Variability of selected wood characteristics in 40 half-sib families of black spruce (Picea mariana)

Summary Based on 15-year-old black spruce (Picea mariana) trees from 40 half-sib families sampled from 9 blocks of a family test in New Brunswick, this study examined intertree and intratree variation in various wood density and ring width characteristics. Of various variance components of the intertree variation, a remarkable variance component due to family was found in wood density characteristics (viz. average wood density, average earlywood density and latewood density of the tree), and these characteristics are thus under strong genetic control (h _i ² ranging from 0.60 to 0.86, and h _f ² from 0.56 to 0.68). It, to a lesser extent, applies to ring width characteristics at the tree level (viz. average ring width, and average earlywood width, latewood width and latewood percent of the tree) that show a lower heritability (h? from 0.18 to 0.28, and h _f ² from 0.22 to 0.36). Both block and block × family interaction contribute little to the total intertree variation encountered in 40 families from 9 blocks, while tree-to-tree variation within the family accounts for most (over 3/4) of the total intertree variation.Compared to the intertree variation (tree-to-tree variation within the family), the intratree variation in various wood characteristics studied is considerably larger in this species. It appears that most intraring wood density characteristics show a relatively smaller intertree variation but a relatively larger intratree variation as compared to ring width characteristics (except latewood width and latewood percent). Latewood width and latewood percent show the smallest intertree variation and the largest intratree variation. Between the two sources of the radial intratree variation, cambial age explains much more variation in most intraring wood density characteristics, while ring width accounts for more variation in earlywood width, latewood width and intraring density variation. This indicates that wood density of growth rings in this species is dependent more on cambial age than ring width (growth rate). Among various wood density and ring width characteristics studies, maximum (latewood) density shows the strongest response to calendar year. This characteristics is thus a useful dendroclimatic parameter in this species.I would like to thank Dr. E.K. Morgenstern and Mr. D. Simpson for their involvement in the planning of this study. Thanks are also due to G. Chauret, T. Keenam, R. Ploure, V. Steel and C. Reitlingshoefer for their technical assistance     

Genetic control of intra-ring wood density variation in hybrid larch (Larix gmelinii var. japonica × L. kaempferi) F1

Genetic parameters for various wood density traits were estimated in 29-year-old trees of 18 full-sib families of hybrid larch (Larix gmelinii var. japonica × Larix kaempferi) F₁. Intra-ring density variation (IDV) was also evaluated using a model that expresses the pattern curve from earlywood to latewood as a power function. A high IDV indicates an abrupt change in wood density from earlywood to latewood. The ring width and wood density traits of individual rings were determined by X-ray densitometry. Overall wood density (RD) was shown to increase with increasing ring number, ranging from 0.42–0.59 g/cm³, whereas IDV of individual rings decreased gradually from pith outwards. Estimates of individual tree narrow-sense heritability of RD and IDV were 0.66 and 0.67, respectively. IDV showed negative genetic and phenotypic correlations with RD (r _g = −0.99, r _p = −0.72). The predicted genetic gains in latewood proportion and IDV were higher than that of RD. These results suggest that the intra-ring density variation is under moderate genetic control equivalent to wood density. The trend of increasing wood density from earlywood to latewood was associated with changes in both tracheid diameter and cell wall thickness.     

Genetic parameters for early wood and latewood densities and development with increasing age in Scots pine

? Each annual ring in pines consists of earlywood and latewood with considerable difference in density and width. To get a better determination of the genetic regulation of total wood density in Scots pine (Pinus sylvestris L.), density and width of those ring sections were measured in annual ring numbers 12 to 21 of Scots pines in a full-sib progeny test. Tree height and stem diameter were also measured.

? Heritabilities for the annual ring sections increased with age for earlywood density from 0.08 to approximately 0.25; latewood density showed similar reductions. Heritability over all 10 annual rings was 0.25 for earlywood density, 0.22 for latewood density, 0.29 for height and 0.10 for stem diameter. Genetic correlations between earlywood and latewood density and growth traits were negative, while they were strongly positive between densities of adjacent annual rings (0.70–1.0).

? Despite the higher heritability of earlywood density, the strong positive genetic correlation between those traits indicates little benefit from focusing solely on earlywood density when selecting for wood density. Analysing earlywood and latewood separately does not benefit from including the width of the corresponding ring section as a covariate. Juvenile wood may possibly turn into mature wood 15–20 y from the pith.

