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.

Wood properties and processing outcomes for plantation grown African mahogany (Khaya senegalensis) trees from Clare, Queensland (18 and 20 year-old trees) and Katherine, Northern Territory (14 year-old trees).

Khaya senegalensis, African mahogany, a high-value hardwood, was introduced in the Northern Territory (NT) in the 1950s; included in various trials there and at Weipa, Q in the 1960s-1970s; planted on ex mine sites at Weipa (160 ha) until 1985; revived in farm plantings in Queensland and in trials in the NT in the 1990s; adopted for large-scale, annual planting in the Douglas-Daly region, NT from 2006 and is to have the planted area in the NT extended to at least 20,000 ha. The recent serious interest from plantation growers, including Forest Enterprises Australia Ltd (FEA), has seen the establishment of some large scale commercial plantations. FEA initiated the current study to process relatively young plantation stands from both Northern Territory and Queensland plantations to investigate the sawn wood and veneer recovery and quality from trees ranging from 14 years (NT – 36 trees) to 18-20 years (North Queensland – 31 trees). Field measures of tree size and straightness were complemented with log end splitting assessment and cross-sectional disc sample collection for laboratory wood properties measurements including colour and shrinkage. End-splitting scores assessed on sawn logs were relatively low compared to fast grown plantation eucalypts and did not impact processing negatively. Heartwood proportion in individual trees ranged from 50% up to 92 % of butt cross-sectional disc area for the visually-assessed dark coloured central heartwood and lighter coloured transition wood combined. Dark central heartwood proportion was positively related to tree size (R2 = 0.57). Chemical tests failed to assist in determining heartwood – sapwood boundary. Mean basic density of whole disc samples was 658 kg/m3 and ranged among trees from 603 to 712 kg/m3. When freshly sawn, the heartwood of African mahogany was orange-red to red. Transition wood appeared to be pinkish and the sapwood was a pale yellow colour. Once air dried the heartwood colour generally darkens to pinkish-brown or orange-brown and the effect of prolonged time and sun exposure is to darken and change the heartwood to a red-brown colour. A portable colour measurement spectrophotometer was used to objectively assess colour variation in CIE L*, a* and b* values over time with drying and exposure to sunlight. Capacity to predict standard colour values accurately after varying periods of direct sunlight exposure using results obtained on initial air-dried surfaces decreased with increasing time to sun exposure. The predictions are more accurate for L* values which represent brightness than for variation in the a* values (red spectrum). Selection of superior breeding trees for colour is likely to be based on dried samples exposed to sunlight to reliably highlight wood colour differences. A generally low ratio between tangential and radial shrinkages was found, which was reflected in a low incidence of board distortion (particularly cupping) during drying. A preliminary experiment was carried out to investigate the quality of NIR models to predict shrinkage and density. NIR spectra correlated reasonably well with radial shrinkage and air dried density. When calibration models were applied to their validation sets, radial shrinkage was predicted to an accuracy of 76% with Standard Error of Prediction of 0.21%. There was also a strong predictive power for wood density. These are encouraging results suggesting that NIR spectroscopy has good potential to be used as a non-destructive method to predict shrinkage and wood density using 12mm diameter increment core samples. Average green off saw recovery was 49.5% (range 40 to 69%) for Burdekin Agricultural College (BAC) logs and 41.9% (range 20 to 61%) for Katherine (NT) logs. These figures are about 10% higher than compared to 30-year-old Khaya study by Armstrong et al. (2007) however they are inflated as the green boards were not docked to remove wane prior to being tallied. Of the recovered sawn, dried and dressed volume from the BAC logs, based on the cambial face of boards, 27% could potentially be used for select grade, 40% for medium feature grade and 26% for high feature grades. The heart faces had a slightly higher recovery of select (30%) and medium feature (43%) grade boards with a reduction in the volume of high feature (22%) and reject (6%) grade boards. Distribution of board grades for the NT site aged 14 years followed very similar trends to those of the BAC site boards with an average (between facial and cambial face) 27% could potentially be used for select grade, 42% for medium feature grade, 26% for high feature grade and 5% reject. Relatively to some other subtropical eucalypts, there was a low incidence of borer attack. The major grade limiting defects for both medium and high feature grade boards recovered from the BAC site were knots and wane. The presence of large knots may reflect both management practices and the nature of the genetic material at the site. This stand was not managed for timber production with a very late pruning implemented at about age 12 years. The large amount of wane affected boards is indicative of logs with a large taper and the presence of significant sweep. Wane, knots and skip were the major grade limiting defects for the NT site reflecting considerable amounts of sweep with large taper as might be expected in younger trees. The green veneer recovered from billets of seven Khaya trees rotary peeled on a spindleless lathe produced a recovery of 83% of green billet volume. Dried veneer recovery ranged from 40 to 74 % per billet with an average of 64%. All of the recovered grades were suitable for use in structural ply in accordance to AS/NZ 2269: 2008. The majority of veneer sheets recovered from all billets was C grade (27%) with 20% making D grade and 13% B grade. Total dry sliced veneer recovery from the logs of the two largest logs from each location was estimated to be 41.1%. Very positive results have been recorded in this small scale study. The amount of colour development observed and the very reasonable recoveries of both sawn and veneer products, with a good representation of higher grades in the product distribution, is encouraging. The prospects for significant improvement in these results from well managed and productive stands grown for high quality timber should be high. Additionally, the study has shown the utility of non-destructive evaluation techniques for use in tree improvement programs to improve the quality of future plantations. A few trees combined several of the traits desired of individuals for a first breeding population. Fortunately, the two most promising trees (32, 19) had already been selected for breeding on external traits, and grafts of them are established in the seed orchard.

