Juvenile-mature wood transition in pine: correlation between wood properties and candidate gene expression profiles

There is a strong desire to exploit transcriptomics data from model species for the genetic improvement of non-model crops. Here, we use gene expression profiles from the commercial model Pinus taeda to identify candidate genes implicated in juvenile-mature wood transition in the non-model relative, P. sylvestris. Re-analysis of 'public domain' SAGE data from xylem tissues of P. taeda revealed 283 mature-abundant and 396 juvenile-abundant tags (P < 0.01), of which 70 and 137, respectively matched to genes with known function. Based on sequence similarity, we then isolated 16 putative homologues of genes that in P. taeda exhibited widest divergence in expression between juvenile and mature samples. Candidate expression levels in P. sylvestris were almost invariably differential between juvenile and mature woody tissue samples among two cohorts of five trees collected from the same seed source and selected for genetic uniformity by genetic distance analysis. However, the direction of differential expression was not always consistent with that described in the original P. taeda SAGE data. Correlation was observed between gene expression and juvenile-mature wood anatomical characteristics by OPLS analysis. Four candidates (alpha-tubulin, porin MIP1, lipid transfer protein and aquaporin like protein) apparently had greatest influence on the wood traits measured. Speculative function of these genes in relation to juvenile-mature wood transition is briefly explored. Thus, we demonstrate the feasibility of exploiting SAGE data from a model species to identify consistently differentially expressed candidates in a related non-model species.

Determination of fiber length and juvenile and mature wood zones from Hevea brasiliensis trees grown in Brazil

The aim of this study was to verify the average fiber length and the juvenile and mature wood zones from Hevea brasiliensis proveniences from a reforestation area in São Paulo, Brazil. For this purpose, five Hevea brasiliensis trees were randomly collected from a 50-year-old plantation, located in Balsamo, São Paulo, Brazil. The trees were cut and five flat sawn boards were obtained. The juvenile and mature wood zones were determined by fiber length measurement from the pith to the bark. The results showed that: (a) the juvenile wood of this species occurred approx. between 40 and 55 mm, from the pith, and from this point forward, the mature wood zone was found; (b) there was a significant difference between the average fiber length of juvenile wood (1.26 mm) and mature wood (1.51 mm).

Effect of early release from intense competition within high density natural regeneration on the properties of the juvenile and mature wood of 40-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.)

Peer reviewed

The effect of initial spacing and thinning on wood quality and graded sawn recovery of mature Caribbean pine grown in Queensland.

A mature Caribbean pine (Pinus caribaea var. hondurensis) silviculture experiment provided initial square spacing treatments of 1.8 m2, 2.4 m2, 3.0 m2 and 3.6 m2 (equal to 3088, 1737, 1111 and 772 stems/ha) that were thinned at age 10 years to 600, 400 and 200 stems/ha, retaining an unthinned control for each initial spacing. The trial was destructively sampled at age of 28 years and discs taken along 8 various stem heights were analysed for variation in basic density and SilviScan wood properties. In addition, the logs from ten stocking × thinning treatments were processed in a sawing study. Results indicate thinning effects were generally more pronounced than initial spacing effects. Fast growing trees produced wood with significantly higher average wood densities and higher average stiffness values. Detailed SilviScan densitometry results obtained radially and at various stem heights enabled construction of tree maps for wood properties, providing insights into the variation in juvenile to mature wood proportion across the initial and post-thinning stocking treatments studied. Dried dressed recovery was strongly related to tree size, and log value decreased consistently from butt to top logs across all treatments. The estimated value per hectare was highest in unthinned plots due to values being multiplied by high stem numbers per hectare. However, a complete economic analysis considering all cost structures is required to investigate the optimal silviculture to maximise economic returns to growers and processors. Improved understanding of the relationship between initial spacing, post-thinning stocking and wood and end-product quality should help to customize future forest management strategies required to produce better quality wood and wood products.

Quantitative and molecular genetics of juvenile wood traits in radiata and slash/Caribbean pines.

