Does Photosynthetic Bark have a Role in the Production of Core vs. Outer Wood?
Keywords:
Core wood, crown wood, wood quality, tracheid length, fiber length, periderm, tracheid, juvenile wood, cambium, phellogenAbstract
This paper hypothesizes a correlation in some species between the cambial age of transition from core (juvenile) to outer (mature) wood and the cambial age of transition from photosynthetic to non-photosynthetic bark. Secondly, this paper hypothesizes that the relationship is causal: a signal produced in relation to the photosynthetic bark affects wood development a few millimeters away. It is further hypothesized that the photosynthetic periderm is replaced by a non-photosynthetic one at light levels below its light compensation point. In T'suga heterophylla and Pseudotsuga menziesii var. menziesii, the cambial age at which the first periderm dies (the base of photosynthetic bark) ranges from 16 to 33 and 12 to 43 years, respectively, for four Oregon Coast Range populations. These values are in the same range as the cambial ages of transition from core to outer wood, as shown by literature values and data reported here on tracheid length in T. heterophylla. In both species, the cambial age at the base of the live crown is not coincident with, nor consistently higher or lower than, the height of the lowest photosynthetic bark. Data presented here are consistent with the photosynthetic bark hypothesis of formation of core wood, but manipulative studies are needed to further explore the relationship.References
Abdel-Gadir, A. Y., and R. L. Krahmer. 1993. Genetic variation in the age of demarcation between juvenile and mature wood in Douglas-fir. Wood Fiber Sci. 25(4):384-394.nAbdel-Gadir, A. Y., R. L. Krahmer, and M. D. McKimmy. 1993. Estimating the age of demarcation of juvenile and mature wood in Douglas-fir. Wood Fiber Sci. 25(3):242-249.nAloni, R., and M. H. Zimmermann. 1983. The control of vessel size and density along the plant axis: A new hypothesis. Differentiation 24:203-208.nBallaré, C. L., A. L. Scopel, and R. A. Sanchez. 1990. Far-red radiation reflected from adjacent leaves: An early signal of competition in plant canopies. Science 247: 329-332.nBorger, G. A. 1973. Development and shedding of bark. Pages 205-236 in T. T. Kozlowski, ed. Shedding of plant parts. Academic Press, New York, NY.nClark, A., III, and J. R. Saucier. 1989. Influence of initial planting density, geographic location, and species on juvenile wood formation in southern pine. Forest Prod. J. 39:42-48.nComstock, J., and J. Ehleringer. 1990. Effect of variations in leaf size on morphology and photosynthetic rate of twigs. Funct. Ecol. 4:209-221.nCown, D. J. 1992. Corewood (juvenile wood) in Pinus radiata — Should we be concerned? N.z. J. For. Sci. 22: 87-95.nCown, D. J., and D. L. McConchie. 1980. Wood property variations in an old-crop stand of radiata pine. N.Z. J. For. Sci. 10:508-520.nCown, D. J., and M. L. Parker. 1979. Densitometric analysis of wood from five Douglas-fir provenances. Silvae Genetica 28:48-53.nDi Lucca, C. M. 1989. Juvenile-mature wood transition. Pages 23-38 in R. M. Kellogg, ed. Second growth Douglas-fir: Its management and conversion for value; a report of the Douglas-fir task force. Special Publication No. SP-32. Forintek Canada Corp, Vancouver, BC.nFoote, K. C., and M. Schaedle. 1978. The contribution of aspen bark photosynthesis to the energy balance of the stem. Forest Sci. 24:569-573.nFukazawa, K. 1984. Juvenile wood of hardwoods judged by density variation. IAWA Bull. n.s. 5:65-73.nKrahmer, R. L. 1966. Variation of specific gravity in western hemlock trees. Tappi 49:227-229.nLarcher, W. 1980. Physiological plant ecology. Springer-Verlag, Berlin, Germany.nLarson, P. R. 1962. A biological approach to wood quality. Tappi 45:443-448.nLarson, P. R. 1969. Wood formation and the concept of wood quality. Yale University School of Forestry Bull. 74:1-54. New Haven, CT.nLev-Yadun, S., and R. Aloni. 1993. Bark structure and mode of canopy regeneration in trees of Melia azedarach L. Trees 7:144-147.nLittle, C. H. A., and R. P. Pharis. 1995. Hormonal control of radial and longitudinal growth in the tree stem. Pages 281-319 in B. L. Gartner, ed. Plant stems: Physiology and functional morphology. Academic Press, San Diego, CA.nMaguire, D. A., J. A. Kershaw, Jr., and D. W. Hann. 1991. Predicting the effects of silvicultural regime on branch size and crown wood core in Douglas-fir. Forest Sci. 37:1409-1428.nMcKimmy, M. D., and R. K. Campbell. 1982. Genetic variation in the wood density and ring width trend in coastal Douglas-fir. Silvae Genetica 31:43-51.nMegraw, R. A. 1985. Wood quality factors in loblolly pine: The influence of tree age, position in tree, and cultural practice on wood specific gravity, fiber length, and fibril angle. Tappi Press, Atlanta, GA.nOlesen, P. O. 1982. The effect of cyclophysis on tracheid width and basic density in Norway spruce. Forest Tree Improv. 15: 1-80.nRasband, W. 1992. Image, Version 1.47. National Institutes of Health, Research Services Branch, NIMH, Bethesda, MD.nSandberg, G., P. Gardeström, F. Sitbon, and O. Olsson. 1990. Presence of indole-3-acetic acid in chloroplasts of Nicotiana tabacum and Pinus sylvestris. Planta 180: 562-568.nSchaedle, M. 1975. Tree photosynthesis. Ann. Rev. Plant Physiol. 26:101-115.nSenft, J. F., M. J. Quanci, and B. A. Bendtsen. 1985. Property profile of 60-year-old Douglas-fir. Pages 17-28 in Juvenile wood — What does it mean to forest management and forest products? Proceedings 47309. Forest Products Research Society, Madison, WI.nSrivastava, L. M. 1964. Anatomy, chemistry, and physiology of bark. Int. Rev. For. Res. 1:203-277.nSundberg, B., A. Ericsson, C. H. A. Little, T. Näsholm, and R. Gref. 1993. The relationship between crown size and ring width in Pinus sylvestris L. stems: Dependence on indole-3-acetic acid, carbohydrates, and nitrogen in the cambial region. Tree Physiol. 12:347-362.nVargas-Hernandez, J., W. T. Adams, and R. L. Krahmer. 1994. Family variation in age trends of wood density traits in young coastal Douglas-fir. Wood Fiber Sci. 26(2):229-236.nYang, K. C., C. A. Benson, and J. K. Wong. 1986. Distribution of juvenile wood in two stems of Larix laricina. Can. J. Forest Res. 16:1041-1049.nYang, K. C., Y. S. Chen, and C. Chiu. 1994. Formation and vertical distribution of juvenile and mature wood in a single stem of Cryptomeria japonica. Can. J. Forest Res. 24:969-975.nZobel, B. J., and J. P. Van Buijtenen. 1989. Wood variation: Its causes and control. Springer-Verlag, Berlin, Germany.n
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