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  • Print publication year: 2010
  • Online publication date: June 2012

Chapter 11 - Secondary xylem

Summary

Perspective

Most of the major taxa of vascular plants produce secondary xylem derived from the vascular cambium. Pteridophytes (except some extinct taxa), most monocotyledons, and a few species of largely aquatic dicotyledons, however, produce only primary vascular tissues. In woody plants secondary xylem comprises the bulk of the tissue in the stems and roots. It is the most important supporting tissue in arborescent dicotyledons and most gymnosperms, and the major tissue for the transport of water and essential minerals in woody plants. Secondary xylem is a complex tissue that consists not only of non-living supporting and conducting cells but also of important living components (rays and axial wood parenchyma) which, with those in the secondary phloem, comprise a three-dimensional symplastic pathway through which photosynthate and other essential molecular substances are transported thoughout the secondary tissues of the plant (Chaffey and Barlow, 2001; see pp. 206–207 for more detail). Additional increments of this tissue are added during each growing season (usually annually), but in older regions of most woody species only the outer increments are functional in transport although the number of increments that remain functional varies greatly among different species. Older increments gradually become plugged by the deposition in them of waste metabolites such as resins, tannins, and in some species by the formation of tyloses (balloon-like extensions of axial or ray parenchyma cells into adjacent conducting cells). The inner non-functional secondary xylem is called heartwood, the outer functional secondary xylem, sapwood.

