Tree Genetics & Genomes

, Volume 8, Issue 4, pp 895–910 | Cite as

Genetic parameters and provenance variation of Pinus radiata D. Don. ‘Eldridge collection’ in Australia 2: wood properties

  • Washington J. GapareEmail author
  • Miloš Ivković
  • Shannon K. Dillon
  • Fiona Chen
  • Robert Evans
  • Harry X. Wu
Original Paper


Provenance variation and genetic parameters for wood properties of mature radiata pine (Pinus radiata D. Don) were studied by sampling three provenance/progeny trials in southeast Australia. Among the mainland provenances, Monterey and Año Nuevo had higher density and modulus of elasticity (at one site) than Cambria. Basic density and predicted modulus of elasticity (MoE) for the island provenances, Guadalupe and Cedros, were ∼20% higher at Billapaloola compared to mainland provenances grown at Green Hills and Salicki, differences that may or may not be linked to site differences. Heritability estimates of density, predicted MoE and microfibril angle were significant and \( {\bar{h}^2} \) > 0.45, suggesting moderate to strong genetic control. The estimated genetic correlations between diameter at breast height and wood properties in the current study were weaker (less negative) than the mean estimated from the current breeding population generation in radiata pine. Of the wood properties, density showed the strongest adverse genetic correlations with growth (mean r A  = −0.23 ± 0.09). Selection for MoE may produce greater gain than selection for density because MoE had almost twice the estimated additive genetic coefficient of variation (\( {\overline {\text{CV}}_A} \)) compared to density. Estimated type B genetic correlations (r B) for all wood quality traits were typically high, conforming to the trend that wood properties have low genotype-by-environment interaction (G × E). Significant differences in wood properties among provenances, families and/or individual trees provide an opportunity for breeding programmes to select superior trees for solid wood production that will combine superior growth with desirable wood traits.


Pinus radiata Provenances Genetic parameters Density Modulus of elasticity Microfibril angle 



This research was jointly funded by the Commonwealth Scientific and Industrial Research Organisation, Forest and Wood Products Australia, Radiata Pine Breeding Company, Forests New South Wales and the Southern Tree Breeding Association. Thanks to Forestry New South Wales and HVP Plantations for the trial maintenance and assessments over the years. Also, thanks are extended to a large number of people who assisted at various stages of the work, backdating to 1978. We extend special gratitude to our colleague the late Dr K. G. Eldridge for his efforts and vision on gene conservation and utilisation of radiata pine genetic resources. Ian Knight and David Spencer helped with the core extraction. Liming Bian helped with basic density measurements. We thank Drs. Xinguo Li, Adrian Hathorn and Paul Cotterill and an anonymous reviewer for their comments and suggestions on the earlier versions of this paper, and associate editor (Dr. Rowland Burdon), for the helpful comments and suggestions for editing the manuscript.


