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.     

Root hydraulic conductivity and root growth capacity of black spruce (Picea mariana) seedlings

The relationship between root elongation and root hydraulic conductivity was investigated in 1-year-old, overwintered black spruce (Picea mariana (Mill.) seedlings. Hydraulic conductivity was estimated by observing water flux through decapitated roots under positive pressure. Five hydraulic conductivity parameters were estimated: (1) water flux under a minimal pressure of 0.2 MPa (J(v, min)); (2) pressure at which a linear relationship between water flux and pressure began (P(min)); (3) slope of the linear water flux-pressure relationship (L(v)); (4) pressure at which the linear relationship between water flux and pressure ended (P(max)); and (5) maximum water flux (J(v, max)). Between day 1 and day 2 after thawing of the growing medium, there were significant increases in L(v) and J(v, max) but there was little, if any, root elongation. Root elongation averaged 5.4 cm seedling(-1) 5 days after thawing and 88.2 cm seedling(-1) 20 days after thawing. Root hydraulic conductivity increased with new root length initially. But between days 20 and 30, when new root length nearly doubled, P(min) was the only measure of hydraulic conductivity that increased significantly. There were significant correlations (P < 0.05) between white root length and both J(v, min) (r(2) = 0.90) and J(v, max) (r(2) = 0.91).     

Responses of black spruce (Picea mariana) and tamarack (Larix laricina) to flooding and ethylene

Black spruce (Picea mariana (Mill.) BSP) and tamarack (Larix laricina (Du Roi) K. Koch) are the predominant tree species in the boreal peatlands of Alberta, Canada, where low nutrient availability, low soil temperature and a high water table limit their growth. Effects of flooding for 28 days on morphological and physiological responses were investigated in greenhouse-grown black spruce and tamarack seedlings in a growth chamber. Flooding reduced root hydraulic conductance, net assimilation rate and stomatal conductance, and increased water-use efficiency (WUE) and needle electrolyte leakage in both species. Although flooded black spruce seedlings maintained higher net assimilation rates and stomatal conductance than flooded tamarack seedlings, flooded tamarack seedlings were able to maintain higher root hydraulic conductance than flooded black spruce seedlings. Needles of flooded black spruce developed tip necrosis and electrolyte leakage after 14 days of flooding, and these symptoms were subsequently more prominent than in needles of flooded tamarack seedlings. Flooded tamarack seedlings exhibited no visible injury symptoms and developed hypertrophied lenticels at their stem base. Application of exogenous ethylene resulted in a significant reduction in net assimilation, stomatal conductance and root respiration, whereas root hydraulic conductivity increased in both species. Thus, although flooded black spruce seedlings maintained a higher stomatal conductance and net assimilation rate than tamarack seedlings, black spruce did not cope with the deleterious effects of prolonged soil flooding and exogenous ethylene as well as tamarack.     

Uncertainty in detecting climate change impact on the projected yield of black spruce (Picea mariana)

The empirical growth and yield (G&Y) curves used in most timber supply models assume that growth conditions are invariable over time, which may not be correct given the projected future climate. However, errors in G&Y models can be quite large, and we therefore wanted to know the probability of detecting climate change-induced growth anomalies as a departure from current G&Y predictions. The work was carried out in a boreal forest study area in Eastern Canada, where Picea mariana Mill. BSP (black spruce) is the dominant species. Climate change effects were incorporated into G&Y predictions as correction factors obtained from process-based simulations of growth using current or projected climate. Uncertainty in G&Y projections was quantified through the inclusion of sampling and model errors in a bootstrap re-sampling scheme that yielded a percentile distribution of possible differences between G&Y curves that included or did not include climate change effects. Results yielded an average projected increase in productivity of 29% under future climate conditions for the black spruce strata in the study area. The probability of the climate change-modified G&Y predictions being significantly different from the current G&Y projections is 75% when considering the sampling error alone, and of 67% when both the sampling and modelling errors are included. Although incomplete in terms of what sources of error have been included, this study demonstrates the type of information that can be generated on the propagation of uncertainty in G&Y projections and, ultimately, on timber supply.     

Root anchorage and stem strength of black spruce (Picea mariana) trees in regular and irregular stands

A tree winching experiment was conducted, simulating wind actionand resulting damage, in order to assess mechanical resistanceof black spruce (Picea mariana (Mill.) B.S.P.) trees in standsof regular and irregular structures. The critical bending moment(M_c) of winched trees was determined and the relationship betweenM_c and tree and stand characteristics was investigated throughstatistical analyses. Stem mass (SW), average spacing betweentrees (S) and height:diameter ratio (H/d.b.h.) were initiallyselected by a screening procedure to predict M_c. Potential differencesbetween stand structures were tested in mixed models using differentsubsets of the data. Included fixed effects varied between thedifferent models. SW was the most useful and significant variablein all models and H/d.b.h. was significant only when snappedtrees were included in the analysis. When decayed samples wereexcluded, resistance to uprooting was higher in irregular stands.Decay seemed to play an important role in irregular stands andshould be investigated further. Since no difference was observedin the relationship between stem mass and critical turning momentbetween distant sites, relationships should be applicable acrosswide regions. This study provides some of the basic relationshipsrequired to model windthrow risk in irregular stands. However,the effects of stand structure on wind load for individual treeswill also need to be considered.     

