Evolutionary history shapes variation of wood density of tree species across the world

The effect of evolutionary history on wood density variation may play an important role in shaping variation in wood density,but this has largely not been tested.Using a comprehensive global dataset including 27,297 measurements of wood density from 2621 tree species worldwide,we test the hypothesis that the legacy of evolutionary history plays an important role in driving the variation of wood density among tree species.We assessed phylogenetic signal in different taxonomic(e.g.,angiosperms and gymnosperms)and ecological(e.g.,tropical,temperate,and boreal)groups of tree species,explored the biogeographical and phylogenetic patterns of wood density,and quantified the relative importance of current environmental factors(e.g.,climatic and soil variables)and evolutionary history(i.e.,phylogenetic relatedness among species and lineages)in driving global wood density variation.We found that wood density displayed a significant phylogenetic signal.Wood density differed among different biomes and climatic zones,with higher mean values of wood density in relatively drier regions(highest in subtropical desert).Our study revealed that at a global scale,for angiosperms and gymnosperms combined,phylogeny and species(representing the variance explained by taxonomy and not direct explained by long-term evolution process)explained 84.3%and 7.7%of total wood density variation,respectively,whereas current environment explained 2.7%of total wood density variation when phylogeny and species were taken into account.When angiosperms and gymnosperms were considered separately,the three proportions of explained variation are,respectively,84.2%,7.5%and 6.7%for angiosperms,and 45.7%,21.3%and 18.6%for gymnosperms.Our study shows that evolutionary history outpaced current environmental factors in shaping global variation in wood density.

Genetic Control of Air-Dried Wood Density, Mechanical Properties and Its Implication for Veneer Timber Breeding of New Triploid Clones in Populus tomentosa Carr

The wood samples of 9 triploid clones of Populus tomentosa Carr. taken from a 9 year old clonal test site were analyzed in order to investigate the genetic variation of wood properties, including air dried wood density and some mechanical properties. The results showed that significant or extremely significant difference in air dried wood density and the mechanical properties existed among the clones, this means these wood properties were under moderate or strong genetic controls and could be improved by genetic manipulations. The radial and vertical variation patterns of air dried wood density were also studied and the results were found to coordinate with other previous research results. The vertical variation patterns of most mechanical properties within the individual tree also conformed to the general wood theories except the modulus of elasticity and cross section hardness. Among the mechanical properties, modulus of elasticity (MOE) and tangent section hardness were under strong genetic control, with the clonal repeatabilities being 0 90 and 0 80, respectively. However, the clonal repeatabilities of other mechanical properties under study were a little lower than above two indexes. Genetic correlation analysis indicated that super clonal selection and breeding for veneer timber could be realized through indirect selection of wood density and form indexes.

Wood Density Determination with the Perspective to Decarbonisation of Tropical Forest Species from the Luki Biosphere Reserve in the Democratic Republic of the Congo

Specimens of the forest species such as Pentaclethra macrophylla, Petersianthus macrocarpus, Pycnanthus angolensis and Terminalia superba have been sampled from LUKI Biosphere reserve in the Democratic Republic of the Congo in order to determine their wood density with the perspective to decarbonisation. These parameters have been found out experimentally utilizing a drying technique in an oven including techniques of immersion in an Erlenmeyer full of water. The corresponding results indicated that the four species wood density is respectively 0.85, 0.80, 0.77 and 0.51. These preliminary results will be useful in our ongoing project on carbon dioxide absorption capacity of Congo rainforest tree species.

Relating estimates of wood properties of birch to stem form, age and species

Birch has long suffered from a lack of active forest management,leading many researchers to use mate-rial without a detailed management history.Data collected from three birch(Betula pendula Roth,B.pubescens Ehrh.)sites in southern Sweden were analyzed using regression analysis to detect any trends or differences in wood proper-ties that could be explained by stand history,tree age and stem form.All sites were genetics trials established in the same way.Estimates of acoustic velocity(AV)from non-destructive testing(NDT)and predicted AV had a higher correlation if data was pooled across sites and other stem form factors were considered.A subsample of stems had radial profiles of X-ray wood density and ring width by year created,and wood density was related to ring number from the pith and ring width.It seemed likely that wood density was negatively related to ring width for both birch species.Linear models had slight improvements if site and species were included,but only the youngest site with trees at age 15 had both birch species.This paper indicated that NDT values need to be considered separately,and any predictive models will likely be improved if they are specific to the site and birch species measured.