Forest and Wood Products Research and Development Corporation (FWPRDC) Cypress exatractives

Study to investigate the use of cypress pine extractives as a novel timber preservative.

Strength and stiffness assessment technologies for improving grading effectiveness of radiata pine wood.

This work was designed to provide the Australian structural radiata pine processing industry with some indications for improving stress grading methods and/or technologies to give an increase in structural grade yields, and significantly reduce processing costs without compromising product quality. To achieve this, advanced statistical techniques were used in conjunction with state-of-the-art property measurement systems applied to the same sample of sawn timber. Acoustic vibration analyses were conducted on green and dry boards. Raw data from existing in-line systems was captured on the same boards. The Metriguard HCLT stress rating system was used as the "reference" machine grading because of its current common use in the industry. A WoodEye optical scanning system and an X-ray LHG scanner were also able to provide relevant information on knots. The data set was analyzed using classical and advanced statistical tools to provide correlations between data sets, and to develop efficient strength and stiffness prediction equations. Reductions in non-structural dry volumes can be achieved..

Evaluation of wood characteristics of tropical post-mid rotation plantation Eucalyptus cloeziana and E. pellita: Part (a) Natural durability of timber.

This project evaluated the timber quality, processing and performance characteristics of 19-year-old Eucalyptus cloeziana (Gympie messmate) and 15-year-old Eucalyptus pellita (red mahogany). Studies were undertaken to evaluate wood and mechanical properties, accelerated seasoning and veneer and plywood production. Above-ground and in-ground durability field tests were established at three locations in Queensland. Ground proixmity tests and L-joint tests were installed to gather data applicable to above-ground, weather-exposed end-use applications, and stake tests were installed to gather data applicable to in-ground, weather-exposed end-use applications.

The use of genetic correlations to evaluate associations between SNP markers and quantitative traits

Open-pollinated progeny of Corymbia citriodora established in replicated field trials were assessed for stem diameter, wood density, and pulp yield prior to genotyping single nucleotide polymorphisms (SNP) and testing the significance of associations between markers and assessment traits. Multiple individuals within each family were genotyped and phenotyped, which facilitated a comparison of standard association testing methods and an alternative method developed to relate markers to additive genetic effects. Narrow-sense heritability estimates indicated there was significant additive genetic variance within this population for assessment traits ( h ˆ 2 =0.28to0.44 ) and genetic correlations between the three traits were negligible to moderate (r G = 0.08 to 0.50). The significance of association tests (p values) were compared for four different analyses based on two different approaches: (1) two software packages were used to fit standard univariate mixed models that include SNP-fixed effects, (2) bivariate and multivariate mixed models including each SNP as an additional selection trait were used. Within either the univariate or multivariate approach, correlations between the tests of significance approached +1; however, correspondence between the two approaches was less strong, although between-approach correlations remained significantly positive. Similar SNP markers would be selected using multivariate analyses and standard marker-trait association methods, where the former facilitates integration into the existing genetic analysis systems of applied breeding programs and may be used with either single markers or indices of markers created with genomic selection processes.

