Skip to main content

Official Journal of the Japan Wood Research Society

  • Original Article
  • Published:

Ultrastructure of the S2 layer in relation to lignin distribution inPinus radiata tracheids

Abstract

The ultrastructure of the S2 layer in relation to its lignin distribution was examined using transmission electron microscopy in the tracheids ofPinus radiata. The S2 layer had a striated appearance at low magnification. Observations at higher magnifications showed lignin to be distributed inhomogeneously in this layer, appearing as a mosaic of electron-dense and electron-lucent regions. These regions are scattered, showing a pattern of often interconnecting sinuous features in a predominantly radial profile. The significance of these features of the S2 layer is discussed, particularly in relation to the available information from recent ultrastructural observations on the appearance of cellulose microfibrils and the pattern of their distribution in the S2 layer using rapid freeze-deep etching in conjunction with transmission electron microscopy. Predictions are made as to the likely distribution and arrangement of cellulose microfibrils in the S2 layer based on the pattern of lignin distribution observed in this layer.

References

  1. Fergus BJ, Goring DAI (1970) The distribution of lignin in birch wood as determined by ultraviolet microscopy. Holzforschung 24:118–124

    Article  CAS  Google Scholar 

  2. Fukazawa K (1974) The distribution of lignin in compression- and lateral-wood ofAbies sachalinensis using ultraviolet microscopy. Res Bull Coll Exp For Hokkaido Univ 31:87–114

    Google Scholar 

  3. Saka S, Thomas RJ (1982) Evaluation of the quantitative assay of lignin distribution by SEM-EDXA technique. Wood Sci Technol 16:1–18

    Article  CAS  Google Scholar 

  4. Donaldson LA (1985) Within- and between-tree variation in lignin concentration in the tracheid cell wall ofPinus radiata. NZ J For Sci 15:361–369

    CAS  Google Scholar 

  5. Westermark U (1985) Bromination of different morphological parts of spruce wood (Picea abies Karst). Wood Sci Technol 19:323–328

    Article  CAS  Google Scholar 

  6. Saka S, Goring DAI (1988) The distribution of lignin in white birch wood as determined by bromination with TEM-EDXA. Holzforschung 42:149–153

    Article  CAS  Google Scholar 

  7. Daniel GF, Nilsson T, Pettersson B (1991) Poorly and non-lignified regions in the middle lamella cell corners of birch (Betula verrucosa) and other wood species. IAWA Bull 12:70–83

    Article  Google Scholar 

  8. Downes GM, Ward JV, Turvey ND (1991) Lignin distribution across tracheid cell walls of poorly lignified wood from deformed copper deficientPinus radiata (D Don). Wood Sci Technol 25:7–14

    Article  CAS  Google Scholar 

  9. Donaldson LA, Singh AP, Yoshinaga A, Takabe K (1999) Lignin distribution in mild compression wood ofPinus radiata D Don. Can J Bot 77:41–50

    CAS  Google Scholar 

  10. Singh AP, Donaldson LA (1999) Ultrastructure of tracheid cell walls in radiata pine (Pinus radiata) mild compression wood. Can J Bot 77:32–40

    Google Scholar 

  11. Kerr AJ, Goring DAI (1975) The ultrastructural arrangement of the wood cell wall. Cellulose Chem Technol 9:563–573

    Google Scholar 

  12. Daniel GF, Nilsson T (1984) Studies on the S2 layer ofPinus sylvestris. Report 154. Department of Forest Products, Swedish University of Agricultural Sciences, Uppsala, Sweden, p 34

    Google Scholar 

  13. Singh AP (1997) The ultrastructure of the attack ofPinus radiata mild compression wood by erosion and tunnelling bacteria. Can J Bot 75:1095–1102

    Article  Google Scholar 

  14. Singh AP, Sell J, Schmitt U, Zimmermann T, Dawson B (1998) Radial striation of the S2 layer in mild compression wood tracheids. ofPinus radiata. Holzforschung 52:563–566

    Article  CAS  Google Scholar 

  15. Larsen MJ, Winandy JE, Green F (1995) A proposed model of the tracheid cell wall of southern yellow pine having an inherent radial structure in the S2 layer. Mater Org 29:197–210

    Google Scholar 

  16. Sell J, Zimmermann T (1993) Radial fibril agglomerations on the S2 on transverse fracture surfaces of tracheids of tension-loaded spruce and white fir. Holz Roh Werkstoff 51:384

    Article  Google Scholar 

  17. Spurr AR (1969) A low viscosity embedding medium for electron microscopy. J Ultrastruct Res 26:31–43

    Article  CAS  PubMed  Google Scholar 

  18. Ruel K, Barnoud F, Goring DAI (1978) Lamellation in the S2 layer of softwood tracheids as demonstrated by scanning transmission electron microscopy. Wood Sci Technol 12:287–291

    Article  Google Scholar 

  19. Tirumalai VC, Agarwal UP, Obst JR (1996) Heterogeneity of lignin concentration in cell corner middle lamella of white birch and black spruce. Wood Sci Technol 30:99–104

    Article  CAS  Google Scholar 

  20. Singh AP, Schmitt U (2000) High variability in the distribution of lignin in the middle lamella of rubber wood (Hevea brasiliensis). In: Kim YS (ed) Proceedings of the 4th Pacific Regional Wood Anatomy Conference, Chonnam National University, Kwangju, South Korea, pp 203–207

    Google Scholar 

  21. Hepler PK, Fosket DE, Newcomb EH (1970) Lignification during secondary wall formation in Coleus: an electron microscope study. Am J Bot 57:85–96

    Article  Google Scholar 

  22. Maurer A, Fengel D (1990) A process for improving the quality and lignin staining of ultrathin sections from wood tissue. Holzforschung 44:453–460

    Article  CAS  Google Scholar 

  23. Bland DE, Foster RC, Logan AF (1971) The mechanism of permanganate and osmium tetroxide fixation and the distribution of lignin in the cell wall ofPinus radiata. Holzforschung 25:137–168

    Article  CAS  Google Scholar 

  24. Hoffmann P, Parameswaran N (1976) On the ultrastructural localisation of hemicelluloses within lignified tracheids of spruce. Holzforschung 30:62–70

    Article  Google Scholar 

  25. Hafrén J, Fujino T, Itoh T (1999) Changes in cell wall architecture of differentiating tracheids ofPinus thunbergii during lignification. Plant Cell Physiol 40:533–541

    Article  Google Scholar 

  26. Hafrén J (1999) Ultrastructure of the wood cell wall. PhD thesis, Royal Institute of Technology, Stockholm

    Google Scholar 

  27. Donaldson LA, Singh AP (1998) Bridge-iike structures between cellulose microfibrils in radiata pine (Pinus radiata D Don) kraft pulp and holocellulose. Holzforschung 52:449–454

    Article  CAS  Google Scholar 

  28. Kataoka Y, Saiki H, Fujita M (1992) Arrangement and superimposition of cellulose microfibrils in the secondary walls of coniferous tracheids. Mokuzai Gakkaishi 38:327–335

    CAS  Google Scholar 

Download references

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Adya Singh.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Singh, A., Daniel, G. & Nilsson, T. Ultrastructure of the S2 layer in relation to lignin distribution inPinus radiata tracheids. J Wood Sci 48, 95–98 (2002). https://doi.org/10.1007/BF00767284

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/BF00767284

Key words