Advertisement

Trees

, Volume 29, Issue 5, pp 1395–1413 | Cite as

A model of stem growth and wood formation in Pinus radiata

  • David M. Drew
  • Geoff Downes
Original Paper
Part of the following topical collections:
  1. Biomechanics

Abstract

Key message

A model of wood formation processes in pines predicted 80 % of mean wood density variation from inputs of carbohydrate allocation and tree water status from several varied sites.

Abstract

Numerous factors determine how wood properties vary as a tree grows. In order to model wood formation, a framework that considers the various xylogenetic processes is required. We describe a new model of xylem development and wood formation in pines (parameterised for the commercially important species, Pinus radiata D. Don). In this paper, we use as inputs simulated daily data from the CaBala stand growth model which, in turn, takes into account site and daily weather conditions, and silviculture. It incorporates a first attempt at predicting microfibril angle (the angle of cellulose microfibrils relative to the vertical axis of the cell, MFA) based on metrics of cambial vigour and carbohydrate allocation. It also predicts tracheid dimensions and wall thickness, and from these data, wood density. Pith-to-bark and intra-annual variation in predicted wood properties was realistic across a wide range of site types, although juvenile wood properties were weakly predicted. The model was able to explain 50 % of the variation in outerwood MFA and 70–80 % of the variation in outerwood and mean sample wood density respectively, from 17 study sites. The model, early results from which are very promising, provides a useful framework for testing concepts of how formation occurs, and to provide insights into areas where further research is needed.

Keywords

Cambium Pinus radiata Secondary thickening Carbohydrate allocation Wood density Microfibril angle Xylem 

Notes

Acknowledgments

This work was funded by Forest and Wood Products Australia (FWPA), Forestry SA, Hancock Victoria Plantations (HVP), Scion and the CSIRO Sustainable Agriculture Flagship. Thank you to Warwick Gill for embedding and sectioning work, Jody Bruce and Michael Battaglia for their advice during the setting up of model scenarios in Cabala. Also thanks to staff at CSIRO (Dale Worledge), Forestry SA (Jim O’Hehir, Don McGuire and Stuart Adam), HVP (Stephen Elms and Ross Gillies) and Scion (Jonathan Harrington), all of whom contributed to the work in various important ways. Thanks to Chris Beadle, Daniel Mendham and Patrick Mitchell for helpful comments on earlier versions of the manuscript.

Conflict of interest

The authors declare that they have no conflict of interest.