  1. Ades PK, Garnier-Géré PH (1997) Making sense of provenance x environment interactions in Pinus radiata. In: Burdon RD, Moore JM (eds) IUFRO’97 Genetics of Radiata Pine: proceedings of NZFRI–IUFRO Conference, December 1–4, and Workshop December 5 1997, Rotorua, NZ. New Zealand Forest Research Institute, FRI Bull. No. 203, pp 113–119Google Scholar
  2. Apiolaza LA (2011) Basic density of radiata pine in New Zealand: genetic and environmental factors. Tree Genet Genomes. doi: 10.1007/s11295-011-0423-1
  3. Baltunis BS, Wu HX, Powell MB (2007) Inheritance of density, microfibril angle, and modulus of elasticity in juvenile wood of Pinus radiata at two locations in Australia. Can J Forest Res 37:2164–2174CrossRefGoogle Scholar
  4. Baltunis BS, Gapare WJ, Wu HX (2010) Genetic parameters and genotype by environment interaction in radiata pine for growth and wood quality traits in Australia. Silvae Genet 59:113–124Google Scholar
  5. Burdon RD (1977) Genetic correlation as a concept for studying genotype–environment interaction in forest tree breeding. Silvae Genet 26:168–175Google Scholar
  6. Burdon RD (1992) Genetic survey of Pinus radiata. 9: general discussion and implications for genetic management. NZ J For Sci 22:274–298Google Scholar
  7. Burdon RD (2008) Short note: coefficients of variation in variables with bounded scales. Silvae Genet 57:179–180Google Scholar
  8. Burdon RD, Low CB (1992) Genetic survey of Pinus radiata. 6: wood properties: variation, heritabilities, and interrelationships with other traits. NZ J For Sci 22(2/3):228–245Google Scholar
  9. Burdon RD, Bannister MH, Low CA (1992) Genetic survey of Pinus radiata. 2: population comparisons for growth rate, disease resistance and morphology. NZ J For Sci 22(2/3):138–159Google Scholar
  10. Burdon RD, Firth A, Low CB, Miller MA (1998) Multi-site provenance trials of Pinus radiata in New Zealand. FAO, Rome. For Genet Resour 26:3–8Google Scholar
  11. Burdon RD, Britton RAJ, Walford GB (2001) Wood stiffness and bending strength in relation to density in four native provenances of Pinus radiata. NZ J For Sci 31:130–146Google Scholar
  12. Burdon RD, Kibblewhite RP, Walker JCF, Megraw RA, Evans R, Cown DJ (2004) Juvenile versus mature wood: a new concept, othoganal to corewood versus outerwood, with special reference to Pinus radiata and P. taeda. For Sci 50(4):399–415Google Scholar
  13. Burdon RD, Carson MJ, Shelbourne CJA (2008) Achievements in forest tree improvement in Australia and New Zealand 10. Pinus radiata in New Zealand. Aust For 71:263–279Google Scholar
  14. Cave ID, Walker JCW (1994) Stiffness of wood in fast-grown plantation softwoods: the influence of microfibril angle. For Prod J 44:43–48Google Scholar
  15. Comstock RE, Moll RH (1963) Genotype–environment interactions. In: Hanson WD, Robinson HF (eds) Statistical genetics and plant breeding. NAS-NRC Pub. 982, Washington, pp 164–194Google Scholar
  16. Costa e Silva J, Dutkowski GW, Borralho NMG (2005) Across-site heterogeneity of genetic and environmental variances in the genetic evaluation of Eucalyptus globulus trials for height growth. Ann For Sci 62:183–191CrossRefGoogle Scholar
  17. Cotterill PP, Dean CA (1990) Successful tree breeding with index selection. Division of Forestry and Forest Products, CSIRO, MelbourneGoogle Scholar
  18. Cown DJ (1992) Corewood (juvenile wood) in Pinus radiata: should we be concerned? NZ J For Sci 22:87–95Google Scholar
  19. Cown DJ, van Wyk L (2004) Profitable wood processing—what does it require? Good wood! NZ J For Sci 49:10–14Google Scholar
  20. Cown DJ, Young GD, Burdon RD (1992) Variation in wood characteristics of 20-year-old half-sib families of Pinus radiata. NZ J For Sci 22:63–76Google Scholar
  21. Cown DJ, Hebert J, Ball R (1999) Modelling Pinus radiata lumber characteristics. Part 1: mechanical properties of small clears. NZ J For Sci 29:203–213Google Scholar