Wood density and carbon concentration of coarse woody debris in native forests,Brazil

Background: With the objective of increasing knowledge on biomass and carbon stocks, and thus improving the accuracy of published estimates, the present study explored wood density and carbon concentration of coarse woody debris (diameter≥10) by decay class in a Seasonal Semi-deciduous Forest (SSF) area in the Atlantic Rain Forest and in a Cerrado sensu-stricto (CSS) area (Brazilian savanna), in Brazil. Two strata were identified in each area and ten sampling units were systematic located in each stratum. Data were collected according to the line intersect sampling method. Each tallied element, the diameter, length, and perpendicular width were recorded at the transect intersection point. Each element was classified into a decay class, and the species was identified when possible. Sample discs were cut from each element, from which cylindrical samples were extracted and oven-dried to determine density. These cylinders were milled and analyzed using a LECO-C632 to determine carbon concentration as percentage of mass. Results: In both areas, wood density decreased as the decay class increased. For SSF the mean carbon concentration of all analyzed samples was 49.8% with a standard deviation of 3.3, with a range of 27.9–57.0% across 506 observations. For CSS the general mean was 49.6% with a standard deviation of 2.6, with a range of 31.2–54.5% over 182 observations. Carbon concentration barely change between decay classes. Carbon stock was estimated at 3.3 and 0.7 MgC/ha for the SSF and the CSS, respectively. Similar results were obtained when using a 50% conversion constant. Conclusions: The present study concludes that wood density decreases as the woody debris becomes more decomposed, a pattern found in many previous studies. The carbon concentration, however, barely changes between decay classes, and that result is consistent with most of the literature reviewed. Our carbon concentrations are very close to the 50% used most commonly as a conversion factor. We strongly recommend that future studies of CWD evaluate wood density and carbon concentration by decay class to address the uncertainty still found in the literature.

Growth and wood properties of a 38-year-old Populus simonii 3 P.nigra plantation established with different densities in semi-arid areas of northeastern China

To characterize the effects of plantation densities on the growth characteristics(diameter at breast level,tree height and volume)and the common wood properties,38-year-old Populus simonii 9 P.nigra clones planted with four levels of spacing(2 m 92 m,3 m 93 m,4 m 94 m,and 5 m 95 m)in a semi-arid area in northeastern China were examined.The results of ANOVA showed significant differences(P\0.01)for all the investigated growth traits and wood properties under different plantation densities,except for the chemical composition of wood.The repeatability and phenotypic variations of all the traits varied from 0.34 to 0.99 and from 13.45 to 59.65%,respectively.Except for wood density,which was significantly negatively correlated with the growth traits,a positive correlation was observed between the growth traits and all the other wood mechanical properties.However,most of the correlations between the growth traits and the chemical composition of the wood were not significant.The path analysis for the wood mechanical characteristics and the growth in the prediction of volume were significant and ranged from 0.18 to 0.72 for wood density and diameter at breast height,respectively,while those for the chemical composition of wood ranged from 0.001 to 0.336,which showed a low impact on the volume.The highest stand volume(610 m3)per hectare was observed with the 2 m 92 m spacing,which consequently provided a high total price and income,while a high individual volume growth per tree was observed with the 5 m 95 m spacing.The results suggested that for the poplar trees younger than 40 years in a semi-arid area in China,2 m 92 m spacing is suitable for obtaining a high volume per hectare,while 5 m 95 m spacing is best for obtaining a high individual volume per tree.