     

Age trends in the genetic parameters of wood density and the relationship with growth rates in hybrid larch (Larix gmelinii var. japonica × L. kaempferi) F1

Age trends in variance components and heritability of overall wood density, earlywood and latewood density, and latewood proportion were investigated in 29-year-old trees of 19 full-sib families of hybrid larch (Larix gmelinii var. japonica × Larix kaempferi) F₁. The age–age correlation and optimum selection age for these traits were also estimated and genetic and phenotypic correlations between wood density and radial growth rate were calculated for each growth ring. Intraring wood density data were obtained using X-ray densitometry. The coefficient of additive genetic variance was stable over all ages, whereas the coefficient of environmental variances gradually decreased with increasing age, resulting in increases in heritability estimates with age for overall density. The latewood proportion had the highest heritability estimates at all ages, ranging from 0.44 to 0.66. Overall density and its various components at 28 years of age showed strong genetic correlations with their respective traits at all younger ages. Optimum selection ages for the wood density traits ranged from 8 to 14 years, at which point maximum gain efficiencies per year were obtained. There were negative correlations between wood density and radial growth rate at early ages, although these relationships tended to be weaker with increasing age. These results suggest that selection at a young age is effective for wood density, but particular care must be taken in selecting trees with an improved radial growth rate because rapid growth will result in a low-density wood product, especially in the early growth period.     

Genetic parameters of growth and wood quality traits in Picea abies

Genetic parameters were estimated for wood and growth traits in two 19-yr-old clonal trials and a 40-yr-old full-sib progeny trial of Norway spruce [Picea abies (L.) Karst.]. In the clonal trials high (>0.4) broad-sense heritabilities were found for wood density traits, lignin content, number of internal cracks, growth traits, spiral grain and number of resin canals. Moderate (0.2–0.4) heritabilities were found for tracheid lumen diameter and cell wall thickness, microfibril angle and tracheid length, while low heritabilities (<0.2) were found for pulp yield, fibre strength, wood stiffness and wood colour. Lignin content and pulp yield showed low genetic variation, whereas the genotypic coefficient of variation for most other traits ranged between 5 and 15%. Most traits showed low levels of genotype by environment interaction. Among the wood properties, latewood proportion, earlywood density and ring density showed significant, adverse correlations with volume in both clonal trials.     

Quantitative analysis of lignin using an UV microscopic image analyser. variation within one growth increment

Summary The variation of the lignin content within one growth increment of Abies sachalinensis was investigated with the aid of an ultraviolet microscopic image analyser. The lignin content was determined continuously in each cell within a growth increment. The direct photometric scanning method of UV image is believed to give accurate results for determination of the lignin content of the cell wall, since it contains fewer assumptions. The lignin content of the earlywood was higher than that of latewood in adult wood as same as the others. It was high, however, in the terminal zone of the latewood. The trends of the juvenile wood were quite different from those of the adult wood. The lignin content increased from earlywood to latewood.This paper was presented at the Wood Anatomy Congress of IAWA, Aug. 27, 1979, Amsterdam. The authors wish to express their gratitude to Prof. Dr. J. Bauch for helpful advice     

Does selecting for improved growth affect wood quality of Pinus pinaster in Portugal?

The selection criteria for the first generations in the Portuguese Pinus pinaster improvement program have been the growth rate and form traits. In this work we study the consequences of this selection on wood quality traits. This study assesses genetic and phenotypic correlation between growth, wood density components, lignin content and mechanical traits (radial modulus of elasticity and radial modulus of rupture) of 46 half sib families from a progeny trail located in Leiria, Portugal, originated from seed collected in a clonal seed orchard. A total of 552 seventeen-years-old trees (about half of full rotation age) were sampled at 2 m height. Height measured at 12 years old presented a higher genetic control (h² = 0.34) relatively to DBH, measured at 12 and 17 years old respectively (h² = 0.17 and h² = 0.15). The results of this study also showed that DBH growth is more dependent on latewood components than earlywood components and that higher growth in Mediterranean regions can be due to an increase of the period of latewood formation. Further, we can conclude genetic selection based on growth will not result in a decrease of wood density, will not affect the occurrence of spiral grain, and is possible to obtain an increase in the radial modulus of elasticity. The present study also showed that it is possible to select for increased growth with lower lignin proportion. Results also suggest that selection for growth at 12 years will probably not affect negatively the wood properties at 17 years.     