  22. Dean CA (1990) Genetics of growth and wood density in radiata pine, Unpublished Ph.D. thesis, University of QueenslandGoogle Scholar
  23. Dillon SK, Nolan M, Li W, Bell C, Wu HX, Southerton SG (2010) Allelic variation in cell wall candidate genes affecting solid wood properties in natural populations and land races of Pinus radiata. Genetics 185:1477–1487PubMedCrossRefGoogle Scholar
  24. Donaldson LA (1993) Variation in microfibril angle among three genetic groups of Pinus radiata trees. NZ J For Sci 23:90–100Google Scholar
  25. Donaldson LA, Burdon RD (1995) Clonal variation and repeatability of microfibril angle in Pinus radiata. NZ J For Sci 25:164–174Google Scholar
  26. Dungey HS, Matheson AC, Kain D, Evans R (2006) Genetics of wood stiffness and its component traits in Pinus radiata. Can J Forest Res 36:1165–1178CrossRefGoogle Scholar
  27. Dungey HS, Brawner JT, Burger F, Carson M, Henson M, Jefferson P, Matheson AC (2007) A new breeding strategy for Pinus radiata in New Zealand and New South Wales. Silvae Genet 58:28–38Google Scholar
  28. Eldridge KG (1978) Refreshing the genetic resources of radiata pine plantations In: Division of forest research: genetics section Report Number 7 CSIRO. 1-120Google Scholar
  29. Evans R (2006) Wood stiffness by X-ray diffractometry. In: Stokke DD, Groom LH (eds) Characterization of the cellulosic cell wall. Wiley, Hoboken, pp 138–148CrossRefGoogle Scholar
  30. Evans R, Ilic J (2001) Rapid prediction of wood stiffness from microfibril angle and density. For Prod J 51:53–57Google Scholar
  31. Evans R, Stuart SA, Van Der Touw J (1996) Microfibril angle scanning of increment cores by X-ray diffractometry. Appita J 49:411–414Google Scholar
  32. Evans R, Hughes MA, Menz D (1999) Microfibril angle variation by scanning X-ray diffractometry. Appita J 52:363–367Google Scholar
  33. Evans R, Ilic J, Matheson AC (2000) Rapid estimation of solid wood stiffness using SilviScan. In: Proc. 26th Forest Products. Research Conf. CSIRO Forestry and Forest Products, Clayton, Australia. pp 49–50Google Scholar
  34. Falconer DS, Mackay TFC (1996) Introduction to quantitative genetics, 4th edn. Longman, Harlow, p 464Google Scholar
  35. Gapare WJ, Wu HX, Abarquez A (2006) Genetic control of the time of transition from juvenile to mature wood in Pinus radiata D. Don. Ann For Sci 63:871–878CrossRefGoogle Scholar
  36. Gapare WJ, Ivkovic M, Baltunis BS, Matheson AC, Wu HX (2010) Genetic stability of wood density and diameter in Pinus radiata D. Don plantation estate in Australia. Tree Genet Genomes 6:113–125CrossRefGoogle Scholar
  37. Gapare WJ, Baltunis BS, Ivkovich M, Low CB, Jefferson P, Wu HX (2011a) Performance differences among ex-situ native-provenance collections of Pinus radiata D. Don. 1: potential for infusion into breeding populations in Australia and New Zealand. Tree Genet Genomes 7:409–419CrossRefGoogle Scholar
  38. Gapare WJ, Ivkovich M, Dutkowski GW, Spencer DJ, Buxton P, Wu, HX (2011b) Genetic parameters and provenance variation of Pinus radiata D. Don. ‘Eldridge collection’ in Australia 1: growth and form traits. doi: 10.1007/s11295-011-0449-4
  39. Garnier-Géré PH, Matheson AC, Ades PK (1997) Assessment of the genetic potential for adaptation of natural provenances: case study of Pinus radiata. In: Proceedings of the NZFRI–IUFRO Conference: IUFRO’97 Genetics of Radiata Pine, 1–4 December 1997, Rotorua, NZ, Burdon RD and Moore JM (eds) New Zealand Forest Research Institute, FRI Bull. No. 203, pp 42–49Google Scholar
  40. Gilmour AR, Gogel BJ, Cullis BR, Thompson R (2009) ASReml User Guide Release 3.0. VSN International Ltd., Hemel Hempstead, HP1 1ES, UK 372 ppGoogle Scholar
  41. Haslett AN, Simpson IG, Kimberley MO (1991) Utilization of 25-year-old Pinus radiata. Part 2. Warp of structural timber in drying. NZ J For Sci 21:228–234Google Scholar
