, Volume 27, Issue 4, pp 973–983 | Cite as

Figured grain in aspen is heritable and not affected by graft-transmissible signals

  • Youran Fan
  • Kendal Rupert
  • Alex C. Wiedenhoeft
  • Keith Woeste
  • Christian Lexer
  • Richard MeilanEmail author
Original Paper


Figure can add value to wood products, but its occurrence is unpredictable. A first step in reliably producing figured wood is determining whether it is faithfully transmitted to progeny via sexual and asexual reproduction. We describe a 26-year-old male aspen genotype, designated ‘Curly Poplar’, which was shown to be a Populus × canescens hybrid using microsatellite markers. All rooted cuttings of this genotype exhibited an undulating pattern on the radial surface that was not seen in the control trees, all of which showed a smooth radial surface and straight grain. We observed spiral grain with a magnitude of 2.77 ± 0.12°/cm from vertical in 11-month-old, field-grown rooted Curly Poplar cuttings, but spiral grain was not apparent in wood from the 26-year-old mature ortet that supplied these cuttings. Veneer cut from the mature tree exhibited a novel type of figure that we called ‘Scattered Moiré’. Reciprocal grafts between Curly Poplar and various non-figured aspens showed that a graft-transmissible signals did not appear to be involved in figure formation in Curly Poplar or the induction of figure in straight-grained trees. Curly Poplar was crossed to a straight-grained clone to test the inheritance of the gene(s) responsible for figure. Samples from the resulting population revealed that 79 out of 377 seedlings exhibited figure. A Chi-square test led to the rejection of a 1:1 segregation ratio between figured and non-figured phenotypes (p < 0.01), but not of a 1:3 segregation ratio (p 0.0793). Overall, these analyses showed that figure in Curly Poplar is under genetic control, but its inheritance may not be simple.


Curly Poplar Scattered Moiré Figured wood Heritability Populus × canescens Wood quality 



We are grateful to Dr. Richard Hall (Iowa State University) who provided Populus cultivars Crandon, Sherrill, and 4877; and Dr. Michael Cunningham (ArborGen, Inc., Summerville, SC), who provided Ca-2-75, the straight-grained P. × canescens. We also thank Mr. Patrick McGovern for producing seeds for the F1 population, and providing the female parent, A502. For their assistance with field work, we thank James McKenna, Brian Beheler, Burk Thompson, Stuart Christopher, and William Skrobutt. This project was partially funded by generous donations from Dr. Samuel Grober and his son, David. Additional support was provided by the Hardwood Tree Improvement and Regeneration Center, Northern Research Station, USDA Forest Service, and by a grant from the British Natural Environment Research Council (NERC).

Conflict of interest

The authors declare that they have no conflict of interest. The use of trade names is for the information and convenience of the reader and does not imply official endorsement or approval by the United States Department of Agriculture or the Forest Service of any product to the exclusion of others that may be suitable.