References

  1. Abe H, Nakai T (1999) Effect of the water status within a tree on tracheid morphogenesis in Cryptomeria japonica. Trees 14:124–129Google Scholar
  2. Almeida AC, Landsberg JJ, Sands PJ, Ambrogi MS, Fonseca S, Barddal SM, Bertolucci FL (2004) Needs and opportunities for using a process-based productivity model as a practical tool in Eucalyptus plantations. For Ecol Manag 193:167–177CrossRefGoogle Scholar
  3. Anfodillo T, Deslauriers A, Menardi R, Tedoldi L, Petit G, Rossi S (2012) Widening of xylem conduits in a conifer tree depends on the longer time of cell expansion downwards along the stem. J Exp Bot 63:837–845PubMedCentralCrossRefPubMedGoogle Scholar
  4. Apiolaza L, Chauhan S, Hayes M, Nakada R, Sharma M, Walker J (2013) Selection and breeding for wood quality A new approach. N Z J For 58:32–37Google Scholar
  5. Auty D, Gardiner B, Achim A, Moore J, Cameron A (2013) Models for predicting microfibril angle variation in Scots pine. Ann For Sci 70:209–218CrossRefGoogle Scholar
  6. Barnett JR (1973) Seasonal Variation in the ultrastructure of the cambium in New Zealand grown Pinus radiata D. Don. Ann Bot Lond 37:1005–1011Google Scholar
  7. Barnett JR, Bonham VA (2004) Cellulose microfibril angle in the cell wall of wood fibres. Biol Rev 79:461–472CrossRefPubMedGoogle Scholar
  8. Battaglia M, Sands P (1997) Modelling site productivity of Eucalyptus globulus in response to climatic and site factors. Aust J Plant Physiol 24:831–850CrossRefGoogle Scholar
  9. Battaglia M, Sands P, White D, Mummery D (2004) CABALA: a linked carbon, water and nitrogen model of forest growth for silvicultural decision support. For Ecol Manag 193:251–282CrossRefGoogle Scholar
  10. Bauerle WL, Oren R, Way DA, Qian SS, Stoy PC, Thornton PE, Bowden JD, Hoffman FM, Reynolds RF (2012) Photoperiodic regulation of the seasonal pattern of photosynthetic capacity and the implications for carbon cycling. Proc Natl Acad Sci 109:8612–8617PubMedCentralCrossRefPubMedGoogle Scholar
  11. Boardman R (1988) Living on the edge—the development of silviculture in South Australian pine plantations. Aust For 51:135–156CrossRefGoogle Scholar
  12. Bogoslavsky L, Neumann PM (1998) Rapid regulation by acid pH of cell wall adjustment and leaf growth in Maize plants responding to reversal of water stress. Plant Physiol 118:701–709PubMedCentralCrossRefPubMedGoogle Scholar
  13. Bollhöner B, Prestele J, Tuominen H (2012) Xylem cell death: emerging understanding of regulation and function. J Exp Bot 63:1081–1094CrossRefPubMedGoogle Scholar
  14. Burdon RD, Kibblewhite RP, Walker JC, Megraw RA, Evans R, Cown DJ (2004) Juvenile versus mature wood: a new concept, orthogonal to corewood versus outerwood, with special reference to Pinus radiata and P. taeda. For Sci 50:399–415Google Scholar
  15. Catesson AM, Roland JC (1981) Sequential changes associated with cell wall formation and fusion in the vascular cambium. IAWA Bull 2:151–162CrossRefGoogle Scholar
  16. Chan J (2012) Microtubule and cellulose microfibril orientation during plant cell and organ growth. J Microsc 247(1):23–32Google Scholar
  17. Cosgrove DJ (2001) Wall structure and wall loosening. A look backwards and forwards. Plant Physiol 125:131–134PubMedCentralCrossRefPubMedGoogle Scholar
  18. Cosgrove DJ (2005) Growth of the plant cell wall. Nat Rev Mol Cell Biol 6:850–861CrossRefPubMedGoogle Scholar
  19. Deckmyn G, Evans SP, Randle TJ (2006) Refined pipe theory for mechanistic modelling of wood development. Tree Physiol 26:703–717CrossRefPubMedGoogle Scholar
  20. Deleuze C, Houllier F (1998) A simple process-based xylem growth model for describing wood microdensitometric profiles. J Theor Biol 193:99–113CrossRefGoogle Scholar