Genetic variation of wood tracheid traits and their relationships with growth and wood density in clones of Pinus tabuliformis

To improve wood quality for pulpwood industries, it is important to examine not only wood density but also its components, especially tracheid characteristics. We studied genetic variations in the following tracheid traits by earlywood （EW） and latewood （LW）： tracheid length （TL）, double wall thickness （WT）, radial lumen diameter （R_D1）, tangential lumen diameter （T_D1）, radial central diameter （R_D2）, and tangential central diameter （T_D2）. We also studied the relationship with the following growth traits： diameter at breast height （DBH）, height （H）, crown breadth south-north axis （NSC）, crown breadth east-west axis （EWC）, ring width （RW）, latewood percentage （LWP）, and wood density （WD）. All sample materials were collected from a 33-year old clonal seed orchard of Pinus tabuliformis Carr. Genetic variation among clones was moderate for all tracheid traits, 9.49-26.03%. Clones significantly affected WT, R_D1, R_D2, T_D1, T_D2, and the two ratios WT/R_D1 and TL/T_D2 in EW but had no effects in LW. Clones significantly affected TL in LW but had no effects in EW. H2/C was higher in LW （0.50） than in EW （0.20） for TL, while H 2/C was higher in EW （0.27-0.46） for other tracheid traits and the two ratios （TL/T_D2 and WT/R_D1） than in EW （0.06-0.22）. WD and TL were significantly positively correlated, but WT and TL were negatively correlated both at individual and clone levels; all tracheid diameters and the four ratio values （EW_WT/ R_D1, LW WT/R_D1, EW_TL/T_D2 and LW_TL/ T_D2）, were strongly positively correlated with DBH, H, NSC, WEC and RW, and strongly negatively correlated with WD both at individual and clone levels. The most important variables for predicting WD were LW_TL, EW_WT and R_D1 in both EW and LW （r2= 0.22）. Selecting the top 10% of the clones by DBH would improve DBH growth by 12.19% （wood density was reduced by 0.14%） and produced similar responses between EW and LW for all tracheid traits： a reduction of 0.94 and 3.69% in tracheid length and increases in tracheid diameters （from 0.36 to 5.24%） and double wall thickness （0.07 and 0.87%）. The two ratios WT/R_D1 and TL/T_D2 across tissues （EW and LW） declined 0.59 and 4.56%, respectively. The decreased tracheid length and the ratio between tracheid length and diameter is disadvantageous for pulp production. The unfavorable relationship of tracheid traits with wood density indicate that multiple trait selection using optimal economic weights and optimal breeding strategies are recommended for the current longterm breeding program for P. tabuliformis.

Variability of European beech wood density as influenced by interactions between treering growth and aspect

Background： Wood density is considered to be the most important predictor of wood quality but despite its importance, diffuse-porous tree species have been the subject of only a limited number of studies. The importance of European beech forests for Central Europe calls for profound research to examine the potential impact of a warmer climate on the quality of beech timber. Methods： In this study we analysed the influence of tree-ring width and tree-ring age on the wood density of beech, and whether the wood density response to these two parameters is modified by aspect. A linear mixed-effects model for wood density was constructed for mean density data measured with high frequency densitometry on stem discs from 72 beech trees sampled from two different aspects （northeast -NE and southwest -SW） of a valley in southwestern Germany. Results： Part of the variability of mean annual wood density was explained by cambial age： an increase in cambial age resulted in an increase in mean wood density. Tree-ring width and aspect had only a small influence on wood density, Wood density on the SW aspect was lower than on the NE with a difference of approximately 0.006 g/cm3. The between-tree variability was very high. Conclusions： The significant interaction between cambial age and aspect reflects the importance of site conditions at older tree ages： with increasing cambial age the difference between aspects becomes stronger. Our results give a better understanding of the importance of site conditions on the wood quality of beech.

Environmental load of solid wood floor production from larch grown at different planting densities based on a life cycle assessment

As one of the main structural units in a building,a solid wood floor has significant strategic research value for low-carbon energy saving.Taking the production line of a solid larch wood floor as a case study,we assessed the environmental load during production based upon a life cycle assessment.Using GaBi 6.0 software,we analyzed the associated carbon sequestration during floor production,with the initial planting density serving as the disturbance factor in a modular analysis.The results indicated that the cutting and finishing steps have relatively intense,negative influences on the environment,whereas transportation,ripping,and trimming do not.Additionally,recycling biomass waste has the potential to reduce greenhouse gas emissions.When the initial planting density was 3.0×3.0 m,carbon sequestration was relatively high.Although the emissions of freshwater pollutants,volatile organic compounds,and fine particulate matter（matter with a 2.5-μm diameter） were comparatively high,the reduction of greenhouse gas emissions was still excellent at this planting density.

