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.     

Effect of pine wood nematode invasion on pine community functions in the Pinghu region, Zhejiang Province, eastern China

In order to investigate the effect of invastion by pine wood nematode （PWN）, this study analyzed severalfunctional indices, i.e., the increment in DBH and stand volume and biomass, in the damaqed stands with various mixedpercentages of Pinus massoniana and P. thunbergii and with different levels of damage. According to the results of rate of change in increment of DBH and stand volume, the forest ecosystem resistance against PWN increased with a reduct on n the m xed ratio of pine. The resistance was highest with a mixed percentage of 50%. The invasion of PWN hanged the corresponding relationship of increment between DBH and stand volume （pure stands 〉 7：3 conifer and roadleaf 〉 6：4 conifer and broadleaf 〉 5：5 conifer and broadleaf） among the P. thunbergii stands when there is no amage, but for P. massoniana stands this phenomenon did not occur. For the increment rate of DBH and stand volume, is significant change in P, thunbergii forest indicates that the resistance of pure P. thunbergii forest was higher than at of P. massoniana. The invasion of PWN accelerates the succession from pure stands to mixed stands and then tohe broadleaf evergreen stands.     

Variation of the chemical composition of Pyrenean oak (<Emphasis Type="Italic">Quercus pyrenaica</Emphasis> Willd.) heartwood among different sites and its relationship with the soil chemical characteristics

This work presents the results of a study on the variation of the chemical composition of Pyrenean oak (Quercus pyrenaica Willd.) heartwood among different sites and its relationship with the soil chemical characteristics. The chemical characteristics of the heartwood are quantified in terms of proximate and ultimate analysis, calorific value and ash composition. Subsequently, the relationship between the wood chemical characteristics and the soil characteristics was also examined. The results showed no significant differences between the wood chemical characteristics from different sites. There are, however, significant differences between trees from the same site regarding the wood ash content, the calorific value and the contents of nitrogen and hydrogen. Strong correlations were observed between the heartwood ash contents of BaO, MnO, CuO and TiO₂ and the soil nitrogen, phosphorous and potassium and the soil exchangeable base contents.     

Direct PCR‐based method for detecting Bursaphelenchus xylophilus,the pine wood nematode in wood tissue of Pinus massoniana

Pine wilt disease (PWD), caused by the pine wood nematode (PWN) Bursaphelenchus xylophilus, leads to serious losses to pine forestry around the world. Pinus massoniana, which is vulnerable to be attacked by the PWN, is the dominant species used in pine forestry in China. The objective of this study is to develop a direct PCR‐based method for detecting B. xylophilus in the wood of P. massoniana without a separate nematode extraction step. A simple procedure was first developed for isolating B. xylophilus DNA in 5 mg pine wood tissue samples harbouring PWN for detection by PCR amplification. A B. xylophilus‐specific amplicon of 403 bp (DQ855275) was generated by PCR from the infested wood tissue. The entire procedure can be completed within 5 h with one pair of primers. This assay can serve as a rapid, cheap and environmentally friendly method to detect B. xylophilus in samples of P. massoniana.     

Comparison of histological responses and tissue damage expansion between resistant and susceptible Pinus thunbergii infected with pine wood nematode Bursaphelenchus xylophilus

Pine wilt disease caused by the pine wood nematode (PWN), Bursaphelenchus xylophilus, has been epidemic and has had disastrous impacts on pine forests and forest ecosystems in eastern Asia. Many pine species in this area are susceptible to this disease. Pinus thunbergii is particularly susceptible. In Japan, tree breeders have selected surviving trees from severely damaged forests as resistant candidates, and have finally established several resistant varieties of P. thunbergii. However, this breeding procedure requires much time and effort due to the lack of physiological and phenotypical information about resistance. To investigate the resistance mechanisms of selected P. thunbergii, we compared histochemical responses, tissue damage expansion, and PWN distribution in resistant and susceptible clones of P. thunbergii after PWN inoculation. The results suggested that the mechanisms of resistance are as follows: damage expansion in the cortex, cambium, and xylem axial resin canals are retarded in resistant trees soon after inoculation, probably due to the induction of wall protein-based defenses. Suppression of PWN reproduction was particularly caused by inhibition of damage expansion in the cambium. The slow expansion of damage in each tissue provides time for the host to complete the biosynthesis of lignin in the walls of cells that surround the damaged regions. This lignification of cell walls is assumed to effectively inhibit the migration and reproduction of the PWNs. The mechanism of initial damage retardation is presumed to be a key for resistance.     