  1. Anonymous (1929) What causes “bird’s-eye” maple. Technical Note 13. USDA Forest Service, Lake State Forest Experiment Station, St. PaulGoogle Scholar
  2. Bao FC, Jiang ZH, Jiang XM, Lu XX, Luo XQ, Zhang SY (2001) Differences in wood properties between juvenile wood and mature wood in 10 species grown in China. Wood Sci Technol 35:363–375CrossRefGoogle Scholar
  3. Beals HO, Davis TC (1977) Figure in wood: an illustrated review. Bulletin 486:88 (Auburn University Agriculture Experiment Station, Auburn University, Auburn)Google Scholar
  4. Boyce SG, Kaiser M (1961) Environmental and genetic variability in the length of fibers of eastern cottonwood. Tappi J 44:363–366Google Scholar
  5. Bragg DC (1999) The birdseye figured grain in sugar maple (Acer saccharum): literature review, nomenclature, and structural characteristics. Can J For Res 29:1637–1648CrossRefGoogle Scholar
  6. Bragg DC, Stokke DD (1993) Field identification of birdseye in sugar maple (Acer Saccharum Marsh). Research Paper NC-317. USDA Forest Service, North Central Forest Experiment Station, St. PaulGoogle Scholar
  7. Bragg DC, Mroz GD, Reed DD, Shetron SG, Stokke DD (1997) Relationship between ‘‘birdseye’’ sugar maple (Acer saccharum) occurrence and its environment. Can J For Res 27:1182–1191CrossRefGoogle Scholar
  8. Brazier JD, Mobbs ID (1993) The influence of planting distance on structural wood yields of unthinned Sitka spruce. Forestry 66:333–352CrossRefGoogle Scholar
  9. Burke J, Arnold M (2001) Genetics and fitness of hybrids. Annu Rev Genet 35:31–52PubMedCrossRefGoogle Scholar
  10. Cameron AD, Lee SL, Livingston AK, Petty JA (2005) Influence of selective breeding on the development of juvenile wood in Sitka spruce. Can J For Res 35:2951–2960CrossRefGoogle Scholar
  11. Campbell MM, Brunner AM, Jones HM, Strauss SH (2003) Forestry’s fertile crescent: the application of biotechnology to forest trees. Plant Biotech J 1:141–154CrossRefGoogle Scholar
  12. Chaffey N (1999) Wood formation in forest trees: from Arabidopsis to Zinnia. Trends Plant Sci 4:203–204PubMedCrossRefGoogle Scholar
  13. Cheng WW, Bensend DW (1979) Anatomical properties of selected Populus clones grown under intensive culture. Wood Sci 11:182–187Google Scholar
  14. Eklund L, Säll H (2000) The influence of wind on spiral grain formation in conifer trees. Trees Struct Funct 14:324–328CrossRefGoogle Scholar
  15. Eklund L, Säll H, Linder S (2003) Enhanced growth and ethylene increases spiral grain formation in Picea abies and Abies balsamea trees. Trees Struct Funct 17:81–86CrossRefGoogle Scholar
  16. Farmer R (1964) Sex ratio and sex-related characteristics in eastern cottonwood. Silvae Genetica 13:116–118Google Scholar
  17. Fussi B, Lexer C, Heinze B (2010) Phylogeography of Populus alba (L.) and Populus tremula (L.) in Central Europe: secondary contact and hybridisation during recolonisation from disconnected refugia. Tree Genet Genomes 6:439–450CrossRefGoogle Scholar
  18. Gehring M, Choi Y, Fischer RL (2004) Imprinting and seed development. Plant Cell 16:S203–S213PubMedCrossRefGoogle Scholar
  19. Grant M, Mitton J (1979) Elevational gradients in adult sex-ratios and sexual-differentiation in vegetative growth-rates of Populus tremuloides. Evol Int J Org Evol 33:914–918CrossRefGoogle Scholar
  20. Grober S (1942) The botanical, erosion control, and economic significance of white poplar in Maryland. University of Maryland, DissertationGoogle Scholar
  21. Harada T (2010) Grafting and RNA transport via phloem tissue in horticultural plants. Sci Hort 125:545–550CrossRefGoogle Scholar
  22. Harris JM (1989) Spiral grain and wave phenomena in wood formation. Springer-Verlag, New YorkCrossRefGoogle Scholar