Effect of site and elevation on wood density and shrinkage and their relationships in Carpinus betulus

This study examined the effect of site and elevation on the oven-dry density and volumetric shrinkage of hornbeam （Carpinus betulus） wood. For this purpose, 45 normal hornbeam trees from three different sites （Mazandaran, Guilan and Golestan） at three elevations, 300 m （low elevation）, 700 m （intermediate elevation） and 1100 m （high elevation） from natural forest in northern Iran were selected. Disks were cut at breast height from mature wood and test samples were prepared to determine physical proper- ties, i.e., wood density and volumetric shrinkage. The results show that there are significant differences between site and elevation on these physical properties of wood. The interaction effects between site ~ elevation on wood density and volumetric shrinkage were also statistically significant. At any fixed site, wood density and volumetric shrinkage increased with elevation. As well, their values at Guilan were higher than those at the other two sites. The relationship between wood density and volumetric shrinkage was deter- mined by simple regression equations. Strong, positive relationships between wood density and volumetric shrinkage at each site were found. Again, these were stronger at the Guilan site than at the other two sites. Given a fixed site, the correlation at high eleva- tion was stronger than at the other two elevations.

Variation of basic density,calorific value and volumetric shrinkage within tree height and tree age of Ugandan grown Eucalyptus grandis wood

Insufficient knowledge on Ugandan grown Eucalyptus grandis W.Hill ex Maiden wood properties,high demand,and processing challenges led to a study into its physical properties.We obtained the variation of basic density(BD),calorific value(CV)and volumetric shrinkage(VS)within tree height and tree-age of E.grandis,and its appropriate use based on these properties.Trees with good boles were harvested from Kabarole District in western Uganda to produce specimens as prescribed by British Standards and ASTM standard wood testing procedures.Secondary data reviews and statistical analysis using ANOVA,Tukey’s test and multivariate analysis were done to obtain property estimates and their variation within trees and amongst tree ages.The mean BD of E.grandis is 413.6,380.5,471.0,and 501.1 kg m^(−3)at 3,6,9,and 12 years,respectively,showing significant increase with tree age(p=0.003).The pattern of BD with tree height showed a reduction with tree height although with higher values in the middle portion of the tree.The CV increases(p=0.014)with tree age and reduces with tree height with values of 14,560.32,15,447.3,16,079.11,and 16,932.6 kJ kg^(−1)at 3,6,9,and 12 years,respectively.The percentage VS was 11.02,9.84,12.31,and 14.45 for 3-,6-,9-,and 12-year-old trees,respectively,and it did not vary significantly(p=0.088)with tree height.Basing on these property values,E.grandis wood could be used for scaffolding,light constructions and fuel wood production.Its seasoning needs to be longer with well monitored drying schedules to reduce seasoning defects caused by the high VS.Further studies on strength properties,seasoning schedules,panel products properties and tree-age chemical variations would improve the knowledge about its wood quality and would enhance its efficient utilization.

Basic density, extractive content and moisture sorption properties of Pinus pinaster wood infected with the pinewood nematode, Bursaphelenchus xylophilus

The pinewood nematode（PWN）, Bursaphelenchus xylophilus, has become one of the most severe threats to pine forest worldwide. Nematodes, migrating through resin canals and feeding on the living cells, induce rapid metabolic changes in ray parenchyma cells, create cavitation areas, decrease xylem water content and oleoresin exudation, and cause necrosis of parenchyma and cambial cells. This study focused on the impact of PWN infection on technological parameters of wood and evaluated the impact of anatomic and biochemical incidences of tree defense reactions on basic density, extractive content and moisture sorption properties of Pinus pinaster wood.Samples of infected and uninfected wood were studied.The presence of nematodes reduced wood basic density by2 % and decreased the total content of extractives in infected wood as compared with uninfected（5.98 and8.90 % of dry wood mass, respectively）. Extractives in infected trees had inverse distribution along the trunk as compared with uninfected trees. The adsorption isotherms for infected and uninfected wood had similar positioning.We recorded differences（some statistically significant） in the equilibrium moisture content of infected and uninfected wood under varying environmental conditions. Despite the verified differences in wood basic density, extractive content and moisture sorption properties, the overall conclusion is that the PWN had a slight impact on these characteristics of wood.