The relationship between terpenes and flammability of leaf litter

Many studies have assumed that plant terpenes favor fire due to their enormous flammability. However, only a few of them, all performed on green leaves, have demonstrated this. In the present work we investigated the question of whether litter terpene content can be used to estimate flammability and temperatures reached during fire. Epiradiator and burn table tests were used to compare flammability of leaf litter of P. pinaster, P. halepensis, P. pinea, C. albidus, C. ladanifer, C. laurifolius and the mixture of litter of P. pinaster with that of the other five species (e.g. P. pinaster + P. halepensis). Tests with burn table showed increasing spread rates and shorter combustion times under higher terpene contents. Flame height was triggered both with higher a terpene content and bed thickness, whereas the percentage of burned biomass was only significantly correlated to bed height. Epiradiator tests indicated that terpene concentration in leaf litter was positively correlated to flame height and negatively correlated to both flame residence time and ignition delay. Flammability was high for P. pinaster, P. halepensis, and hence for P. pinaster + P. halepensis, intermediate for C. albidus, P. pinea and P. pinaster combined with each of these species, and low for C. laurifolius, C. ladanifer and P. pinaster combined with them. Accordingly, their terpene content was high, intermediate and low. We concluded that plants might influence fire intensity, by having stored terpenes in their dead leaves, in addition to having developed traits to survive fire. Thus, a correct management of dead aboveground fuels rich in terpene concentrations, such as those of P. pinaster and P. halepensis, could prove helpful in reducing the hazard of fire.     

Resin tapping in Pinus pinaster: effects on growth and response function to climate

Pinus pinaster is of great ecological and economic importance and has traditionally been subjected to intensive uses such as wood and resin extraction. In the last decade, dendrochronological methods are increasingly being used to analyze the effects of climatic factors on the growth of the maritime pine, although tapped trees were generally avoided because it was thought that their growth was affected by resin extraction. In Spain, however, it is hard to find a long-lived forest of P. pinaster that has not been subjected to tapping for resin. In the present paper, we performed dendrochronological analyses of this species from wood cores and cross sections taken from both resin-tapped trees and resin-untapped trees killed by a fire in 2008 in the central Iberian region. On the one hand, we reconstructed the history of forest management by means of analysis of resin scars in the cross sections of resin-tapped trees. This facet of dendrochronological dating had not heretofore been developed, and little is therefore known about it. We dated 46 scars, which indicate a history of intensive resin extraction in the 1920–1950 period. Moreover, we attempted to answer the question: Have the old resin extractions in P. pinaster precluded the use of their growth rings for dendrochronological and dendroclimatic studies? We found that resin extraction did not alter general short-wavelength variability, and we developed a local chronology with all synchronized series, and the response function with respect to climate is similar to other oldest P. pinaster forests studied in Spain. The information we have recorded can be of use for providing tools to land managers for interpreting forest dynamics in resin-tapped regions.     

白桦木材的变色类型及变色前后化学成分分析

通过化学方法、材面色差分析法、扫描电镜观察和化学成分分析,研究了白桦木材的蓝变和黄变,探讨了变色的原因及主要类型。研究结果表明:白桦蓝变材的变色是由变色菌侵蚀木材引起的,变色深入木材内部。木材薄壁细胞腔内充满了大量的变色菌丝体,菌丝体本身的颜色及其分泌的色素是木材变色的原因。蓝变材部分半纤维素发生降解,其各项抽提物含量比正常材高;而黄变材是由于自然环境变化引起的创伤性木材变色,存在于木材局部区域,其部分导管和少量木纤维细胞被胶状物所填充,内含物增加,木材三大组分没有明显变化。     

Wood colour in Lapacho (Tabebuia serratifolia): chemical composition and industrial implications

This study is aimed at relating the colour of Tabebuia serratifolia (Bignoniaceae family) wood to its chemical composition and technological characteristics, such as hardness and gluing, important for the manufacture of wood flooring. Samples with standard wood colour and evident chromatic anomalies were subjected to CIELAB colorimetric and gas-chromatography analyses. Red and yellow wood colour is associated with the presence of lapachol and dehydro-α-lapachone, respectively, whereas large amounts of polyphenols were found in black parts, together with other quinone compounds. Wood is hardest where lapachol gives rise to a yellow chromatic zone (L1). Delamination tests have shown that polyphenols affect lamella bonding strength. The results may be used by the wood flooring industry to identify the chemical constituents of T. serratifolia according to its colour and foresee possible problems during the production process.     