日本落叶松无性系木材性质的遗传变异

对10个10年生日本落叶松无性系的木材基本密度、管胞参数进行了测定.结果表明:木材基本密度,早、晚材管胞宽度和早材长宽比无性系间差异显著;木材基本密度,早、晚材管胞长度,早晚材管胞宽度和早晚材长宽比径向变异模式相似,即从髓心向外以曲线形式不断增加,有时亦有起伏;早材从髓心向外以近似直线的形式缓慢增加,晚材从髓心向外以曲线形式增加,初期增加幅度较大,到一定年龄后趋于水平变化并略有波动;材质性状与树木年轮间的关系以对数方程、幂函数方程、指数方程拟合效果较好;除了晚材壁腔比和早材壁厚外,其它木材性质的重复力均在0.5以上,受中度或中度以上的遗传制约,按照20%的选择率,长宽比和晚材管胞长能获得较高的遗传增益.     

Genetic correlations between wood quality traits of <Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait

• ? It is essential to understand how characteristics are related to each other in breeding programmes to select wood properties, in order to avoid that, in selecting for one trait, we are negatively affecting another. Moreover, measuring wood properties is time consuming and expensive.
• ? This study assesses genetic and phenotypic correlations between wood density components and spiral grain of 46 half-sib families of Pinus pinaster in seventeen-year-old trees.
• ? Results showed that genetic correlations for all wood density components were higher than corresponding phenotypic correlations. Furthermore, all wood density components were highly genetically correlated with ring density, and also closely associated among themselves. Results showed a higher genetic correlation of ring density with earlywood density (r _g = 0.96) than with latewood density r _g = 0.79). A moderate to high positive genetic correlation was found between spiral grain and wood density characteristics (0.29–0.61).
• ? We conclude that ring density (overall wood density) can be improved by increasing either earlywood density, latewood percent, or both of these traits, and spiral grain can be modify in future plantations.

     

The effects of within-species and between-species variation in wood density on the photodegradation depth profiles of sugi (Cryptomeria japonica) and hinoki (Chamaecyparis obtusa)

Low density wood is more rapidly eroded than denser wood when exposed to the weather, possibly because it is more susceptible to photodegradation. Fourier transform infrared microscopy was used to examine: (1) the depth of photodegradation in earlywood and latewood of sugi (Japanese cedar) and earlywood of hinoki (Japanese cypress) exposed for up to 1500 h to artificial sunlight emitted by a xenon lamp (375 W/m² within the 300 to 700 nm spectral range); and (2) the relationship between the density of wood tissues and depth of photodegradation. The depth of photodegradation varied between species (sugi and hinoki) as well as within a growth ring (sugi earlywood and latewood), and there was an inversely proportional relationship between depth of photodegradation and wood density. These findings may explain why low density earlywood is more rapidly eroded than latewood during weathering, and more generally, why there is an inverse relationship between the density of wood species and their rate of erosion during artificial and natural weathering. Part of this work was presented at the 54th Annual Meeting of the Japan Wood Research Society, Sapporo, August 2004     

The effects of drying on the structure and permeability of the wood of Abies grandis

Summary The effects of air-drying and solvent-drying on the sapwood of Abies grandis have been investigated by a new method for the determination of the size and number of conducting tracheid lumina and pit membrane pores which involves the measurement of gaseous permeability at various mean pressures. Both earlywood and latewood tracheids (83% of the total) were found to be conducting in solvent-dried wood, but in air-dried wood only latewood tracheids (32% of the total) were conducting. In solvent-dried wood there were on average 27,000 pit membrane pores per conducting tracheid compared with only 600 in air-dried wood. In both, the average pit membrane pore radius was about 0.1 m.Liquid permeabilities have been predicted from the calculated radii and numbers. The liquid permeability of solvent-dried wood was 31 times greater than that of air-dried wood in which the lumina were responsible for 13% of the total resistance to flow. The lumina were responsible for 39% of the resistance in solvent-dried wood and it is suggested that in first-formed earlywood the lumina may cause more than half the total resistance.A new method is described for the cleaning of direct carbon replicas of wood. In this the cellulose is removed by cellulase instead of sulphuric acid, and no wax backing is required. This provides much cleaner replicas. Electron micrographs have been obtained of both earlywood and latewood dried by the two methods.The authors wish to thank Mr. A. R. Sayers for preparing the computer programme used in this work, Dr. R. Ph. C. Johnson for his help and advice regarding the electron microscopy and Professors Matthews and Weatherley for their advice and encouragement.     

