Genetic variation and relationships to growth traits for microfibril angle,wood density and modulus of elasticity in a Picea abies clonal trial in southern Sweden

Abstract

Genetic variation in wood density, microfibril angle (MFA), wood stiffness (MOE), height, diameter and volume was investigated in a 26-year-old Norway spruce [(Picea abies (L.) Karst.] clonal trial in southern Sweden. Wood quality measurements were performed on 10 mm increment cores using SilviScan. For MFA, mean values of annual rings showed the highest value (30°) at ring 2 counting from the pith, followed by a steep decrease and a gradual stabilization around ring 12 at approximately 14°. MOE showed a monotonic increase from 5 GPa to 14 GPa when moving from pith to bark. High broad-sense heritability values were found for wood density (0.48), MFA (0.41) and MOE (0.50). All growth traits displayed heritability values of similar magnitudes as reported in earlier studies. The generally high age–age correlations between different sections of the wood cores suggested that early selection for wood quality traits would be successful. Owing to unfavorable genetic correlations between volume and MOE, the correlated response indicated that selection for volume only at age 10 would result in a 0.27% decrease in weighted MOE at age 26 for every 1% increase in volume.     

Radial and between-family variations of the microfibril angle and the relationships with bending properties in Picea jezoensis families

With emphasis on tree breeding for wood quality in Picea jezoensis, we aimed to evaluate radial and between-family variations in the microfibril angle (MFA) of the S₂ layer in the latewood tracheids in 10 open-pollinated families of 43-year-old P. jezoensis trees. In addition, the relationships between MFA/wood density with the modulus of elasticity (MOE) or modulus of rupture (MOR) were investigated. Significant differences in MFA between families were found from the pith toward the bark. MFA showed higher values around the pith area, although some families showed relatively lower values than others around this area. In addition, due to a larger coefficient of variations of MFA near the pith, the potential for juvenile wood MFA improvement may be greater compared with mature wood. MOE was correlated with MFA in juvenile wood and with wood density in mature wood, whereas MOR was mainly correlated with wood density at radial positions in both woods. Therefore, to improve the MOE and MOR of P. jezoensis wood, both MFA and wood density would be factors to consider in both juvenile and mature woods. On the other hand, there are indications that, only wood density would be an important criterion for improving mature wood properties.     

Variation in wood properties among five full-sib families of Norway spruce (Picea abies)

Genetic- and environmental variation and correlation patterns were characterized for modulus of elasticity (MOE), modulus of rupture (MOR) and related wood traits: latewood proportion, wood density, spiral grain, microfibril angle and lignin content in five full-sib families of Norway spruce. The families were evaluated on the basis of clearwood specimens from the juvenile -mature wood transition zone of 93 sampled trees at age 30 year from seed. Family-means varied significantly (p < 0.05) for all wood traits studied except lignin content. MOE varied between 7.9–14.1 GPa among trees and 9.4–11.0 GPa among families. MOR varied between 47–87 MPa among trees and 61–71 MPa among families. Families remained significantly different in an analysis of specific MOE (MOE/density) and MOR (MOR/density). Hence, solely relying on wood density as a wood quality trait in tree breeding would not fully yield the potential genetic gain for MOE and MOR. Correlations between wood structural traits and specific MOE and MOR are presented and discussed.     

Provenance variation in growth and wood properties of juvenile Cyclocarya paliurus

Cyclocarya paliurus is a highly valued and multiple function tree species. There has been increasing interest in planting and managing C. paliurus for timber production and medical use owing to loss of harvestable acreage. Seed from six provenances was collected from the main natural range of this species. Significant variation in growth and wood properties was measured among the six provenances at age 7 years. Provenance mean height and DBH varied significantly from 730–991 and 6.7–10.0 cm, whereas provenance means of wood basic density and crystallinity ranged from 463–554 kg m^?3 and 51.4–74.1 %, respectively. Mean provenance microfibril angle (MFA) at breast height ranged from 18.1° to 23.2°, while MFA at breast height varied from 11.0° to 34.5° among growth rings which showed a consistent pith-to-bark trend of declining angles. There was no significant relationship of growth rate with latitude or longitude of seed sources, however, provenances from low latitude and longitude grew faster at the trial site. Wood quality was significantly related to latitude of seed sources, showing a positive correlation for both wood basic density and wood crystallinity, but a highly negative association with MFA. Significant correlations between wood properties measured indicated that there exists a great opportunity to improve wood quality of C. paliurus through selection of juvenile trees with low MFA.     