  42. Ivković M, Wu HX, McRae TA, Powell MB (2006) Developing breeding objective for Pinus radiata structural wood production I: bioeconomic model and economic weights. Can J Forest Res 36:2920–2931CrossRefGoogle Scholar
  43. Ivković M, Gapare WJ, Abaquez A, Ilic J, Powell MB, Wu HX (2009) Prediction of wood stiffness, strength, and shrinkage in juvenile wood of radiata pine. Wood Sci Tech 43:237–257CrossRefGoogle Scholar
  44. Johnson GR, Gartner BL (2006) Genetic variation in basic density and modulus of elasticity of coastal Douglas-fir. Tree Genet Genomes 3:25–33CrossRefGoogle Scholar
  45. Johnson IG, Ades PK, Eldridge KG (1997) Growth of natural Californian provenances of Pinus radiata in New South Wales, Australia. NZ J For Sci 27:23–38Google Scholar
  46. Kennedy G (2004) Variation in wood density and diameter growth between inter- and intra-provenance crosses of Pinus radiata D. Don. Unpublished BSc Hons Thesis, Australian National University, Canberra, AustraliaGoogle Scholar
  47. Kininmonth JA, Whitehouse LJ (eds) (1991) Properties and uses of New Zealand radiata pine, vol 1, Wood properties. New Zealand Forest Research Institute, RotoruaGoogle Scholar
  48. Kumar S (2004) Genetic parameter estimates for wood stiffness, strength, internal checking, and resin bleeding for radiata pine. Can J Forest Res 34:2601–2610CrossRefGoogle Scholar
  49. Kumar S, Jayawickrama KJS, Lee J, Lausberg M (2002) Direct and indirect measures of stiffness and strength show high heritability in a wind-pollinated radiata pine progeny test in New Zealand. Silvae Genet 51(5–6):256–261Google Scholar
  50. Kumar S, Dungey HS, Matheson AC (2006) Genetic parameters and strategies for genetic improvement of stiffness in radiata pine. Silvae Genet 55:77–84Google Scholar
  51. Kumar S, Burdon RD, Stovold GT (2008) Wood properties and stem diameter of Pinus radiata in New Zealand: clonal and seedling material. NZ J For Sci 38:88–101Google Scholar
  52. Lachenbruch B, Johnson GR, Downes GM, Evans R (2010) Relationships of density, microfibril angle, and sound velocity with stiffness and strength in mature wood of Douglas-fir. Can J Forest Res 40:55–64CrossRefGoogle Scholar
  53. Li L, Wu HX (2005) Efficiency of early selection for rotation-aged growth and wood density traits in Pinus radiata. Can J Forest Res 35:2019–2029CrossRefGoogle Scholar
  54. Lindström H, Evans R, Reale M (2005) Implications of selecting tree clones with high modulus of elasticity. NZ J For Sci 35:50–71Google Scholar
  55. Low CB, Smith T (1997) Use of the Guadalupe provenance in Pinus radiata improvement in New Zealand. In: Burdon RD, Moore JM (eds) IUFRO’97 Genetics of Radiata Pine: proceedings of NZFRI–IUFRO Conference, December 1–4, and Workshop December 5 1997, Rotorua, NZ. New Zealand Forest Research Institute, FRI Bull. No. 203, pp 57–61Google Scholar
  56. Lundgren C (2004) Microfibril angle and density patterns of fertilized and irrigated Norway spruce. Silva Fenn 38:107–117Google Scholar
  57. Matheson AC, Raymond CA (1984) The impact of genotype × environment interactions on Australian Pinus radiata breeding programs. Aust For Res 14:11–25Google Scholar
  58. Matheson AC, Gapare WJ, Ilic J, Wu HX (2008) Inheritance and genetic gain in wood stiffness in radiata pine assessed acoustically in young standing trees. Silvae Genet 57:56–64Google Scholar
  59. Megraw RA, Leaf G, Bremer D (1998) Longitudinal shrinkage and microfibril angle in loblolly pine. In: Butterfield BG (ed) Microfibril angle in wood. University of Canterbury Press, Christchurch, pp 27–61Google Scholar
  60. Moran GF, Bell JC, Eldridge KG (1988) The genetic structure and the conservation of the five natural-populations of Pinus radiata. Can J Forest Res 18:506–514CrossRefGoogle Scholar
  61. Myszewski JH, Bridgewater FE, Lowe WJ, Byram TD, Megraw RA (2004) Genetic variation in the microfibril angle of loblolly pine from two test sites. South J App For 28:196–204Google Scholar