Variation in wood density and ring width in Acacia melanoxylon at four sites in Portugal

The radial variation in wood density in Acacia melanoxylon R. Br. was studied using microdensitometry by sampling 20 trees with a 40-cm diameter class at four sites in Portugal. The measurements were taken from pith to bark at breast height. A. melanoxylon had an average ring density of 0.607 g cm^?3, ranging from 0.556 to 0.630 g cm^?3. The mean growth was 6.0 mm year^?1. Latewood corresponded, on average, to 34 % of the ring width. Between-tree variability at each site was the main source of variation in the density components, representing between 30 and 54 % of the total variation. Between-sites variability represented from 0 to 21 % of the total variation in density components. The environmental effects (site related) were more pronounced on latewood, while the genetic effect (tree related) was more evident in earlywood. Ring width, latewood percentage and heterogeneity index were independent from site, trees in site and age effects. The values of wood density and radial growth revealed that A. melanoxylon can be important as a commercial timber species in Portugal.     

Variations and correlations of various ring width and ring density features in European oak: implications in dendroclimatology

Summary Variations and correlations of various ring width and ring density features were analyzed in 18 European oak (Quercus petraea andQuercus robur) trees from northeastern France. In light of these analyses, the goodness of various tree-ring features as a climatic parameter was discussed. In general, ring density features (viz. earlywood density, latewood density, average ring density, minimum density and maximum density) show a stronger response to calendar year, a comprehensive climatic variable, than ring width features (viz. earlywood width, latewood width, total ring width and latewood percent). The response of latewood features (viz. latewood width, latewood density and maximum latewood density) is stronger than that of earlywood features (viz. earlywood width, earlywood density and minimum earlywood density). Average ring density seems to be the most sensible tree-ring feature in European oak in terms of the response to calendar year. Moreover, total ring width as a climatic parameter is not as good as latewood width, and maximum (latewood) density and minimum (earlywood) density appears not to contain as much climatic information as (average) latewood density and (average) earlywood density, respectively.This project was supported by the institut National de la Recherche Agromonique (INRA). Thanks are due to Dr. G. Nepveu for his constant encouragement and assistance during this study     

Effect of age on the variation, correlations and inheritance of selected wood characteristics in black spruce (Picea mariana)

Summary The variation, correlations and inheritance of various intraring wood characteristics in 15-year-old black spruce (Picea mariana) trees from 40 half-sib families grown in New Brunswick were examined. The emphasis was placed on the effect of age on these genetic parameters. With increasing (cambial) age, ring width and ring density tend to exhibit a constant tree-to-tree variation whereas most other characteristics exhibit a smaller tree-to-tree variation. The heritability for latewood characteristics does not seem to change appreciably. Other characteristics (except RDmi), however, tend to be less heritable when the tree grows older. Correlations between ring density and most intraring characteristics tend to be weaker with increasing age. This implies that fast-growth in this species will have less negative effect on wood density when the tree grows older.The author wishes to thank Dr. E.K. Morgenstern of the Genetic Resource Consultants, Permbroke, Ontario and Mr. D. Simpson of the Canadian Forest Service in Fredericton, New Brunswick for their involvement in the initial planning of this project. Thanks are also due to G. Chauret, T. Keenam, R. Ploure, V. Steel and C. Reitlingshoefer for their technical assistance.     

Interpretation of X-ray diffractograms of wood for assessments of microfibril angles in fibre cell walls

Summary Graphical models have been developed to represent X-ray diffraction patterns for microfibril arrangements in each of the characteristic secondary wall layers of fibres in normal earlywood, latewood, compression wood, and tension wood. Models for usual combinations of typical layers S₁, S₂, and S₃, and for complex tissues including more than one S₂ layer class indicate a basis for a new analytical technique for diffractograms.Diffractograms of tissues from earlywood or latewood zones may involve effects of three to four S₂ layer variations, possibly including tension wood or compression wood. The new technique enables assessment of the microfibril angle for each. Corresponding probable experimental errors are considered. Thus it is demonstrated that, even without direct calibration by other methods for measuring microfibril angle, realistic comparative values may be obtained for all S₂ layer classes substantially represented. Such data constitute significantly more reliable indices of actual values than those provided by other techniques. Also, the data give qualitative information on other aspects of the variability of fibre types within each specimen.     

Control of tracheid cross-sectional dimensions in Norway spruce and Scots pine wood raw material

Abstract

Wood properties, including tracheid cross-sectional dimensions, show a large degree of variation. To improve the properties of products made from wood, different methods to control variation have been developed. This study aims to determine the theoretical efficiency of three control strategies: the fractionation of pulped tracheids into earlywood and latewood, the separation of juvenile and mature wood, and sorting of logs according to tree size. The efficiency of each method was studied by first constructing virtual trees from measured tracheid cross-sectional dimensions, then simulating the efficiency of above-mentioned methods. The tracheid dimension data include Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.). The simulations show that separation into earlywood and latewood classes has the highest theoretical efficiency and yields the lowest variances in raw material. Classification into juvenile and mature wood groups is the second most efficient method, and the sorting of logs according to the size class of the tree is the least efficient method. It was also concluded that the variation in cell-wall thickness and radial diameter mainly originates from differences between earlywood and latewood, whereas the variation in tangential diameter mainly originates from differences between mature and juvenile wood.     