  21. Denne MP (1971) Temperature and tracheid development in Pinus sylvestris seedlings. J Exp Bot 22:362–370CrossRefGoogle Scholar
  22. Denne MP (1976) Effects of environmental change on wood production and wood structure in Picea sitchensis seedlings. Ann Bot Lond 40:1017–1028Google Scholar
  23. Denne MP, Dodd RS (1981) The environmental control of xylem differentiation. In: Barnett JR (ed) Xylem cell developement. Castle House Publications, Kent, pp 236–255Google Scholar
  24. Dodd RS, Fox P (1990) Kinetics of tracheid differentiation in Douglas-fir. Ann Bot Lond 65:649–657Google Scholar
  25. Donaldson L (2008) Microfibril angle: measurement, variation and relationships—a review. IAWA J 29:345–386CrossRefGoogle Scholar
  26. Downes GM, Drew DM (2008) Climate and growth influences on wood formation and utilisation. South For 70:155–167Google Scholar
  27. Downes GM, Beadle C, Gensler W, Mummery D, Worledge D (1999) Diurnal variation and radial growth of stems in young plantation eucalypts. In: Wimmer R, Vetter RE (eds) Tree ring analysis. Biological, methodological and environmental aspects. CAB International, New York, pp 83–104Google Scholar
  28. Downes GM, Wimmer R, Evans R (2004) Interpreting sub-annual wood property variation in terms of stem growth. Wood fibre cell walls: methods to study their formation, structure and properties. Swedish University of Agricultural Sciences, Department of Wood Science, pp 267–283Google Scholar
  29. Drew DM, Pammenter NW (2007) Developmental rates and morphological properties of fibres in two eucalypt clones at sites differing in water availability. South Hemisph For J 69:71–79CrossRefGoogle Scholar
  30. Drew DM, Downes GM, Battaglia M (2010) CAMBIUM, a process-based model of daily xylem development in Eucalyptus. J Theor Biol 264:395–406CrossRefPubMedGoogle Scholar
  31. Drew DM, Richards AE, Cook GD, Downes GM, Gill W, Baker PJ (2013) The number of days on which increment occurs is the primary determinant of annual ring width in Callitris intratropica. Trees: 1–10Google Scholar
  32. Duan H, Amthor JS, Duursma RA, O’Grady AP, Choat B, Tissue DT (2013) Carbon dynamics of eucalypt seedlings exposed to progressive drought in elevated [CO2] and elevated temperature. Tree Physiol 65(5):1313–1321Google Scholar
  33. Duchesne L, Houle D, D’Orangeville L (2012) Influence of climate on seasonal patterns of stem increment of balsam fir in a boreal forest of Québec. Canada Agric For Meteorol 162–163:108–114CrossRefGoogle Scholar
  34. Escamez S, Tuominen H (2014) Programmes of cell death and autolysis in tracheary elements: when a suicidal cell arranges its own corpse removal. J Exp Bot 65(5):1313–1321CrossRefPubMedGoogle Scholar
  35. Evans R (1994) Rapid measurement of the transverse dimensions of tracheids in radial wood sections from Pinus radiata. Holzforschung 48:168–172CrossRefGoogle Scholar
  36. Evans R, Ilic J (2001) Rapid prediction of wood stiffness from microfibril angle and density. For Prod J 51:53–57Google Scholar
  37. Feikema PM, Morris JD, Beverly CR, Collopy JJ, Baker TG, Lane PNJ (2010) Validation of plantation transpiration in south-eastern Australia estimated using the 3PG+ forest growth model. For Ecol Manage 260:663–678CrossRefGoogle Scholar
  38. Fernández MP, Norero A, Vera JR, Pérez E (2011) A functional–structural model for radiata pine (Pinus radiata) focusing on tree architecture and wood quality. Ann Bot Lond 108:1155–1178CrossRefGoogle Scholar
  39. Fritts HC (1976) Tree rings and climate. Academic Press, New YorkGoogle Scholar
  40. Fritts HC, Shashkin A, Downes GM (1999) TreeRing 3: a simulation model of conifer ring growth and cell structure. In: Wimmer R, Vetter RE (eds) Tree ring analysis: biological, methodological and environmental aspects. CAB International, Oxford, pp 3–32Google Scholar