Within-tree distribution and attractant sampling of propagative pinewood nematode, Bursaphelenchus xylophilus: An early diagnosis approach

Early detection is of primary importance to enable rapid actions to prevent the spread and introduction of invasive species. The pinewood nematode (PWN), Bursaphelenchus xylophilus, a serious invasive and destructive species, is listed as a quarantine pest in the legislation of more than 40 countries. However, Baermann funnel extractions of wood from discs cut from trees at breast-height often do not detect the presence of PWN in infested trees. A serious consequence of such false negatives is the loss of the best window for implementation of eradication or quarantine measures to prevent establishment of incipient PWN infestations. Here we document the within-tree horizontal and vertical distribution of PWN in infested stands in China, using a newly developed kairomonal trapping technique. Our results provide a simple, effective, rapid and non-destructive sampling method that takes into account the changes of PWN within-tree distribution in relation to pine wilt disease (PWD) symptom development. When 60–80% of the foliage has become pale green, PWN is recovered from larger diameter branches. As disease symptoms progress, PWN moves into and down the trunk. As the needles turn yellow, PWN was recovered from the trunk at 1–2 m above the ground. The correlation between the within-tree distribution of PWN and the expression of symptoms indicated a strong association between the distribution of PWN and physiological and pathological changes that develop in attacked pines through the interaction between PWN and tree. This systematic sampling technique takes into account the within-tree distribution of the nematode and should greatly enhance early detection of PWN in field surveys, monitoring and phytosanitary inspections.     

A mid-IR multivariate analysis study on the gross calorific value in longleaf pine: impact on correlations with lignin and extractive contents

Twenty 70-year-old longleaf pine trees from a spacing, thinning, and pruning study were harvested, from which samples were analyzed for gross calorific value (GCV). A strong correlation was found between GCV and extractive contents for the unextracted wood samples. Although lignin content should impact GCV, no correlation was found between the variation in GCV with lignin content for the extractive-free wood samples. Mid-IR spectroscopy coupled with multivariate analysis provided strong correlations between the mid-IR-predicted and calorimetry-determined values for the unextracted wood samples. Plotting the regression coefficients for GCV and extractive contents showed that the same mid-IR bands were responsible for the strength of these models. Spectral differences were observed between the different extract samples, and relative peak intensities appeared to be dependent upon the extractive contents from the wood samples. Thus, models were also built based on the corresponding wood extractive contents using the actual extracts. These provided good correlations, suggesting a proportional change in extractive compositions coinciding with the total amount of extractives present in the unextracted wood samples.     

Variability of some wood properties of eastern cottonwood (Populus deltoides Bartr.) clones

The objective of this article is to investigate the variability in chemical composition, fiber length, and basic wood density (specific gravity) amongst clones. Wood samples were taken from an experimental plantation in a dense 2.5 × 2.5 m wood, i.e., 1600 trees/ha. The plantation consisted of 127 clones which were selected in the nursery from 11 eastern cottonwood (P. deltoides Bartr., section Aigeiros) half-sib progenies. The physical, structural, and chemical characteristics of 40 poplar clones aged four years were investigated in three model trees from each clone. There are significant differences between the contents of the individual components of the wood chemical composition. The values of the heritability factor are high and they are similar for lignin (0.936), holocellulose (0.937) and for the extractives content (0.999). The mean value of the basic wood density is 380 kg/m³ (coefficient of heritability is 0.938), and the mean value of the fiber length is 863 m (coefficient of heritability 0.611).     

Pine wilt disease: detection of the pinewood nematode (Bursaphelenchus xylophilus) as a tool for a pine breeding programme

The pinewood nematode (PWN), Bursaphelenchus xylophilus, is a serious quarantine pest first detected in Portugal and Europe in 1999. It is the causal agent of pine wilt disease (PWD). A resistance breeding programme has been initiated to contribute to control the evolution of the disease. Five hundred and four adult maritime pine, Pinus pinaster, trees were phenotypically selected as candidate trees for this programme from an area affected by PWD. To identify tolerance to the nematode, the selected trees were monitored monthly. Over the course of 1 year, 57 candidate trees died and were tested for the presence/absence of the PWN. As accuracy of detection is of major importance, an ITS‐PCR‐based method applied directly to wood from adult maritime pine trees was tested and compared with a standard morphological identification method. The results showed that the use of PCR to detect the pathogen provided more rapid and accurate results in comparison with the standard morphological identification. Thus, this method is suitable to be used in the survey of the breeding population for resistance/tolerance to PWD.     

Treatment of Softwoods with CCA at High Moisture Levels

A literature study on the feasibility of treating wood with aqueous solutions at high moisture levels revealed a positive indication. The objective of the present study was to determine treatability of Pinus radiata and P. pinaster lumber at high moisture levels with commercial CCA in controlled experiments. The wood samples were treated at levels of moisture content ranging from 20 to 100%.

A statistical analysis of the data showed that specific gravity and moisture content influence solution-absorption, the effect being different for different species. Chemical analyses showed a screening out of copper- and arsenic compounds. The effect of higher moisture contents on screening out was negligible. The results indicated that certain wood species can be treated satisfactorily with CCA at moisture levels higher than 25%.     