The Juvenile Wood Initiative (JWI) project has been running successfully since July 2003 under a Research Agreement with FWPA and Letters of Association with the consortium partners STBA (Southern Tree Breeding Association), ArborGen and FPQ (Forestry Plantations Queensland). Over the last five and half years, JWI scientists in CSIRO, FPQ, and STBA have completed all 12 major milestones and 28 component milestones according to the project schedule. We have made benchmark progress in understanding the genetic control of wood formation and interrelationships among wood traits. The project has made 15 primary scientific findings and several results have been adopted by industry as summarized below. This progress was detailed in 10 technical reports to funding organizations and industry clients. Team scientists produced 16 scientific manuscripts (8 published, 1 in press, 2 submitted, and several others in the process of submission) and 15 conference papers or presentations. Primary Scientific Findings. The 15 major scientific findings related to wood science, inheritance and the genetic basis of juvenile wood traits are: 1. An optimal method to predict stiffness of standing trees in slash/Caribbean pine is to combine gravimetric basic density from 12 mm increment cores with a standing tree prediction of MoE using a time of flight acoustic tool. This was the most accurate and cheapest way to rank trees for breeding selection for slash/Caribbean hybrid pine. This method was also recommended for radiata pine. 2. Wood density breeding values were predicted for the first time in the STBA breeding population using a large sample of 7,078 trees (increment cores) and it was estimated that selection of the best 250 trees for deployment will produce wood density gains of 12.4%. 3. Large genetic variation for a suite of wood quality traits including density, MFA, spiral grain, shrinkage, acoustic and non-acoustic stiffness (MoE) for clear wood and standing trees were observed. Genetic gains of between 8 and 49% were predicted for these wood quality traits with selection intensity between 1 to 10% for radiata pine. 4. Site had a major effect on juvenile-mature wood transition age and the effect of selective breeding for a shorter juvenile wood formation phase was only moderate (about 10% genetic gain with 10% selection intensity, equivalent to about 2 years reduction of juvenile wood). 5. The study found no usable site by genotype interactions for the wood quality traits of density, MFA and MoE for both radiata and slash/Caribbean pines, suggesting that assessment of wood properties on one or two sites will provide reliable estimates of the genetic worth of individuals for use in future breeding. 6. There were significant and sizable genotype by environment interactions between the mainland and Tasmanian regions and within Tasmania for DBH and branch size. 7. Strong genetic correlations between rings for density, MFA and MoE for both radiata and slash/Caribbean pines were observed. This suggests that selection for improved wood properties in the innermost rings would also result in improvement of wood properties in the subsequent rings, as well as improved average performance of the entire core. 8. Strong genetic correlations between pure species and hybrid performance for each of the wood quality traits were observed in the hybrid pines. Parental performance can be used to identify the hybrid families which are most likely to have superior juvenile wood properties of the slash/Caribbean F1 hybrid in southeast Queensland. 9. Large unfavourable genetic correlations between growth and wood quality traits were a prominent feature in radiata pine, indicating that overcoming this unfavourable genetic correlation will be a major technical issue in progressing radiata pine breeding. 10. The project created the first radiata pine 18 k cDNA microarray and generated 5,952 radiata pine xylogenesis expressed sequence tags (ESTs) which assembled into 3,304 unigenes. 11. A total of 348 genes were identified as preferentially expressed genes in earlywood or latewood while a total of 168 genes were identified as preferentially expressed genes in either juvenile or mature wood. 12. Juvenile earlywood has a distinct transcriptome relative to other stages of wood development. 13. Discovered rapid decay of linkage disequilibrium (LD) in radiata pine with LD decaying to approximately 50% within 1,700 base pairs (within a typical gene). A total of 913 SNPS from sequencing 177,380 base pairs were identified for association genetic studies. 14. 149 SNPs from 44 genes and 255 SNPs from a further 51 genes (total 95 genes) were selected for association analysis with 62 wood traits, and 30 SNPs were shortlisted for their significant association with variation of wood quality traits (density, MFA and MoE) with individual significant SNPs accounting for between 1.9 and 9.7% of the total genetic variation in traits. 15. Index selection using breeding objectives was the most profitable selection method for radiata pine, but in the long term it may not be the most effective in dealing with negative genetic correlations between wood volume and quality traits. A combination of economic and biological approaches may be needed to deal with the strong adverse correlation.

Effect of reduced rotation length on Araucaria wood quality.

The review of existing information has identified the following: - the juvenile core in Araucaria is probably contained within the first 15 growth rings in the pith, with spiral grain being a chief determinant of its extent within the stem; - a reduction in rotation length for a given site index will reduce ASV and mature wood volume, with an increase in the proportion of juvenile wood; - for a given rotation length, lower ASV stems were estimated to contain a lower proportion of juvenile wood (based on the assumptions made and crude simulations using WEEDS, PL YSIM and STEPS software); regardless of juvenile wood proportions, smaller stems will yield a higher proportion of pith-in material; - an increase in the proportion of juvenile wood, due to a reduction in rotation length, could affect wood quality due to an increase in the proportion of the recovery containing high spiral grain, shorter tracheids and higher micellar angle; - high spiral grain and high micellar angles adversely impact on wood quality through their influence on twist and longitudinal shrinkage, respectively; - positive outcomes from a reduction in rotation length might include an increase in the proportion of live knots in upper stem sections and a reduction in the extent of brown-stain heartwood; - the uniformity in basic density within Araucaria stems means reduced rotation lengths and lower stem ASVs are unlikely to have a major impact on this wood property, and - the effect of a reduction in rotation length on the incidence of compression wood and timber susceptible to kiln staining could not be established from the available information.