Estimates of genetic parameters for growth and wood properties in Eucalyptus pellita F. Muell. to support tree breeding in Vietnam

Key message Eucalyptus pellita demonstrated good growth and wood quality traits in this study, with young plantation grown timber being suitable for both solid and pulp wood products. All traits examined were under moderate levels of genetic control with little genotype by environment interaction when grown on two contrasting sites in Vietnam. Context Eucalyptus pellita currently has a significant role in reforestation in the tropics. Research to support expanded of use of this species is needed: particularly, research to better understand the genetic control of key traits will facilitate the development of genetically improved planting stock. Aims This study aimed to provide estimates of the heritability of diameter at breast height over bark, wood basic density, Kraft pulp yield, modulus of elasticity and microfibril angle, and the genetic correlations among these traits, and understand the importance of genotype by environment interactions in Vietnam. Methods Data for diameter and wood properties were collected from two 10-year-old, open-pollinated progeny trials of E. pellita in Vietnam that evaluated 104 families from six native range and three orchard sources. Wood properties were estimated from wood samples using near-infrared (NIR) spectroscopy. Data were analysed using mixed linear models to estimate genetic parameters (heritability, proportion of variance between seed sources and genetic correlations). Results Variation among the nine sources was small compared to additive variance. Narrow-sense heritability and genetic correlation estimates indicated that simultaneous improvements in most traits could be achieved from selection among and within families as the genetic correlations among traits were either favourable or close to zero. Type B genetic correlations approached one for all traits suggesting that genotype by environment interactions were of little importance. These results support a breeding strategy utilizing a single breeding population advanced by selecting the best individuals across all seed sources. Conclusion Both growth and wood properties have been evaluated. Multi-trait selection for growth and wood property traits will lead to more productive populations of E. pellita both with improved productivity and improved timber and pulp properties.

Estimates of genetic parameters for growth and wood properties in Eucalyptus pellita F. Muell. to support tree breeding in Vietnam

Eucalyptus pellita demonstrated good growth and wood quality traits in this study, with young plantation grown timber being suitable for both solid and pulp wood products. All traits examined were under moderate levels of genetic control with little genotype by environment interaction when grown on two contrasting sites in Vietnam. Eucalyptus pellita currently has a significant role in reforestation in the tropics. Research to support expanded of use of this species is needed: particularly, research to better understand the genetic control of key traits will facilitate the development of genetically improved planting stock. This study aimed to provide estimates of the heritability of diameter at breast height over bark, wood basic density, Kraft pulp yield, modulus of elasticity and microfibril angle, and the genetic correlations among these traits, and understand the importance of genotype by environment interactions in Vietnam. Data for diameter and wood properties were collected from two 10-year-old, open-pollinated progeny trials of E. pellita in Vietnam that evaluated 104 families from six native range and three orchard sources. Wood properties were estimated from wood samples using near-infrared (NIR) spectroscopy. Data were analysed using mixed linear models to estimate genetic parameters (heritability, proportion of variance between seed sources and genetic correlations). Variation among the nine sources was small compared to additive variance. Narrow-sense heritability and genetic correlation estimates indicated that simultaneous improvements in most traits could be achieved from selection among and within families as the genetic correlations among traits were either favourable or close to zero. Type B genetic correlations approached one for all traits suggesting that genotype by environment interactions were of little importance. These results support a breeding strategy utilizing a single breeding population advanced by selecting the best individuals across all seed sources. Both growth and wood properties have been evaluated. Multi-trait selection for growth and wood property traits will lead to more productive populations of E. pellita both with improved productivity and improved timber and pulp properties.