  41. Fromm J (2013) Xylem development in trees: from cambial divisions to mature wood cells. In: Fromm J (ed) Cellular aspects of wood formation, vol 20. Springer, Berlin Heidelberg, pp 3–39CrossRefGoogle Scholar
  42. Gavran M, Parsons M (2011) Australian plantation statistics 2011. Australian Bureau of Agricultural and Resource Economics and Sciences, CanberraGoogle Scholar
  43. Gričar J, Zupančič M, Čufar K, Oven P (2007) Regular cambial activity and xylem and phloem formation in locally heated and cooled stem portions of Norway spruce. Wood Sci Technol 41:463–475CrossRefGoogle Scholar
  44. Harashima H, Schnittger A (2010) The integration of cell division, growth and differentiation. Curr Opin Plant Biol 13:66–74CrossRefPubMedGoogle Scholar
  45. Haygreen JG, Bowyer JL (1982) Forest products and wood science: an introduction. Iowa State University Press, Ames, Iowa, p 495Google Scholar
  46. Hölttä T, Mäkinen H, Nöjd P, Mäkelä A, Nikinmaa E (2010) A physiological model of softwood cambial growth. Tree Physiol 30:1235–1252CrossRefPubMedGoogle Scholar
  47. Horacek P, Slezingerova J, Gandelova L, Wimmer R, Vetter R (1999) Effects of environment on the xylogenesis of Norway spruce (Picea abies [L.] Karst.). In: Vetter RE (ed) Tree-ring analysis: biological, methodological and environmental aspects, pp 35–53Google Scholar
  48. Kellogg RM, Wangaard FF (1969) Variation in the cell wall density of wood. Wood Fibre Sci 1:180–204Google Scholar
  49. Kramer EM (2002) A mathematical model of pattern formation in the vascular cambium of trees. J Theor Biol 216:147–158CrossRefPubMedGoogle Scholar
  50. Kutschera U (2004) The biophysical basis of cell elongation and organ maturation in coleoptiles of rye seedlings: implications for shoot development. Plant Biol 6:158–164CrossRefPubMedGoogle Scholar
  51. Lachaud S (1989) Participation of auxin and abscisic acid in the regulation of seasonal variations in cambial activity and xylogenesis. Trees 3:125–137CrossRefGoogle Scholar
  52. Lachenbruch B, Moore J, Evans R (2011) Radial variation in wood structure and function in woody plants, and hypotheses for its occurrence. In: Meinzer FC, Lachenbruch B, Dawson TE (eds) Size- and age-related changes in tree structure and function, vol 4. Springer, Netherlands, pp 121–164CrossRefGoogle Scholar
  53. Landsberg JJ, Sands P (2010) Physiological ecology of forest production: principles, processes and models. Academic Press, London, p 352Google Scholar
  54. Larson P (1969) Wood formation and the concept of wood quality. Yale University, New Haven, p 54Google Scholar
  55. Larson P (1994) The vascular cambium: development and structure. Springer-Verlag, New YorkCrossRefGoogle Scholar
  56. Lasserre J-P, Mason EG, Watt MS, Moore JR (2009) Influence of initial planting spacing and genotype on microfibril angle, wood density, fibre properties and modulus of elasticity in Pinus radiata D. Don corewood. For Ecol Manag 258:1924–1931CrossRefGoogle Scholar
  57. Li X, Wu HX, Southerton SG (2012) Identification of putative candidate genes for juvenile wood density in Pinus radiata. Tree Physiol 32:1046–1057CrossRefPubMedGoogle Scholar
  58. Lloyd C (2006) Microtubules make tracks for cellulose. Science 312:1482–1483CrossRefPubMedGoogle Scholar
  59. Lupi C, Rossi S, Vieira J, Morin H, Deslauriers A (2014) Assessment of xylem phenology: a first attempt to verify its accuracy and precision. Tree Physiol 34:87–93CrossRefPubMedGoogle Scholar
  60. Meicenheimer RD, Larson P (1983) Empirical models for xylogenesis in Populus deltoides. Ann Bot Lond 51:491–502Google Scholar