STUDY ON WOOD COMPOSITES DENSITY UNIFORMITY BY X-RAY

X-ray is irradiated in some wood composites(multi-layer particleboard.sin-gle-layer particleboard,reconsolidated wood,).According to the gray degree principle(0-255 grade degree)and the corresponding relation between density and gray,X-raynegatives are scanned.The numbers and pictures of every degree density are obtained andaccurate composite densities are counted.

EFFECT OF TIMBER DENSITY ON RETENTION OF WOOD PRESERVATIVES

The amount of CCA fixed in the southern pine treated with CCA have been eval-uated.The conclusions from these experiments are:with the timber density increases,the reten-tion based on weight/weight will decrease,but the retention based on weight/volume will bealmost constant.Retention by weight would be more influential than by volume.

Simulation of the growth ring density of Larix olgensis plantation wood with the ARMA model

Because growth ring data have temporal features, time series analysis can be used to simulate and reveal changes in the life of a tree and contribute to plantation management. In this study, the autoregressive(AR) and moving average modeling method was used to simulate the time series for growth ring density in a larch plantation with different initial planting densities. We adopted the Box–Jenkins method for the modeling, which was initially based on an intuitive analysis of sequence graphs followed by the augmented Dickey–Fuller stationarity test. The order p and q of the ARMA(p, q) model was determined based on the autocorrelation and partial correlation coefficient figure truncated on the respective order.Through the residual judgment, the model AR(2) was only fitted to the larch growth ring density series for the plantation with the 1.5 9 2.0 m^2 initial planting density.Because the residuals series for the other three series was not shown as a white noise sequence, the modeling was rerun. Larch wood from the initial planting density of2.0 9 2.0 m^2 was modeled by ARMA(2, 1), and ARMA((1, 5), 3) fitted to the 2.5 9 2.5 m^2 initial planting density,and the 3.0 9 3.0 m^2 was modeled by AR(1, 2, 5).Although the ARMA modeling can simulate the change in growth ring density, data for the different growth ring time series were described by different models. Thus, time series modeling can be suitable for growth ring data analysis, revealing the time domain and frequency domain of growth ring data.

Effects of site conditions on growth and wood properties of Populus×euramericana cv.'74/76'

The growth and wood properties of 240 individual Populus×euramericana cv.’74/76’(hereafter poplar 107)trees planted in Hebei Plain,China was evaluated.Mean annual increments in height,breast height diameter and volume,as well as cellulose,hemicellulose and lignin contents,shrinkage,density,bending strength and modulus of elasticity in the heart wood and sap wood.Environmental factors influencing growth and wood properties were analyzed using correlation and stepwise regres sion.The results show that the coefficients of variation(CVs)of growth traits ranged from 10.6 to 22.4%.The CVs of the chemical properties of heartwood ranged from 4.3 to 30.2%,and for sap wood from 3.2 to 27.5%.The CVs of the physical and mechanical properties of heartwood ranged from 8.6 to 31.7%,and for sapwood from 6.4 to 29.9%.The results of one-way ANOVA showed that there were significant differences in growth traits and wood properties among sites.Soil pH,total and available phosphorus,total potassium,and soil organic matter were key soil factors affecting growth and wood properties of poplar 107,whereas mean annual ground temperatures and precipitation were the main climatic factors.To better cultivate poplar 107,area with less annual rainfall,slightly higher temperature and soil pH value close to neutral should be selected.

Variability in Wood Density and Wood Fibre Characterization of Woody Species and Their Possible Utility in Northeastern Mexico

Context: Preliminary screening has been undertaken by woody species of Tamaulipan thornscrub for wood density and its utilization northeastern Mexico for different purposes and to find possible relationship of density with wood fibre characteristics. This technique can be used in the selection of species with high wood density for possible utility. Aims: It is to determine the variability in wood density and in fiber cell morphology and its length and among wood species of the Tamaulipan thornscrub, northeastern Mexico. Methods: Wood density and wood fibres are characterized in these woody species following standard methodology. Results: The results reveal large variability in wood density and in fibre cell morphology h. The species have been classified on the basis of wood density and its fibber cell morphology and has been recommended for their possible utilization for different purposes. Conclusions: Species desirable for strong furniture making, paper pulp, soft furniture, fence etc. can be selected on the basis of fibre length to breadth ratios as strong fibres for furniture and fibre cells with broad lumen and thin cell wall use for fabrication of paper pulp and other utilities.