High negative genetic correlations between growth traits and wood properties suggest incorporating multiple traits selection including economic weights for the future Scots pine breeding programs

Context

The development of multiple trait selection indices for solid (structure) wood production in the Scots pine (Pinus sylvestris L.) breeding program requires genetic variances and covariances estimated among wood quality traits including stiffness.

Aims

Genetic control and relationships among Scots pine growth, fiber, and wood quality traits were assessed by estimating heritability, phenotypic and genetic correlation using a Scots pine full-sib family trial.

Method

Wood quality traits including clearwood and dynamic acoustic stiffness were measured using SilviScan and Hitman in a 40-year-old progeny trial and by sampling increment cores of 778 trees of 120 families. Genetic parameters were estimated using the mixed model by the ASReml software.

Results

Heritability ranged from 0.147 to 0.306 for growth, earlywood, transition wood and latewood proportion traits and from 0.260 to 0.524 for fiber dimension, wood density, MFA and stiffness traits. The highly unfavorable genetic correlation between diameter and whole core density (?0.479) and clearwood stiffness (?0.506) and dynamic acoustic stiffness (?0.382) was observed in this study.

Conclusion

The unfavorable genetic correlations between growth traits and stiffness indicate that multiple traits selection using optimal economic weights and optimal breeding strategies are recommended for the advanced Scots pine breeding program.     

Influence of initial planting spacing and genotype on microfibril angle,wood density,fibre properties and modulus of elasticity in Pinus radiata D. Don corewood

Pinus radiata D. Don trees from six clones, grown at initial spacings of 2500 stems ha⁻¹ and 833 stems ha⁻¹ were destructively harvested. For these trees wood properties were measured on radial slices sampled at a height of 1.4 m above the ground. Relative to wide spacing, close initial stand spacing significantly reduced microfibril angle (MFA) and ring width and significantly increased dynamic modulus of elasticity (MOE), fibre length, latewood percentage and cell wall thickness. Density and fibre width were not significantly different between spacing treatments. Examination of the influence of genetic population on wood properties indicated that genotype significantly influenced MFA, MOE and ring width. The key wood properties MFA, MOE and fibre length were regressed against tree diameter, height and stem slenderness. All three wood properties were most strongly correlated with stem slenderness. Multiple regression models developed for MFA, MOE and ring width accounted for respectively 62%, 81% and 58% of the variation in these variables. The following changes occurred in sampled properties with increasing ring number: MFA and ring width declined markedly; MOE and fibre length increased markedly; latewood percentage and cell wall thickness increased slightly; and density and fibre width did not show any radial trend.     

Wood density of three oak species in Tennessee, 1940–45 and 1970–75

Three oak species from East Tennessee were core-sampled to determine whether the density of wood produced in a pre-pollution period (1940–45) was different from that of a pollution period (1970–75) using gamma-densitometry. Component densities of average earlywood, average latewood, minimum earlywood, maximum latewood and average periodic density for Quercus alba L. and Quercus prinus L. were significantly greater for 1940–45 than for 1970–75. Differences between the two time periods for mean earlywood (P = 0.05), maximum latewood and mean periodic densities were found for Q. velutina.     

Ring characteristics and screw withdrawal resistance of naturally regenerated <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> var. <Emphasis Type="Italic">formosana</Emphasis> trees

The ring characteristics and screw withdrawal resistance (SWR) of naturally regenerated Taiwan yellow cypress (Chamaecyparis obtusa var. formosana) trees were explored. Significant differences in average ring width (RW), earlywood width, latewood width, ring density (RD), earlywood density (ED), latewood density (LD), highest density (D_max), lowest density (D_min), latewood percentage (LWP), and SWR were observed between trees, rings (SWR excluded), and tree height positions. The RW components in the radial direction increased from the pith outward to about the 3rd to 5th ring and then decreased to about the 25th ring; it was almost constantly sustained toward the bark side. The RD in the radial direction slowly decreased from the pith outward to the bark side. Average ring width and ring density were significantly affected by the various tree growth rates, radial ring numbers, and tree height positions. ED, LD, D_max, D_min, and LWP were the most important factors determining the overall RD. RW did not correlate with tree RD. SWR is correlated with ED, RD, D_min, LWP, and intra-ring density variation (IDV). Thus, the SWR can be used to predict wood density and in nondestructive evaluation of a living tree.