  61. Meinzer FC, Bond BJ, Karanian JA (2008) Biophysical constraints on leaf expansion in a tall conifer. Tree Physiol 28:197–206CrossRefPubMedGoogle Scholar
  62. Nonami H, Boyer JS (1989) Turgor and growth at low water potentials. Plant Physiol 89:798–804PubMedCentralCrossRefPubMedGoogle Scholar
  63. Oda Y, Fukuda H (2012) Secondary cell wall patterning during xylem differentiation. Curr Opin Plant Biol 15:38–44CrossRefPubMedGoogle Scholar
  64. O’Hehir JF, Nambiar EKSN (2010) Productivity of three successive rotations of P. radiata plantations in South Australia over a century. For Ecol Manag 259:1857–1869CrossRefGoogle Scholar
  65. Panteris E, Adamakis I-DS, Daras G, Hatzopoulos P, Rigas S (2013) Differential responsiveness of cortical microtubule orientation to suppression of cell expansion among the developmental zones of Arabidopsis thaliana root apex. PLoS One 8:e82442PubMedCentralCrossRefPubMedGoogle Scholar
  66. Paulina Fernández M, Norero A, Barthélémy D, Vera J (2007) Morphological trends in main stem of Pinus radiata D. Don: transition between vegetative and reproductive phase. Scand J For Res 22:398–406CrossRefGoogle Scholar
  67. Philipson WR, Ward JM, Butterfield BG (1971) The vascular cambium: its development and activity. Chapman and Hall, LondonGoogle Scholar
  68. Pinkard EA, Bruce J (2011) Climate change and South Australia’s plantations: impacts, risks and options for adaptation. Department of Primary Industries of South Australia. http://pir.sa.gov.au/__data/assets/pdf_file/0011/233984/ForestrySA_impact_and_adaptation_report_FINAL_July_2011.pdf
  69. Plomion C, Leprovost G, Stokes A (2001) Wood formation in trees. Plant Physiol 127:1513–1523PubMedCentralCrossRefPubMedGoogle Scholar
  70. Ridoutt BG, Sands R (1993) Within-tree variation in cambial anatomy and xylem cell differentiation in Eucalyptus globulus. Trees 8:18–22CrossRefGoogle Scholar
  71. Ridoutt BG, Sands R (1994) Quantification of the processes of secondary xylem fibre development in Eucalyptus globulus at two height levels. IAWA J 15:417–424CrossRefGoogle Scholar
  72. Rossi S, Anfodillo T, Menardi R (2006a) Trephor: a new tool for sampling microcores from tree stems. IAWA J 27:89–97CrossRefGoogle Scholar
  73. Rossi S, Deslauriers A, Anfodillo T (2006b) Assessment of cambial activity and xylogenesis by microsampling tree species: an example at the alpine timberline. IAWA J 27:383–394CrossRefGoogle Scholar
  74. Rossi S, Deslauriers A, Anfodillo T, Morin H, Saracino A, Motta R, Borghetti M (2006c) Conifers in cold environments synchronize maximum growth rate of tree-ring formation with day length. New Phytol 170:301–310CrossRefPubMedGoogle Scholar
  75. Rossi S, Deslauriers A, Griçar J, Seo J-W, Rathgeber CBK, Anfodillo T, Morin H, Levanic T, Oven P, Jalkanen R (2008) Critical temperatures for xylogenesis in conifers of cold climates. Glob Ecol Biogeogr 17:696–707CrossRefGoogle Scholar
  76. Rossi S, Simard S, Rathgeber C, Deslauriers A, De Zan C (2009) Effects of a 20-day-long dry period on cambial and apical meristem growth in Abies balsamea seedlings. Trees Struct Funct 23:85–93CrossRefGoogle Scholar
  77. Sands PJ (2004) 3PGPJS vsn 2.4—a user-friendly interface to 3-PG, the Landsberg and Waring model of forest productivity. Cooperative Research Centre for Sustainable Production Forestry, HobartGoogle Scholar
  78. Sauter JJ (1980) Seasonal variation of sucrose content in the xylem sap of salix. Z für Pflanzenphysiol 98:377–391CrossRefGoogle Scholar
  79. Sauter JJ (2000) Photosynthate allocation to the vascular cambium: facts and problems. In: Savidge R, Barnett JR, Napier R (eds) Cell and molecular biology of wood formation. BIOS Scientific Publishers, Oxford, pp 71–83Google Scholar