Optimal timing of early genetic selection for sawn timber traits in <Emphasis Type="Italic">Picea abies</Emphasis>

In breeding Norway spruce, selection for improved growth and survival is performed at age 10–15 years in order to optimize genetic gain per year. We investigated whether a selection based on wood traits such as density and grain angle, measured under bark in the field at the same age would be informative enough with respect to structural quality traits of sawn boards. To achieve this objective, a sawing study was conducted on the butt logs of 401 trees from a 34-year-old Norway spruce progeny trial situated in southern Sweden. Stem discs were excised from the top of the logs and radial profile data of grain angle, and wood density was recorded for specific annual rings. The sawn and dried boards were assessed for structural traits such as twist, board density, bending stiffness (static modulus of elasticity, sMoE) and bending strength (modulus of rupture, MoR). Additive genetic correlations (r_a) between single annual ring density measurements and board density, sMoE and MoR were consistently strong (r_a>?0.7) for annual rings 5–13. Genetic correlations of similar magnitude between grain angle and board twist were estimated for all investigated annual rings (from 2 to around 26 under bark). Consequently, it was found that indirect selection for wood density and grain angle at the tree age 10–16 years would result in more genetic gain per year than selection at later ages. This makes it feasible to perform simultaneous selection of progeny in the field for both growth and wood traits at similar ages.     

Prediction of wood stiffness, strength, and shrinkage in juvenile wood of radiata pine

Development of optimal ways to predict juvenile wood stiffness, strength, and stability using wood properties that can be measured with relative ease and low cost is a priority for tree breeding and silviculture. Wood static modulus of elasticity (MOE), modulus of rupture (MOR), radial, tangential, and longitudinal shrinkage (RS, TS, LS), wood density (DEN), sound wave velocity (SWV), spiral grain (SLG), and microfibril angle (MFA) were measured on juvenile wood samples from lower stem sections in two radiata pine test plantations. Variation between inner (rings 1–2 from pith) and outer (rings 3–6 from pith) rings was generally larger than that among trees. MOE and MOR were lower (50%) in inner-rings than in outer-rings. RS and TS were higher (30–50%) for outer-rings than inner-rings, but LS decreased rapidly (>200%) from inner-rings to outer-rings. DEN had a higher correlation with MOR than with MOE, while MFA had a higher correlation with dry wood MOE than with MOR. SLG had higher significant correlation with MOE than with MOR. DEN and MOE had a weak, significant linear relationship with RS and TS, while MOE had a strong negative non-linear relationship with LS. Multiple regressions had a good potential as a method for predicting billet stiffness (R ² > 0.42), but had only a weak potential to predict wood strength and shrinkage (R ² < 0.22). For wood stiffness acoustic velocity measurements seemed to be the most practical, and for wood strength and stability acoustic velocity plus core density seemed to be the most practical measurements for predicting lower stem average in young trees.     

The variation of microfibril angle in South African grown Pinus patula and its influence on the stiffness of structural lumber

Reduction in the rotation ages of softwood saw-log plantations in South Africa is causing increased proportions of low stiffness sawn lumber at final harvest. It has been shown for some species that the microfibril angle (MFA) of the S2 layer of tracheids is strongly related to the modulus of elasticity (MOE) of wood, even more so than wood density, especially in wood formed during juvenile growth. The objectives of this study were to describe the variation in MFA in young Pinus patula trees and to determine the relationship between MFA and the dynamic MOE of sawn P. patula lumber. Thirty 16- to 20-year-old trees from six compartments from the Mpumalanga escarpment were processed into discs and lumber. The MFA, density and ring width were measured at two height levels using Silviscan 3. The average annual ring MFA varied between 7° and 29°; the pattern of variation depended mainly on height level and the ring number from the pith. The MFA in P. patula followed the same within-tree variation trends as in New Zealand-grown Pinus radiata but the average MFA was lower in absolute terms and differences between height levels were less pronounced. The MFA and density exhibited highly significant Pearson correlations of 0.73 and 0.70, respectively, with board dynamic MOE. A multiple regression model, which included MFA, density and ring width, explained 71% of the variation in the dynamic MOE of boards. A sensitivity analysis on the model showed that MFA and density had approximately similar influences on predicting the dynamic MOE of Pinus patula boards.     