Variation in tracheid cross-sectional dimensions and wood viscoelasticity extent and control methods

Printing papers have been the main product of the Finnish paper industry. To improve properties and economy of printing papers, controlling of tracheid cross-sectional dimensions and wood viscoelasticity are examined in this study. Controlling is understood as any procedure which yields raw material classes with distinct properties and small internal variation. Tracheid cross-sectional dimensions, i.e., cell wall thickness and radial and tangential diameters can be controlled with methods such as sorting wood into pulpwood and sawmill chips, sorting of logs according to tree social status and fractionation of fibres. These control methods were analysed in this study with simulations, which were based on measured tracheid cross-sectional dimensions. A SilviScan device was used to measure the data set from five Norway spruce (Picea abies) and five Scots pine (Pinus sylvestris) trunks. The simulation results indicate that the sawmill chips and top pulpwood assortments have quite similar cross-sectional dimensions. Norway spruce and Scots pine are on average also relatively similar in their cross-sectional dimensions. The distributions of these species are somewhat different, but from a practical point of view, the differences are probably of minor importance. The controlling of tracheid cross-sectional dimensions can be done most efficiently with methods that can separate fibres into earlywood and latewood. Sorting of logs or partitioning of logs into juvenile and mature wood were markedly less efficient control methods than fractionation of fibres. Wood viscoelasticity affects energy consumption in mechanical pulping, and is thus an interesting control target when improving energy efficiency of the process. A literature study was made to evaluate the possibility of using viscoelasticity in controlling. The study indicates that there is considerable variation in viscoelastic properties within tree species, but unfortunately, the viscoelastic properties of important raw material lots such as top pulpwood or sawmill chips are not known. Viscoelastic properties of wood depend mainly on lignin, but also on microfibrillar angle, width of cellulose crystals and tracheid cross-sectional dimensions.

Physical and chemical changes in juvenile and mature woods of Pinus elliottii var. elliottii by thermal modification

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Kraft pulp from juvenile and mature woods of corymbia citriodora

Kraft pulp produced from juvenile and mature wood from thirty-two-year-old Corymbia citriodora trees was evaluated. The stem was subdivided into regions of juvenile and mature wood, and then it was transformed into chips. These materials were then cooked in the Laboratory of Pulp and Paper at São Paulo State University (UNESP, Botucatu, SP, Brazil) and the physico-mechanical properties of the pulps were determined. The results showed that: (1) the pulp yields of mature wood were up to 4.4% greater in comparison to the juvenile wood, (2) the juvenile wood pulp required a shorter refining time than mature wood to reach the same Schopper-Riegler degree, (3) the juvenile wood pulp presented lower specific volume, and (4) the mature wood pulp presented greater air resistance, tensile, tear and burst index values, stress-strain factor, and stretch than the juvenile wood pulp.

Effect of thermal modification on the physical properties of juvenile and mature woods of Eucalyptus grandis

This study aimed to evaluate the effect of thermal treatment on the physical properties of juvenile and mature woods of Eucalyptus grandis. Boards were taken from 30-year-old E. grandis trees. The boards were thermally modified at 180 °C in the Laboratory of Wood Drying and Preservation at UNESP, Botucatu, Sao Paulo state, Brazil. The results showed that thermal modification caused: (1) decrease of 6.8% in the density at 0% equilibrium moisture content of mature wood; (2) significant decreases of 14.7% and 35.6% in the maximum volumetric swellings of juvenile and mature woods, respectively; (3) significant decreases of 13.7% and 21.3% in the equilibrium moisture content of juvenile and mature woods, respectively. The influence of thermal modification in juvenile wood was lower than in mature wood and caused greater uniformity in the physical variations between these types of wood in E. grandis.

Properties of juvenile and mature woods of Hevea brasiliensis untapped and with tapping panels

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

X-ray Microdensitometry Applied to Determination of Wood Density Variation of Eucalyptus grandis W. Hill trees

This study aimed to analyze the variation of wood density in the radial and longitudinal trunk of Eucalyptus grandis trees. Six 23 years old trees were selected and cross sections were cut in three longitudinal positions (DBH, 3.70, 6.10 m) of the log. The results showed that the apparent density of wood (i) increases in the radial direction, characterizing the juvenile wood and mature wood, (ii) no significant variation in base-top log direction was observed. Based on the radial profiles of density, the mature wood of Eucalyptus grandis can be applied in the manufacture of products with higher aggregated value (PMVAs).