  80. Savidge RA, Wareing PF (1981) A tracheid-differentiation factor from pine needles. Planta 153:395–404CrossRefPubMedGoogle Scholar
  81. Shepherd KR (1964) Some observations on the effect of drought on the growth of Pinus radiata D. Don. Aust For 28:7–22CrossRefGoogle Scholar
  82. Skene DS (1969) The period of time taken by cambial derivatives to grow and differentiate into tracheids in Pinus radiata. Ann Bot Lond 33:253–262Google Scholar
  83. Skene DS (1972) The kinetics of tracheid development in Tsuga canadensis and its relation to tree vigour. Ann Bot Lond 36:179–187Google Scholar
  84. Steppe K, Lemeur R (2007) Effects of ring-porous and diffuse-porous stem wood anatomy on the hydraulic parameters used in a water flow and storage model. Tree Physiol 27:43–52CrossRefPubMedGoogle Scholar
  85. Uggla C, Magel E, Moritz T, Sundberg B (2001) Function and dynamics of auxin and carbohydrates during earlywood/latewood transition in scots pine. Plant Physiol 125:2029–2039PubMedCentralCrossRefPubMedGoogle Scholar
  86. Vaganov EA, Hughes MK, Shashkin AV (2006) Growth dynamics of conifer tree rings: an image of past and future environments. Springer-Verlag, New YorkGoogle Scholar
  87. Vavrčík H, Gryc V, Vichrová G (2013) Xylem formation in young Norway spruce trees in Drahany Highland, Czech Republic. IAWA J. 34:231–234CrossRefGoogle Scholar
  88. Walcroft AS, Silvester WB, Whitehead D, Kelliher FM (1997) Seasonal changes in stable carbon isotope ratios within annual rings of Pinus radiata reflect environmental regulation of growth processes. Funct Plant Biol 24:57–68Google Scholar
  89. Wardrop AB (1981) Lignification and xylogenesis. In: Barnett JR (ed) Xylem cell development. Castle House Publications, KentGoogle Scholar
  90. Wardrop AB, Harada H (1965) The formation and structure of the cell wall in fibres and tracheids. J Exp Bot 16:356–371CrossRefGoogle Scholar
  91. Wilson BF (1964) A model of cell production by the cambium of conifers. In: Zimmerman MH (ed) The formation of wood in forest trees. Academic Press, New York, pp 19–36CrossRefGoogle Scholar
  92. Wilson BF, Howard RA (1968) A computer model for cambial activity. For Sci 14:77–90Google Scholar
  93. Wimmer R, Downes GM, Evans R (2002) Temporal variation of microfibril angle in Eucalyptus nitens grown in different irrigation regimes. Tree Physiol 22:449–457CrossRefPubMedGoogle Scholar
  94. Winship LJ, Obermeyer G, Geitmann A, Hepler PK (2010) Under pressure, cell walls set the pace. Trends Plant Sci 15:363–369PubMedCentralCrossRefPubMedGoogle Scholar
  95. Woodruff DR, Bond BJ, Meinzer FC (2004) Does turgor limit growth in tall trees? Plant Cell Environ 27:229–236CrossRefGoogle Scholar
  96. Wu HX, Ivković M, Gapare WJ, Baltunis BS, Powell MB, McRae TA (2008) Breeding for wood quality and profit in radiata pine: a review of genetic parameters. N Z J For Sci 38(1)Google Scholar
  97. Zhang J, Nieminen K, Serra JAA, Helariutta Y (2014) The formation of wood and its control. Curr Opin Plant Biol 17:56–63CrossRefPubMedGoogle Scholar
  98. Zweifel R, Zimmerman L, Newbery DM (2005) Modeling tree water deficit from microclimate: an approach to quantifying drought stress. Tree Physiol 25:147–156CrossRefPubMedGoogle Scholar
  99. Zweifel R, Steppe K, Sterck FJ (2007) Stomatal regulation by microclimate and tree water relations: interpreting ecophysiological field data with a hydraulic plant model. J Exp Bot 58:2113–2131CrossRefPubMedGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2015

Authors and Affiliations

  1. 1.CSIRO Sustainable Agriculture FlagshipHobartAustralia
  2. 2.Forest Quality Pty. Ltd.HuonvilleAustralia

Personalised recommendations