 3-7 ppm during the growing season, and decreased to 0.1-0.3 ppm during the winter. In response to extreme drought, sapwood ethylene concentration increased to 30 ppm, followed by a rapid decrease after the onset of rain. Ethylene concentrations in gas samples from the heartwood were consistently lower than 1 ppm. The lowest values, about 0.1 ppm, were found during the autumn and early winter, whereas values around 0.5 ppm were typical from February to August.     

Peroxidase activity, isoenzymes and histological localisation in sapwood and heartwood of Scots pine (Pinus sylvestris L.)

Peroxidase activity and isoenzymes of fresh wood samples of the third shoot of 12-year old trees and from the sapwood, transition zone and heartwood of c. 60-year old stems of Scots pine (Pinus sylvestris L.) were investigated. Wood samples were ground at −30°C, extracted, and the extracts concentrated c. 20-fold for peroxidase activity assays (guaiacol method) and for IEF-PAGE. At least 11 major isoenzymes could be found in the gels. Even the heartwood contained some peroxidase isoenzymes. Isoenzyme patterns of the juvenile wood did not change with the season. However, juvenile wood showed the highest peroxidase activity at the end of the growing season. Peroxidase activity decreased from the outer sapwood towards the heartwood. Thin sections of different wood zones stained for peroxidase revealed activity in ray parenchyma and resin canal epithelial cells. Intensive staining was localised in the bordered pits of vertical and ray tracheids, and in the end walls of ray parenchyma cells.     

Heartwood/sapwood variation of western redcedar as influenced by cultural treatments and position in tree

We studied heartwood and sapwood variation in western redcedar (Thuja plicata) at three sites, including a 95-year-old naturally regenerated, unmanaged stand, a 35-year-old planted spacing trial, and a 30-year-old naturally regenerated stand to which thinning and fertilization treatments had been applied. In the 95-year-old stand, we studied within-tree variation in heartwood and sapwood. In the thinning/fertilization trial and the planted spacing trial, we studied effects of cultural practices and growth rate on heartwood and sapwood. In the trees that we studied, sapwood width was generally fairly narrow, rarely exceeding 3.5 cm. Heartwood formation in western redcedar appeared to begin at a relatively small stem diameter (7 cm) and at a young age, probably 10–15 years. The amount and proportion of heartwood increased with distance downward from the top of the tree, with the implication that older trees will contain a greater proportion of heartwood than younger trees. For any given age, it appears that cultural treatments that favor rapid growth will result in stems with greater amounts of both sapwood and heartwood, and a greater proportion of heartwood.     

Radial movement of sapwood-injected rubidium into heartwood of Japanese cedar (<Emphasis Type="Italic">Cryptomeria</Emphasis><Emphasis Type="Italic">japonica</Emphasis>) in the growing period

Rubidium solution was injected in the sapwood of a Japanese cedar cultivar in the growing period, and its radial movement in stem was traced to investigate the accumulation of alkali metals in the heartwood. Sapwood-injected Rb was detected in outer heartwood at 10 days after the treatment, and continued increasing at 20 days after. Radial movement of Rb toward heartwood was considered to occur soon after the treatment, and to decline at a certain point of the time after Rb injection ceased. However, Rb continued moving in heartwood probably by diffusion even after the cease of Rb injection. In a series of injection experiment, radial movement of injected Rb is not corresponding to the seasonality of both cambial activity and cytological changes of ray parenchyma accompanied with heartwood formation. From the results on Rb’s behavior, we conclude that accumulation of K and other alkali metals in heartwood of Japanese cedar has two steps, active transport from sapwood to outer heartwood via ray, and diffusion in heartwood, and that these processes proceed independently from both cambial activity and cytological changes of ray parenchyma.     

Properties of reaction zones associated with decay from pruning wounds in plantation‐grown Eucalyptus nitens

Summary Decay columns resulting from naturally infected pruning wounds in 5‐ to 8‐year‐old plantation‐grown Eucalyptus nitens in Tasmania are interfaced by a reaction zone in the sapwood. The reaction zone is blue‐purple in colour and occasionally associated with a white zone at the reaction zone/healthy sapwood interface. A log incubation experiment has shown that the reaction zone is particularly durable against decay. The reaction zone is significantly drier than healthy sapwood, with lower levels of potassium, and a lower pH than both sapwood and heartwood. The increased total phenols levels and abundant tyloses in the reaction zone may be particularly important in defence.     

Variation of extractives content in heartwood and sapwood of Eucalyptus globulus trees

The variation in extractives content in sapwood and heartwood was investigated among 12 trees in each of four commercial plantations of Eucalyptus globulus in central Portugal. The study was carried out at the 15% height level and extractions used successively dichloromethane, ethanol and water. At all sites, heartwood had significantly more extractives than sapwood, on average 3.8 and 2.4%, respectively. Most extractives consisted of ethanol soluble material (on average 52% of total extractives). Among the sites, there was a statistically significant difference in the content of extractives but the most important source of variation was the within-tree variation between sapwood and heartwood. Differences in the content of extractives were also observed among trees. A strong relation between extractives content and heartwood proportion was found. The potential loss of pulp yield and problems associated with accumulation of extractives are directly related to the heartwood proportion in the eucalypt stems. Forest management should take into account heartwood development and selection for minimising heartwood extractives.     

Separating Norway spruce heartwood and sapwood in dried condition with near-infrared spectroscopy and multivariate data analysis

Norway spruce [Picea abies (L.) Karst.] heartwood and sapwood have differing wood properties, but are similar in appearance. An investigation was made to see whether near-infrared spectroscopy (NIRS) could be used with multivariate statistics for separation between heartwood and sapwood in dry state on tangential longitudinal surfaces. For classification of wood into sapwood and heartwood, partial least square (PLS) regression was used. Orthogonal signal correction (OSC) filtering was used on the spectra. This study shows that a separation of sapwood and heartwood of spruce is possible with NIR spectra measured in a laboratory environment. The visible-wavelength spectra have significant influence on the predictive power of separation models between sapwood and heartwood of spruce. All 44 specimens in the calibration set were correctly classified into heartwood and sapwood. Validation of the model was done with a prediction set of 16 specimens, of which one was classified incorrectly.