  62. Powell MB, McRae TA, Wu HX, Dutkowski GW, Pilbeam DJ (2004) Breeding strategy for Pinus radiata in Australia. 2004 IUFRO Joint Conference of Division 2: Forest Genetics and Tree Breeding in the Age of Genomics: Progress and Future. Charleston, South Carolina, USA, 1–5 November, 2004, pp 308–18Google Scholar
  63. Raymond CA, Henson M, Joe B (2009) Genetic variation amongst and within the native provenances of Pinus radiata D. Don in South-eastern Australia. 2. Wood density and stiffness to age 26 years. Silvae Genet 58:192–204Google Scholar
  64. Shelbourne CJA (1972) Genotype–environment interaction: its study and its implications in forest tree improvement. In: IUFRO Genetics SABRAO joint symposium, TokyoGoogle Scholar
  65. Stram DO, Lee JW (1994) Variance components testing in the longitudinal mixed effects model. Biometrics 50(4):1171–1177PubMedCrossRefGoogle Scholar
  66. Vermaas HF (1988) Combination of a special water immersion method with the maximum moisture content method for bulk wood density determination. Holzforschung 42:131–134CrossRefGoogle Scholar
  67. Vogl C, Karhu A, Moran G, Savolainen O (2002) High resolution analysis of mating systems: inbreeding in natural populations of Pinus radiata. J Evol Biol 15:433–439CrossRefGoogle Scholar
  68. Walker JCF, Butterfield BG (1995) The importance of microfibril angle for the processing industries. NZ For 40:34–40Google Scholar
  69. Walker JFC, Nakada R (1999) Understanding corewood in some softwoods: a selective review on stiffness and acoustics. Int For Rev 1:251–259Google Scholar
  70. Wielinga B, Raymond CA, James R, Matheson AC (2009) Genetic parameters and genotype by environment interactions for green and basic density and stiffness of Pinus radiata D. Don estimated using acoustics. Silvae Genet 58:112–122Google Scholar
  71. Wu HX, Matheson AC (2005) Genotype by environment interaction in an Australia-wide radiata pine diallel mating experiment: implications for regionalized breeding. For Sci 5:1–11Google Scholar
  72. Wu HX, Yang JL, McRae TA, Li L, Ivkovich M, Powell MB (2004) Breeding for wood quality and profits with radiata pine 1: MOE prediction and genetic correlation between early growth, density, microfibril angle and rotation-age MOE. In: Proceedings of Wood quality 2004: practical tools and new technologies to improve segregation of logs and lumber for processing, AlburyGoogle Scholar
  73. Wu HX, Powell MB, Yang JL, Ivković M, McRae TA (2006) Efficiency of early selection for rotation-aged wood quality traits in radiata pine. Ann For Sci 64:1–9CrossRefGoogle Scholar
  74. Wu HX, Eldridge KG, Matheson AC, Powell MB, McRae TA (2007) Achievement in forest tree improvement in Australia and New Zealand: successful introduction and breeding of radiata pine to Australia. Aust For 70:215–225Google Scholar
  75. Young GD, McConchie DL, McKinley (1991) Utilization of 25-year-old Pinus radiata part 1: wood properties. NZ J For Sci 21:217–227Google Scholar
  76. Zamudio F, Baettyg R, Vergara RA, Guerra F, Rozenberg P (2002) Genetic trends in wood density and radial growth with cambial age in a radiata pine progeny test. Ann For Sci 59:541–549CrossRefGoogle Scholar
  77. Zobel BJ, Jett JB (1995) Genetics of wood production. Springer, Berlin, p 337CrossRefGoogle Scholar
  78. Zobel BJ, van Buijtenen JP (1989) Wood variation: its causes and control. Springer, Berlin, p 363CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2012

Authors and Affiliations

  • Washington J. Gapare
    • 1
    Email author
  • Miloš Ivković
    • 1
  • Shannon K. Dillon
    • 1
  • Fiona Chen
    • 2
  • Robert Evans
    • 2
  • Harry X. Wu
    • 1
    • 3
  1. 1.CSIRO Plant IndustryCanberraAustralia
  2. 2.CSIRO Materials Science and EngineeringMelbourneAustralia
  3. 3.Umeå Plant Science Centre, Department Forest Genetics and Plant PhysiologySwedish University of Agricultural SciencesUmeåSweden

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