  23. Hartmann HT, Kester DE, Davies FT, Geneve RL Jr (2002) Plant propagation: principles and practices, 7th edn. Prentice-Hall Inc, Englewood CliffsGoogle Scholar
  24. Heinkinheimo O (1940) The cultivation of figured birch. Skogen 27:165–167Google Scholar
  25. Jansson S, Douglas CJ (2007) Populus: a model system for plant biology. Annu Rev Plant Biol 58:435–458PubMedCrossRefGoogle Scholar
  26. Kalev N, Aloni R (1999) Role of ethylene and auxin in regenerative differentiation and orientation of tracheids in Pinus pinea seedlings. New Phytol 142:307–313CrossRefGoogle Scholar
  27. Kasai A, Bai S, Li T, Harada T (2011) Graft-transmitted siRNA signal from the root induces visual manifestation of endogenous post-transcriptional gene silencing in the scion. PLoS One 6:e16895PubMedCrossRefGoogle Scholar
  28. Kramer EM (2006) Wood grain pattern formation: a brief review. J Plant Growth Regul 25:290–301CrossRefGoogle Scholar
  29. Kramer EM (2009) Auxin-regulated cell polarity: an inside job? Trends Plant Sci 14:242–247PubMedCrossRefGoogle Scholar
  30. Kramer EM, Lewandowski M, Beri S, Bernard J, Borkowski M, Borkowski MH, Burchfield LA, Mathisen B, Normanly J (2008) Auxin gradients are associated with polarity changes in trees. Science 320:1610PubMedCrossRefGoogle Scholar
  31. Kutner M, Nachtsheim C, Neter J, Li W (2004) Applied linear statistical models, 5th edn. McGraw-Hill/Irwin, New YorkGoogle Scholar
  32. Lescot M, Rombauts S, Zhang J, Aubourg S, Mathé C, Jansson S, Rouzé P, Boerjan W (2004) Annotation of a 95-kb Populus deltoides genomic sequence reveals a disease resistance gene cluster and novel class I and class II transposable elements. Theor Appl Genet 109:10–22PubMedCrossRefGoogle Scholar
  33. Lexer C, Fay MF, Joseph JA, Nica M-S, Heinze B (2005) Barrier to gene flow between two ecologically divergent Populus species, P. alba (white poplar) and P. tremula (European aspen): the role of ecology and life history in gene introgression. Mol Ecol 14:1045–1057PubMedCrossRefGoogle Scholar
  34. Lexer C, Joseph JA, van Loo M, Barbará T, Heinze B, Bartha D, Castiglione S, Fay MF, Buerkle CA (2010) Genomic admixture analysis in European Populus spp. reveals unexpected patterns of reproductive isolation and mating. Genetics 186:699–712PubMedCrossRefGoogle Scholar
  35. Macaya-Sanz D, Suter L, Joseph J, Barbara T, Alba N, Gonzalez-Martinez SC, Widmer A, Lexer C (2011) Genetic analysis of post-mating reproductive barriers in hybridizing European Populus species. Heredity 107:478–486PubMedCrossRefGoogle Scholar
  36. MacDaniels LH (1953) Some aspects of the problem of producing curly-grained walnut. In: Proceedings of the 44th annual report of the northern nut growers association, RochesterGoogle Scholar
  37. Matyas C, Peszlen I (1997) Effect of age on selected wood quality traits of poplar clones. Silvae Genetica 46:64–72Google Scholar
  38. Mellerowicz EJ, Baucher M, Sundberg B, Boerjan W (2001) Unravelling cell wall formation in the woody dicot stem. Plant Mol Biol 47:239–274PubMedCrossRefGoogle Scholar
  39. Michalak P (2009) Epigenetic, transposon and small RNA determinants of hybrid dysfunctions. Heredity 102:45–50PubMedCrossRefGoogle Scholar
  40. Panshin A, Zeeuw C (1980) Textbook of wood technology: structure, identification, properties, and uses of the commercial woods of the United States and Canada, 4th edn. McGraw-Hill Inc, New YorkGoogle Scholar
  41. Persson A (1954) Seed orchard production of figured birch. Skogen 41:160–163Google Scholar
  42. Pillow MY (1955) Detection of figured wood in standing trees. Research Paper No. 2034. USDA Forest Service, Forest Products Laboratory, MadisonGoogle Scholar