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.     

Effects of cone-induction treatments on black spruce (Picea mariana) current-year needle development and gas exchange properties

Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.     

Characterization of juvenile wood to mature wood transition age in black spruce (Picea mariana (Mill.) B.S.P.) at different stand densities and sampling heights

The radial pattern of both maximum ring density and ring area of 36 black spruce trees were used to determine the transition age from juvenile wood to mature wood. The data were obtained by X-ray densitometry and both segmented linear and polynomial regressions were used to point out the age of the juvenile wood boundary. Three stand densities (1,790, 2,700 and 3,400 stems/ha) and three sampling heights (2.4, 5.1 and 7.8 m) were studied. Although maximum ring density and ring area presented similar radial patterns, they gave two significantly different results of transition ages. The maximum ring density over-estimated the transition age (17.6 years) in contrast to ring area (14 years). The results show that the transition from juvenile wood to mature wood occurred after 12 years at 7.8 m (versus 13.1 years at a height of 5.1 m, and 17.6 years at 2.4 m). Although transition age occurred later in the high stand density group (21 years), the difference was not significant between the three stand density groups. Nevertheless, transition age remains difficult to determine since no standard definition exists. The transition occurs over years, and most probably a transition wood exists between juvenile wood and mature wood. Estimation of the juvenile wood proportion in volume shows that it remains constant along the stem and it increases with stand density.     

Production of a new generation of seeds through the use of somatic clones in controlled crosses of black spruce (Picea mariana)

To assess the potential to integrate somatic clones (SC) of desired characteristics in production of high genetic quality seed, controlled crosses between different SCs of black spruce (Picea mariana (Mill) B.S.P.) were used to assess their suitability for the production of viable pollen, cones, seeds and seedlings. These SC produced male and female strobili at an early stage. Pollen, cones and seeds produced were characterized (mass, size, germination); their characteristics were similar to those produced by trees in natural forests or seed orchards. A maternal effect was demonstrated for the cone size and seed mass. Although seeds had excellent germination rates, the somatic biparental crosses were divided into three distinct groups with different germination curves using the Weibull function. Seeds from controlled crosses between different SC enabled the production of high morpho-physiological quality seedlings in a forest nursery. Using black spruce as a model, we showed, for the first time, that SC can be used as seed producers. These encouraging results open new perspectives on the tangible integration of somatic embryogenesis (SE) in the chain of seed, vegetative propagation (cuttings and SE) and production of plants for high productivity plantations. Controlled crosses can be made between SC with the desired characteristics (fewer large branches, fewer nodes, good growth, high wood density, performance, improved yield, etc.), vegetative propagules produced and deployed to clonal tests. After elimination of the worst performing SC, clonal tests can be converted into seed orchards that produce a new generation of seeds of high genetic quality. This will allow the rapid introduction of new materials in elite breeding programs of forest species.     

Bromination of different morphological parts of spruce wood (Picea abies)

Summary It has been found that the lignin in a middle lamella fraction isolated by a sieving technique has the same reactivity to bromine as the whole wood lignin. A plataue value of 1 mole bromine/mole C₉-unit was obtained for both samples. The lignin in compression wood had a considerably lower reactivity to bromine than the normal wood lignin. A plataue value of 0.68 mole bromine/mol C₉-unit was obtained with such lignin. The discrepancy between this and earlier results from the bromination of middle lamella and the implication of the results for lignin determination by the SEM- and TEM-EDXA technique are discussed.     

青海云杉半同胞子代苗期生长性状遗传分析

以青海云杉无性系种子园内30个青海云杉半同胞家系为研究对象,在张掖市肃南县塔尔沟苗圃开展苗期试验,测定5a生苗期苗高、地径、新梢长、侧枝数等生长性状。分析结果表明：（1）青海云杉半同胞子代苗家系间苗高、地径、新梢长存在极显著差异;遗传变异系数较高,均在19．76％以上,苗高、地径、新梢长、侧枝数指标的家系遗传力均较大,均在0．830以上;苗高与地径、新梢长间存在极显著正相关,地径与新梢长、侧枝数存在极显著正相关;（2）以苗高为首要选择指标,按照1倍标准差选择出158#、106#、160#、154#、109#、119#6个优良家系,各性状值高于30个家系的平均值。     

Identification of selected internal wood characteristics in computed tomography images of black spruce: a comparison study

The feasibility of identifying internal wood characteristics in computed tomography (CT) images of black spruce was investigated using two promising classifiers: the maximum likelihood classifier (MLC) and the back propagation (BP) artificial neural network (ANN) classifier. Nine image features including one spectral feature (gray level values), a distance feature, and seven textural features were employed to develop the classifiers. The selected internal wood characteristics to be identified included heartwood, sapwood, bark, and knots. Twenty cross-sectional CT images of a black spruce log were randomly selected to develop the two classifiers. The results suggest that both classifiers produced high classification accuracy. Compared with the MLC classifier (80.9% overall accuracy), the BP ANN classifier had better classification performance (97.6% overall accuracy). Moreover, statistical analysis reveals that the heartwood of the black spruce log used in this study is the easiest to identify by either classifier compared with the other three log features. The results also suggest that the separability of one wood characteristic from the other wood characteristics in black spruce CT images is mainly related to moisture content.     