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.     

马尾松无性系种子园半同胞子代变异分析和家系选择

以福建省漳平五一林场马尾松无性系种子园自由授粉子代半同胞家系为研究对象,分析其树高、胸径、材积、冠幅、通直度、高径比、冠径比、木材基本密度、管胞长度和宽度、管胞长宽比等性状,发现家系间除管胞长宽比性状差异显著,其余性状均达差异极显著,暗示着马尾松种子园半同胞家系间存在较为丰富的变异,具选育潜力。8年生林分内各性状的家系遗传力以树高性状为最高(0·691),其次是木材基本密度(0·640)。各性状间的相关性分析结果表明,生长量性状的改良可间接改善通直度,且生长量性状与木材基本密度存在显著正相关关系,木材基本密度与管胞长度和宽度不相关。采用10%入选率,发现5年生林分和8年生林分入选家系一致,材积遗传增益分别达19·74%和19·23%,且5年生林分和8年生林分生长量性状的遗传相关极显著,认为马尾松制浆造纸材短轮伐期的初选年龄可确定为5年生;根据性状遗传力的相对大小和典型相关分析的结果,认为马尾松家系选择可适当注重树高生长量指标。以树高和材积为选择指标,按照10%入选率,兼顾材性和种子园内无性系的开花结实情况,筛选出制浆造纸材短轮伐期优良家系12个。     

Genetic Correlations Among Wood,Growth Capacity and Stem Traits in Pinus sylvestris

Wood density and tracheid length are two traits that significantly affect wood products. Genetic correlations were estimated to evaluate the effect on these traits from a selection for traits included in the Swedish Pinus sylvestris L. breeding programme. Measurements from a non-contiguous single-tree plot progeny trial with controlled matings between 30 parent trees was used. Heritabilities were high for the wood traits, intermediate for the growth capacity traits and low for the stem traits, with the exception of branch angle. Wood density showed no or non-significant negative genetic correlations with the growth capacity traits and a positive correlation with relative branch diameter. Tracheid length showed positive genetic correlations with height and a positive correlation with relative branch diameter. A selection that increased height growth at the age of 13 yrs by 10% was expected to decrease mean wood density at 33 yrs by 1%. The expected correlated response to tracheid length from the same selection was a 3% increase in the juvenile and mature wood.     

Wood properties and their among-family variations in 10 open-pollinated families of Picea jezoensis

This study aimed to evaluate radial and among-family variations of wood properties in Picea jezoensis. A total of 174 trees were randomly selected from 10 open-pollinated families in a progeny trial for measuring stem diameter, dynamic Young’s modulus of log (DMOE_log), annual ring width (ARW), air-dry density (AD), modulus of elasticity (MOE), and modulus of rupture (MOR). Mean values of DMOE_log, AD, MOE, and MOR were 9.60 GPa, 0.41 g/cm³, 9.44 GPa, and 76.6 MPa, respectively. Significant differences among families were observed in all properties. F values obtained by analyzing variance in wood properties were higher than those generally observed in growth traits. In addition, F values in wood properties remained relatively higher from the 1st to 25th annual ring from the pith, although F value in ARW rapidly decreased with each increase in annual ring number. These results indicate that genetic factors largely contributed to the variance in wood properties compared with the growth traits.     

Index selection for growth and construction wood properties in Pinus elliottii open-pollinated families in southern China

The initial introduction of Pinus elliottii (PEE) to China occurred in the 1930s, and the planting of this conifer species has now attained close to 3 million ha in the subtropical zone of southern China. A large-scale genetic improvement program for PEE was implemented in southern China to produce fast-growing trees with high wood quality to address the severe shortage of timber production over the last two decades. In this paper, selection for stem volume, basic wood density (DEN) and modulus of elasticity (MOE) was based on the Smith–Hazel index, and a total of approximately 2 000 individual trees from 158 PEE open-pollinated families were selected at 22 years of age. The DEN and MOE for each tree were determined by non-destructive evaluation techniques using the Pilodyn and Hitman Director ST300® acoustic velocity device. The heritabilities and genetic and phenotypic correlations for the traits that were measured were estimated using the residual maximum likelihood approach in the flexible mixed modelling program ASReml-R. The results showed that the heritability estimates for the wood properties were between 0.292 and 0.309, and the heritabilities of the growth traits ranged from 0.129 to 0.216. The genetic correlation between the DEN_P and acoustic velocity (V?) with MOE_P was 0.45 and 0.95, respectively. An indirect selection based on V was observed to be highly effective for determination of MOE. It indicated that V can be integrated into tree improvement programs as a useful index of MOE by ranking candidate families or individuals within the selection population. The genetic correlations between the growth traits and wood properties were not significant. By contrast, the phenotypic correlations between them were significantly positive, but the correlation coefficients were very low. The appropriate selection index (I₄), which placed 10 times as much weight on DEN and MOE as the equal emphsis method, was determined as the appropriate selection index.     