Wood Density of Ten Native Trees and Shrubs and Its Possible Relation with a Few Wood Chemical Compositions

The present study was undertaken in Forest Science Faculty, Universidad de Nuevo Leon, Mexico on variability of Wood density and its possible relation to few wood chemical composition and wood fiber cell structure anatomy. The results reveal that among 10 specie studied, there exist a large variation in wood density (0.51 to 1.09), and few wood chemical composition such % carbón (37.14 to 44.07), nitrogen (9.18 to 19.22), sulphur (31.45 to 33/82), lignin (15/28 to 24.35), hemicellulose (19.94 to 27.36%), and % cellulose (33.69 to 45.92). In general, though there was no clear relationship between wood density and other chemical composition of wood. It was observed that the species having moderate to high wood density contained >40% carbón, >30% sulphur and >40% cellulose and more or less 20% lignin. It seems that carbón, sulphur, cellulose and lignin content contribute to greater density. The wood fiber cell with wall lignification seems to be related to higher wood density.

Density and Strength of Mortar Made with the Mixture of Wood Ash, Crushed Gneiss and River Sand as Fine Aggregate

The main purpose of this paper is to study the feasibility of using wood bottom ash to partially replace natural fine aggregate or crushed gneiss sand in the manufacturing of mortars. The experiment uses wood ash as fine aggregates, which passes through 5 mm sieve, in proportions of 5%, 10%, 15%, 20% and 25% by weight to replace partially river sand and crushed gneiss, and the both sand of the same size as the aggregate respectively. Experimental results show that density of mortar and the compressive strength of mortar decrease globally with the increase in wood ash content. At 56 days, and for all replacements with wood ash, compressive strengths values of mortar obtained with the mixture of wood ash and river sand is greater than 20 MPa, which is not the case for mortar made with crushed gneiss and wood ash. Moreover, for 5% of replacement with wood ash, compressive strengths of mortar obtained with the mixture of wood ash and river sand and the mixture of wood ash and crushed gneiss are respectively 37 MPa and 32 MPa at 56 days. These values satisfied the strength requirements. Hence, 5% replacement of crushed gneiss with wood ash is suggested and could be benefit for mortar. In addition, the replacement of sand by wood ash is preferable with river sand which contains fewer fines than crushed gneiss. The compressive strength of mortar with 25% wood ash + river sand could be suitable.

Removing harvest residues from hardwood stands affects tree growth, wood density and stem wood nutrient concentration in European beech (Fagus sylvatica) and oak (Quercus spp.)

Background:Higher exportation of harvest residues from forests due to increased demand for woody biomass,has reportedly diminished soil mineral resources and may lead to degraded tree nutrition as well as growth.However,as nutrients become less available in the soil,the remobilization of nutrients in biomass tissues(plant internal cycling)helps sustain tree nutrition.Our study aims to quantify the impact of Removing Harvest Residues and Litter(RHRL)during five years on tree growth,wood density,and stem wood nutrient concentrations in young beech and oak forest stands.Result:Our study found that,RHRL significantly decreased tree growth ring width by 14%,and wood density by 3%,in beech trees,in near bark rings.RHRL also significantly reduced nutrient concentration in near bark and near pith areas of both studied species.Mg,Na and S were found lower by 44%,76%,and 56%,respectively,in near bark area of beech trees.In near bark area of oak trees,K,Ca,Mg,Na,S,and Fe were lower by 20%,25%,41%,48%,41%,and 16%,respectively.K and Mg concentrations decreased more strongly in near pith area compared to near bark area suggesting internal translocation of these two elements.Conclusion:In beech trees,wood density proved to be an important factor while quantifying the effect of removing harvest residuals on tree growth and biomass.Soil nutrient loss intensified the remobilization of nutrients con-tained in older tree rings(close to the pith)towards newly formed rings(close to bark).In our study,in beech trees,K was found to be the most recycled major nutrient.These results demonstrate the potential of such analysis for providing valuable insight into the effect of RHRL in premature stands on the physiological adaptive strategies of trees and an indication of soil fertility status.