Effect of thermal modification temperature on the mechanical properties,dimensional stability,and biological durability of black spruce (Picea mariana)

This study was aimed at evaluating the effect of thermal modification temperature on the mechanical properties, dimensional stability, and biological durability of Picea mariana. The boards were thermally modified at different temperatures 190, 200 and 210 °C. The results indicated that the thermal modification of wood caused a significant decrease in the modulus of rupture (MOR) after 190 °C, while the modulus of elasticity (MOE) seemed less affected with a slight increase up to 200 °C and slight decrease with further increase in temperature. The hardness of the thermally modified wood increased in the axial direction. This increase was also observed in tangential and axial directions but at a lesser extent. The final value was slightly higher in axial direction and lower in radial and tangential directions compared to those of the untreated wood. Dimensional stability improved with thermal modification in the three directions compared to the dimensional stability of unmodified wood. The fungal degradation results showed that the decay resistance of thermally modified wood against the wood-rotting fungi Trametes versicolor and Gloephyllum trabeum improved compared to that of the untreated wood. By contrast, the thermal modification of P. mariana had a limited effect on the degradation caused by the fungus Poria placenta.     

Radial mechanical properties of high-temperature dried Norway spruce (Picea abies) wood

Abstract

Wood drying experiments were conducted in which the temperature and the drying rate were controlled independently. The mechanical properties of dried wood in radial loading were analysed in relation to drying parameters. Mass loss, due to thermal degradation of the wood structural components, occurred predominantly in slow high-temperature drying processes. Despite the higher mass loss, slowly dried wood showed similar radial strength and stiffness to rapidly dried wood. The formation of irreversible hydrogen bonds (hornification) within the wood structure may compete with the effects of mass loss on the radial mechanical behaviour of wood. However, both the mass loss and the hornification resulted in wood specimens with lower hygroscopicity. Application of slow high-temperature drying to reduce microscopic cell-wall damage, caused by anisotropic shrinkage of cell-wall layers, did not seem to affect the radial mechanical properties of wood. The effects of stress relaxation within the wood cell wall on the mechanical behaviour of wood may be offset by the degradation of structural components along with drying. Radial mechanical properties may be improved by rapid high-temperature drying up to high final dryness.     

Polyamines in embryogenic cultures of Norway spruce (Picea abies) and red spruce (Picea rubens)

Embryogenic cultures of red spruce (Picea rubens Sarg.) and Norway spruce (Picea abies (L.) Karst.) were initiated from dissected mature zygotic embryos. The tissues were grown on either proliferation medium or maturation medium. On proliferation medium, the embryogenic tissue continued to produce early stage somatic embryos (organized meristems attached to elongated, suspensor-like cells), whereas on maturation medium fully mature embryos developed from the embryonic tissue. Analysis of polyamines in tissues grown on these two media showed that: (1) both putrescine and spermidine concentrations were always higher in cultures grown on proliferation medium than in cultures grown on maturation medium; (2) in both species, spermidine concentrations declined with time in the tissues grown on maturation medium; and (3) spermine was present in only minute quantities and showed only a small change with time. The presence of difluoromethylornithine in the culture medium had little effect on polyamine concentration, whereas the presence of difluoromethylarginine caused a decrease in putrescine concentrations in both red spruce and Norway spruce tissues grown on proliferation medium or maturation medium.     

The variation of the basic density within the juvenile wood of Sitka spruce (Picea sitchensis)

The variation in the basic‐density level of 45 stems of Sitka spruce from two Danish sites has been examined. It is shown that the basic‐density level, i.e. the basic density as a function of ring width, within the juvenile wood decreases with (1) increasing distance from the pith and (2) increasing height in the stem (same ring number from pith provided). Furthermore, the basic‐density level seems to decrease with increasing stand quality, which may affect the pattern of variation in basic‐density level if the stand quality class varies with age. The basic density of the juvenile wood is markedly higher in Sitka spruce than in Norway spruce, whereby the risk of stem cracks probably is reduced.     

Variation in black spruce (<Emphasis Type="Italic">Picea mariana</Emphasis> (Mill.) BSP) wood quality after thinning

• Introduction   

Commercial thinning (CT) could contribute to increase short-term tree growth and be beneficial in a cold climate, as in boreal regions. Thus, growth rate, ring density and flexural modulus of elasticity (MOE) of trees may change after CT. Moreover, mechanical wood properties vary with position in the tree, and there is a need to develop optimal log allocation strategies in order to allocate logs to their best use.