人工林杉木和杨树木材物理力学性质的株内变异研究

按照中国国家标准研究杉木和I-214杨树木材的抗弯弹性模量、抗弯强度、顺纹抗压强度和密度,同时按照日本国家标准研究2个树种的顺纹抗剪强度.结果表明:杉木的抗弯强度、顺纹抗压强度和密度由胸高直径处向上呈波浪形增加,抗弯弹性模量则稳定降低,但不同高度间杉木的物理力学性质没有显著差异;近树皮处木材的物理力学性质高于近髓心处木材,并有极显著差异.对于I-214杨树,只有抗弯弹性模量从髓心到树皮逐渐增加,其他的物理力学性质,最小值在从髓心到树皮的过渡区,最大值在近树皮处,从髓心到树皮,杨树的物理力学性质有极显著的差异.杉木和杨树的径面顺纹抗剪强度从髓心到树皮有极显著差异,并且近树皮的高于近髓心的木材,而弦面顺纹抗剪强度从髓心到树皮没有显著差异.木材密度与力学性质有很好的线性相关关系,木材密度是一个很好的力学强度的预测手段.     

Variation analysis and fine family selection on half-sib progenies from clonal seed orchard of Pinus massoniana

Variation of traits that include height, diameter at breast height (DBH), stem volume, crown width, straightness, wood basic density, tracheid length and width of open-pollinated progenies from clonal seed orchard of Pinus massoniana was analyzed. Differences in traits such as growth, tree form and wood quality among families was significant, which indicated that the improvement potential was notable. Heritability of height and wood basic density reached 0.691 and 0.640, respectively. According to a correlation analysis between traits, stem straightness could be improved indirectly when growth traits are improved; growth traits were significantly correlated with wood basic density; and wood basic density was not significantly correlated with length and width of the tracheid. Under the selection ratio of 10%, genetic gains were 19.74% and 19.32% respectively, and selected families from the progeny test stands at the age of five years and eight years were the same. Genetic correlation of height, DBH and stem volume was also significant at the level of 0.01. These show that selection of the short rotation of P. massoniana for pulp and paper use would be efficient for five-year-old trees. According to the comparison of the heritability of all the traits and taking into account the result of canonical correlation analysis, height could be considered as the most important index for selection. Given wood quality of all the families, and flowering and fruit settings of the clones in a seed orchard, twelve fine families were selected at 10% selection ratio of height growth and stem volume. Translated from Scientia Silvae Sinicae, 2005, 41(6): 43–49 [译自: 林业科学]     

Genetic correlations among juvenile wood quality and growth traits and implications for selection strategy in Pinus radiata D. Don

? Juvenile wood quality in Pinus radiata is affected by factors such as low density, stiffness, and high microfibril angle, spiral grain, and shrinkage. Adverse genetic correlations between growth and wood quality traits remain as one of the main constraints in radiata pine advanced generation selection breeding program.

? Juvenile wood property data for this study were available from two progeny tests aged 7 and 6 y. We estimated the genetic correlations between stiffness, density, microfibril angle, spiral grain, shrinkage in the juvenile core and DBH growth in radiata pine, and) to evaluated various selection scenarios to deal with multiple objective traits.

? Negative genetic correlations were found for modulus of elasticity (MoE) and density with microfibril angle, spiral grain, shrinkage, and DBH. We observed low to moderate unfavourable genetic correlations between all wood quality traits and DBH growth.