Calorific values and ash contents of different parts of Masson pine trees in southern China

Calorific value of plants is an important parameter for evalu- ating and indexing material cycles and energy conversion in forest eco- systems. Based on mensuration data of 150 sample sets, we analyzed the calorific value （CV） and ash content （AC） of different parts of Masson pine （Pinus massoniana） trees in southern China using hypothesis testing and regression analysis. CV and AC of different tree parts were almost significantly different （P〈0.05）. In descending order, ash-free calorific value （AFCV） ranked as foliage 〉 branch 〉 stem bark 〉 root 〉 stem wood, and AC ranked as foliage 〉 stem bark 〉 root 〉 branch 〉 stem wood. CV and AC of stem wood from the top, middle and lower sections of trees differed significantly. CV increased from the top to the lower sections of the tnmk while AC decreased. Mean gross calorific value （GCV） and AFCV of aboveground parts were significantly higher than those of belowground parts （roots）. The mean GCV, AFCV and AC of a whole tree of Masson pine were 21.54 kJ/g, 21.74 kJ/g and 0.90%, re- spectively. CV and AC of different tree parts were, to some extent, cor- related with tree diameter, height and origin.     

Regeneration dynamics of mixed stands of <Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait. and <Emphasis Type="Italic">Pinus pinea</Emphasis> L. in Central Spain

The dynamics of mixed stands are more complex and less studied than those of monospecific stands. The objective of this work was to analyze the variables involved in seedling occurrence and seedling survival in mixed stands of Pinus pinaster and P. pinea in Mediterranean areas. From 2011 to 2016, regeneration of both species was monitored at two sites located in Central Spain. We installed 72 regeneration plots where seedling dynamics were monitored. All the trees in the study areas were measured and mapped. Additionally, we took hemispherical photographs in each regeneration plot. The average density of P. pinea seedlings over the study period was almost 20 times larger than that of P. pinaster. Our results indicate that the seedlings of both species grow under moderate light conditions. In addition, we found that the occurrence of seedlings of both species was related to the structure of the stand. P. pinea seedlings grew where the density and size of P. pinaster trees were low and where P. pinea trees provided moderately sheltered conditions, whereas the number of P. pinaster seedlings was related to under intermediate densities of P. pinaster trees. Furthermore, seedling survival was positively associated with age of the seedlings and negatively with the August average maximum temperature. The temporal continuity of mixed stands of P. pinea and P. pinaster in the study area is compromised by the observed lack of regeneration of P. pinaster.     

Characterization and photodegradation mechanism of three Algerian wood species

Aging is the irreversible change of mechanical, physical, and chemical properties of materials; the main objective of this work was to study the photochemical degradation and structural changes of three major Algerian wood species. For this, we evaluated the photodegradation mechanism for Maritime Pine (Pinus pinaster), zeen oak (Quercus canariensis), and afares oak (Quercus afares) by accelerated aging in a Xenon test chamber. Degradation of the samples was established by an initial color change (after 30 h exposure), followed by roughening and cracking (120 h exposure) as translated by scanning electron microscopy and Fourier transform infrared spectroscopy. The discoloration of irradiated wood samples was primarily related to the decomposition of lignin which is the key structure in wood photodegradation. As expected, a decrease in mechanical properties was observed; for all samples, the modulus of elasticity decreased after aging, indicating that the wood specimens loss some of their stiffness.     

Migration of Bursaphelenchus xylophilus in cortical and xylem axial resin canals of resistant pines

Migration of the pine wood nematode (PWN), Bursaphelenchus xylophilus, in susceptible and resistant pines was investigated at the tissue level. PWN was inoculated onto the top cross‐cut surface of 20‐cm stem cuttings of susceptible Pinus thunbergii and resistant pines (P. strobus, P. rigida and P. thunbergii of a resistant family Namikata‐(t)‐73 (half‐sib)). PWNs were mainly distributed in cortical resin canals of susceptible P. thunbergii down to 15 cm from the inoculated surface by 6 h after inoculation (HAI) and all tissues (including cortical and xylem resin canals) down to the bottom at 192 HAI. In P. strobus, P. rigida and P. thunbergii family Namikata‐(t)‐73 (half‐sib), PWN was distributed in cortical resin canals down to 5 cm by 6 HAI and down to the base at 192 HAI. However, the distribution of PWN in xylem resin canals of the resistant pines was restricted near inoculated surfaces down to 5 cm, even at 192 HAI. These results demonstrated that migration of PWN in resistant pines was slowed in cortical resin canals and restricted in xylem axial resin canals, features which may be associated with the resistance.