  43. Pritchard JK, Stephens M, Donnelly P (2000) Inference of population structure using multilocus genotype data. Genetics 155:945–959PubMedGoogle Scholar
  44. Rauchfuss J, Speer JH (2006) Age dependence of spiral grain in white oaks (Quercus alba L.) in southwestern Illinois. Tree Ring Res 62:13–14CrossRefGoogle Scholar
  45. Rieseberg L, Carney S (1998) Plant hybridization. New Phytol 140:599–624CrossRefGoogle Scholar
  46. Righter FI (1934) On the cause of bird’s-eye maple. J For 32:626–627Google Scholar
  47. Rioux D, Yamada T, Simard M, Lessard G, Rheault FJ, Biouin D (2003) Contribution to the fine anatomy and histochemistry of birdseye sugar maple. Can J For Res 33:946–958CrossRefGoogle Scholar
  48. Robbins PW (1953) Forest genetics problems needing study in Michigan-minor products (maple syrup). In: Proceedings of the Lake States forest genetics conference, vol. 22. USDA Forest Service Lake States Forest Experiment Station Misc. Report, Eagle River, pp 59–60Google Scholar
  49. Rottenberg A, Nevo E, Zohary D (2000) Genetic variability in sexually dimorphic and monomorphic populations of Populus euphratica (Salicaceae). Can J For Res 30:482–486Google Scholar
  50. Rudolf PO (1954) Forest tree improvement work in the Lake States. J For 52:688–689Google Scholar
  51. Stace CA (1997) New flora of the British Isles, 2nd edn. Cambridge University Press, CambridgeGoogle Scholar
  52. Stettler RF, Bradshaw HD, Heilman PE, Hinckley TM (1996) Biology of Populus and its implications for management and conservation. NRC Research Press, National Research Council of Canada, OttawaGoogle Scholar
  53. Taylor G (2002) Populus: Arabidopsis for forestry. Do we need a model tree? Ann Bot 90:681–689PubMedCrossRefGoogle Scholar
  54. Turnbull CGN, Booker JP, Leyser HMO (2002) Micrografting techniques for testing long-distance signalling in Arabidopsis. Plant J 32:255–262PubMedCrossRefGoogle Scholar
  55. Tuskan GA, DiFazio SP, Jansson S et al (2006) The genome of black cottonwood, Populus trichocarpa (Torr. & Gray). Science 313:1596–1604PubMedCrossRefGoogle Scholar
  56. Václav E (1969) The quality of wood of the technical forms of birch. In: Proceedings of the FAO and IUFRO second world consultation on forest tree breeding. IUFRO, Vienna, pp 438–446 (Publ. No. FO-FTB-69-4/6)Google Scholar
  57. van Loo M, Joseph JA, Heinze B, Fay MF, Lexer C (2008) Clonality and spatial genetic structure in Populus × canescens and its sympatric backcross parent P. alba in a Central European hybrid zone. New Phytol 177:506–516PubMedGoogle Scholar
  58. Wullschleger SD, Jansson S, Taylor G (2002) Genomics and forest biology: Populus emerges as the perennial favorite. Plant Cell 14:2651–2655PubMedCrossRefGoogle Scholar
  59. Yin TM, DiFazio SP, Gunter LE, Riemenschneider D, Tuskan GA (2004) Large-scale heterospecific segregation distortion in Populus revealed by a dense genetic map. Theor Appl Genet 109:451–463PubMedCrossRefGoogle Scholar
  60. Zobel BJ, Sprague JR (1998) Juvenile wood in forest trees. Springer-Verlag, BerlinCrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  • Youran Fan
    • 1
  • Kendal Rupert
    • 1
  • Alex C. Wiedenhoeft
    • 2
  • Keith Woeste
    • 1
    • 3
  • Christian Lexer
    • 4
  • Richard Meilan
    • 1
    Email author
  1. 1.Department of Forestry and Natural ResourcesPurdue UniversityWest LafayetteUSA
  2. 2.USDA Forest ServiceForest Products LaboratoryMadisonUSA
  3. 3.USDA Forest Service, Northern Research StationHardwood Tree Improvement and Regeneration CenterWest LafayetteUSA
  4. 4.Department of BiologyUniversity of FribourgFribourgSwitzerland

Personalised recommendations