? These low to moderate genetic correlations suggest that there may be some genotypes which have high DBH growth performance while also having high wood stiffness and density, and that the adverse correlation between DBH and MoE may not entirely prohibit the improvement of both traits. Results indicate that, in the short term, the optimal strategy is index selection using economic weights for breeding objective traits (MAI and stiffness) in radiata pine.

? In the long-term, simultaneously purging of the adverse genetic correlation and optimizing index selection may be the best selection strategy in multiple-trait selection breeding programs with adverse genetic correlations.

     

Models for predicting microfibril angle variation in Scots pine

Context

Microfibril angle (MFA) is one of the key determinants of solid timber performance due to its strong influence on the stiffness, strength, shrinkage properties and dimensional stability of wood.

Aims

The aim of this study was to develop a model for predicting MFA variation in plantation-grown Scots pine (Pinus sylvestris L). A specific objective was to quantify the additional influence of growth rate on the radial variation in MFA.

Methods

Twenty-three trees were sampled from four mature Scots pine stands in Scotland, UK. Pith-to-bark MFA profiles were obtained on 69 radial samples using scanning X-ray diffractometry. A nonlinear mixed-effects model based on a modified Michaelis–Menten equation was developed using cambial age and annual ring width as explanatory variables.

Results

The largest source of variation in MFA (>90 %) was within trees, while between-tree variation represented just 7 % of the total. Microfibril angle decreased rapidly near the pith before reaching stable values in later annual rings. The effect of ring width on MFA was greater at higher cambial ages.

Conclusion

A large proportion of the variation in MFA was explained by the fixed effects of cambial age and annual ring width. The final model is intended for integration into growth, yield and wood quality simulation systems.     

Genetic improvement of wood density and radial growth in Larix kaempferi: results from a diallel mating test

Context

Wood density is an important component of wood quality, and it is therefore important to assess whether it can be subject to genetic improvement.

Aim and methods

We assessed the potential for genetic improvement of wood density in Larix kaempferi by recording components of annual growth rings. A full diallel mating test based on six plus L. kaempferi trees was used. Trees were 29 years old. Wood density was recorded by soft X-ray densitometry, and genetic parameters and genetic gains were computed.

Results

Wood density of mature wood was highly heritable, and the largest heritability (0.78) was reached at age 25. Specific combining ability and reciprocal effects displayed very low variance. The age–age correlation of overall wood density was very high (>0.94). The genetic correlation between overall wood density and basal cross-sectional area was positive after age 10. Early selection at age 6 would account for 69 % of the genetic gain from direct selection at age 28 in terms of wood density.

Conclusion

Genetic improvement of wood density could be achieved by mass selection and a simultaneous selection for radial increment in L. kaempferi; early selection for wood density can be achieved in this species.     

Fresh-wood bending: linking the mechanical and growth properties of a Norway spruce stem

To provide data and methods for analyzing stem mechanics, we investigated bending, density and growth characteristics of 207 specimens of fresh wood from different heights and radial positions of the stem of one mature Norway spruce (Picea abies L. Karst.) tree. From the shape of each stress-strain curve, which was calculated from bending tests that accounted for shear deformation, we determined the modulus of elasticity (MOE), the modulus of rupture (MOR), the completeness of the material, an idealized stress-strain curve and the work involved in bending. In general, all mechanical properties increased with distance from the pith, with values in the ranges of 5.7-18 GPa for MOE, 23-90 MPa for MOR and 370-630 and 430-1100 kg m(-3) for dry and fresh wood densities, respectively. The first three properties generally decreased with stem height, whereas fresh wood density increased. Multiple regression equations were calculated, relating MOR, MOE and dry wood density to growth properties. We applied these equations to the growth of the entire stem and considered the annual rings as superimposed cylindrical shells, resulting in stem-section values of MOE, MOR and dry and fresh densities as a function of stem height and cambial age. The standing tree exhibits an inner stem structure that is well designed for bending, especially at a mature stage.     

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.     

尾叶桉第1代与第2代群体间生长性状和木材材性的遗传控制对比研究

利用4个尾叶桉（Eucalyptus urophylla）自由授粉子代试验（其中D46、D54 2个试验为第1代子代试验,D135、SE35 2个试验为第2代子代试验）作为研究对象,对其生长性状和木材材性进行遗传参数估算.研究结果表明,生长性状和树皮性状受到较低的遗传控制,其狭义遗传力(h2):胸径(DBH) 0.0...