Studies on the Cell Wall Structure and on the Mechanical Properties of Norway Spruce

The structure and the mechanical properties of wood of Norway spruce (Picea abies [L.] Karst.) were studied using small samples from Finland and Sweden. X-ray diffraction (XRD) was used to determine the orientation of cellulose microfibrils (microfibril angle, MFA), the dimensions of cellulose crystallites and the average shape of the cell cross-section. X-ray attenuation and x-ray fluorescence measurements were used to study the chemical composition and the trace element content. Tensile testing with in situ XRD was used to characterise the mechanical properties of wood and the deformation of crystalline cellulose within the wood cell walls. Cellulose crystallites were found to be 192 284 Å long and 28.9 33.4 Å wide in chemically untreated wood and they were longer and wider in mature wood than in juvenile wood. The MFA distribution of individual Norway spruce tracheids and larger samples was asymmetric. In individual cell walls, the mean MFA was 19 30 degrees, while the mode of the MFA distribution was 7 21 degrees. Both the mean MFA and the mode of the MFA distribution decreased as a function of the annual ring. Tangential cell walls exhibited smaller mean MFA and mode of the MFA distribution than radial cell walls. Maceration of wood material caused narrowing of the MFA distribution and removed contributions observed at around 90 degrees. In wood of both untreated and fertilised trees, the average shape of the cell cross-section changed from circular via ambiguous to rectangular as the cambial age increased. The average shape of the cell cross-section and the MFA distribution did not change as a result of fertilisation. The mass absorption coefficient for x-rays was higher in wood of fertilised trees than in that of untreated trees and wood of fertilised trees contained more of the elements S, Cl, and K, but a smaller amount of Mn. Cellulose crystallites were longer in wood of fertilised trees than in that of untreated trees. Kraft cooking caused widening and shortening of the cellulose crystallites. Tensile tests parallel to the cells showed that if the mean MFA is initially around 10 degrees or smaller, no systematic changes occur in the MFA distribution due to strain. The role of mean MFA in defining the tensile strength or the modulus of elasticity of wood was not as dominant as that reported earlier. Crystalline cellulose elongated much less than the entire samples. The Poisson ratio νca of crystalline cellulose in Norway spruce wood was shown to be largely dependent on the surroundings of crystalline cellulose in the cell wall, varying between -1.2 and 0.8. The Poisson ratio was negative in kraft cooked wood and positive in chemically untreated wood. In chemically untreated wood, νca was larger in mature wood and in latewood compared to juvenile wood and earlywood.

Propriedades físicas dos lenhos juvenil e adulto de Pinus elliottii Engelm var. elliottii e de Eucalyptus grandis Hill ex Maiden

The juvenile wood presents great difference in its anatomical and physical properties in relation to the mature wood. That heterogeneity of the wood causes a series of upset for the processing industry. The present research had as objective the study of the physical properties of the juvenile and mature wood of Pinus elliottii Engelm var. elliottii, with 25 years of age and of Eucalyptus grandis Hill ex Maiden, with 30 years of age, coming from the Experimental Station of Itapetininga and State Forest of Santa Barbara, belonging to the Forest Institute of the state of São Paulo. From both species there were taken from radial boards, specimens with dimensions of 20x30x50 mm, for the determination of density, swelling and swelling coefficient of the juvenile and mature wood. The results of the species showed that the mature compared with the juvenile wood presented: (1) an increase in the values of the basic density, at 0% and at 12% of moisture content and of the volumetric, radial and tangential swelling at 12% moisture content and maximum coefficient of radial and tangential swelling; (2) a reduction in the values of longitudinal swelling at 12% and maximum and of the longitudinal swelling coefficient.

Propriedades de resistência e rigidez da madeira juvenil e adulta de Pinus taeda L.

O objetivo do presente trabalho foi o estudo da variabilidade das propriedades de resistência e rigidez à flexão estática e à densidade aparente (12%) entre a madeira juvenil e adulta de Pinus taeda L., de 37 anos de idade, procedente do Horto Florestal de Manduri, Estado de São Paulo. Na primeira parte do trabalho foram determinadas a região de madeira juvenil, a região de transição e a região de madeira adulta, por meio de estudos anatômicos (comprimento dos traqueídes axiais), segundo as recomendações das normas ABNT e IAWA. Os resultados mostraram que a região de madeira juvenil dessa espécie ocorre aproximadamente até o 18º anel de crescimento. Na segunda parte do trabalho foram analisados a resistência (módulo de ruptura - MOR) à flexão, o módulo de elasticidade (MOE) nessa mesma solicitação e a densidade aparente (12%) para as madeiras juvenil e adulta. Os resultados mostraram que o MOE e o MOR da madeira juvenil foram menores e mais variáveis que aqueles obtidos para madeira adulta. A densidade apresentou a mesma tendência observada nas propriedades avaliadas no ensaio de flexão estática.