REFERENCES
The diploid (2n) cell that results from the fusion of male and female gametes.Abe, H., Ohtani, J., and Fukazawa, K.. 1994. A scanning electron microscopic study of changes in microtubule distributions during secondary wall formation in tracheids. Int. Ass. Wood. Anatomists J. 15: 185–189.
The diploid (2n) cell that results from the fusion of male and female gametes.Aloni, R. 1992. The control of vascular differentiation. Int. J. Plant Sci. 153: S90–S92.
The diploid (2n) cell that results from the fusion of male and female gametes.Aloni, R. 2001. Foliar and axial aspects of vascular differentiation: hypotheses and evidence. J. Plant Growth Reg. 20: 22–34.
The diploid (2n) cell that results from the fusion of male and female gametes.Bailey, I. W. 1953. Evolution of the tracheary tissue of land plants. Am. J. Bot. 40: 4–8.
The diploid (2n) cell that results from the fusion of male and female gametes.Bailey, I. W. 1954. Contributions to Plant Anatomy. Waltham, MA: Chronica Botanica.
The diploid (2n) cell that results from the fusion of male and female gametes.Bailey, I. W. 1957. The potentialities and limitations of wood anatomy in the study of the phylogeny and classification of angiosperms. J. Arnold Arbor. 38: 243–254.
The diploid (2n) cell that results from the fusion of male and female gametes.Bailey, I. W. and Tupper, W. W.. 1918. Size variation in tracheary cells. I. A comparison between the secondary xylems of vascular cryptogams, gymnosperms, and angiosperms. Proc. Amer. Acad. Arts Sci. 54: 149–204.
The diploid (2n) cell that results from the fusion of male and female gametes.Barnett, J. R. 1981. Secondary xylem cell development. In Barnett, J. R., ed., Xylem Cell Development. Tunbridge Wells:Castle House, pp. 47–95.
The diploid (2n) cell that results from the fusion of male and female gametes.Baskin, T. I. 2001. On the alignment of cellulose microfibrils by cortical microtubules: a review and a model. Protoplasma 215: 150–171.
The diploid (2n) cell that results from the fusion of male and female gametes.Baum, S. F., Aloni, R., and Peterson, C. A.. 1991. The role of cytokinin in vessel regeneration in wounded Coleus internodes. Ann. Bot. 67: 543–548.
The diploid (2n) cell that results from the fusion of male and female gametes.Beck, C. B., Coy, K., and Schmid, R.. 1982. Observations on the fine structure of Callixylon wood. Am. J. Bot. 69: 54–76.
The diploid (2n) cell that results from the fusion of male and female gametes.Canny, M. J. 1995. A new theory for the ascent of sap-cohesion supported by tissue pressure. Ann. Bot. 75: 343–357.
The diploid (2n) cell that results from the fusion of male and female gametes.Canny, M. J. 1998. Applications of the compensating pressure theory of water transport. Am. J. Bot. 85: 897–909.
The diploid (2n) cell that results from the fusion of male and female gametes.Canny, M. J. 2001. Contributions to the debate on water transport. Am. J. Bot. 88: 43–46.
The diploid (2n) cell that results from the fusion of male and female gametes.Carlquist, S. 1966. Wood anatomy of Compositae: a summary, with comments on factors affecting wood evolution. Aliso 6: 225--244.
The diploid (2n) cell that results from the fusion of male and female gametes.Carlquist, S. 1975. Ecological Strategies of Xylem Evolution. Berkeley, CA: University of California Press.
The diploid (2n) cell that results from the fusion of male and female gametes.Carlquist, S. 1988. Comparative Wood Anatomy. Berlin: Springer-Verlag.
The diploid (2n) cell that results from the fusion of male and female gametes.Carpenter, C. H. and Leney, L.. 1952. 91 Papermaking Fibers. Technical Publication No. 74. Syracuse, NY: College of Forestry, Syracuse University.
The diploid (2n) cell that results from the fusion of male and female gametes.Chaffey, N. and Barlow, P.. 2001. The cytoskeleton facilitates a three-dimensional symplasmic continuum in the long-lived ray and axial parenchyma cells of angiosperm trees. Planta 213: 811–823.
The diploid (2n) cell that results from the fusion of male and female gametes.Chaffey, N. and Barlow, P.. 2002. Myosin, microtubules and microfilaments: cooperation between cytoskeletal components during cambial cell division and secondary vascular differentiation in trees. Planta 214: 526–536.
The diploid (2n) cell that results from the fusion of male and female gametes.Chaffey, N. J., Barnett, J. R., and Barlow, P. W.. 1997. Cortical microtubule involvement in bordered pit formation in secondary xylem vessel elements of Aesculus hippocastanum L. (Hippocastanaceae): a correlative study using electron microscopy and indirect immunofluorescence microscopy. Protoplasma 197: 64–75.
The diploid (2n) cell that results from the fusion of male and female gametes.Chaffey, N. J., Barnett, J. R., and Barlow, P. W.. 1999. A cytoskeletal basis for wood formation in angiosperm trees: the involvement of cortical microtubules. Planta 208: 19–30.
The diploid (2n) cell that results from the fusion of male and female gametes.Chaffey, N., Barlow, P., and Barnett, J.. 2000. A cytoskeletal basis for wood formation in angiosperm trees: the involvement of microfilaments. Planta 210: 890–896.
The diploid (2n) cell that results from the fusion of male and female gametes.Chaffey, N., Barlow, P., and Sundberg, B.. 2002. Understanding the role of the cytoskeleton in wood formation in angiosperm trees: hybrid aspen (Populus tremula × P. tremuloides) as the model species. Tree Physiol. 22: 239–249.
The diploid (2n) cell that results from the fusion of male and female gametes.Comstock, J. P. 1999. Why Canny's theory doesn't hold water. Am. J. Bot. 86: 1077–1081.
The diploid (2n) cell that results from the fusion of male and female gametes.Core, H. A., Côté, W. A., and Day, A. C.. 1979. Wood: Structure and Identification, 2nd edn. Syracuse, NY: Syracuse University Press.
The diploid (2n) cell that results from the fusion of male and female gametes.Cosgrove, D. J. 1993. Wall extensibility: its nature, measurement and relationship to plant cell growth. New Phytol. 124: 1–23.
The diploid (2n) cell that results from the fusion of male and female gametes.Cosgrove, D. J. 2000. Loosening of plant cell walls by expansins. Nature 407: 321–326.
The diploid (2n) cell that results from the fusion of male and female gametes.Cosgrove, D. J., Li, L. C., Cho, H. T.et al. 2002. The growing world of expansins. Plant Cell Physiol. 43: 1436–1444.
The diploid (2n) cell that results from the fusion of male and female gametes.Côté, W. A. 1967. Wood Ultrastructure: An Atlas of Electron Micrographs. Seattle, WA: University of Washington Press.
The diploid (2n) cell that results from the fusion of male and female gametes.Dixon, H. H. and Joly, J.. 1894. On the ascent of sap. Ann. Bot. 8: 468–470.
The diploid (2n) cell that results from the fusion of male and female gametes.Ehlers, K. and Kollmann, R.. 2001. Primary and secondary plasmodesmata: structure, origin, and functioning. Protoplasma 216: 1–30.
The diploid (2n) cell that results from the fusion of male and female gametes.Esau, K. 1977. Anatomy of Seed Plants, 2nd edn. New York, NY: John Wiley and Sons.
The diploid (2n) cell that results from the fusion of male and female gametes.Fahn, A. 1990. Plant Anatomy, 4th edn. Oxford: Pergamon Press.
The diploid (2n) cell that results from the fusion of male and female gametes.Frost, F. H. 1930a. Specialization in secondary xylem in dicotyledons. I. Origin of vessel. Bot. Gaz. 89: 67–94.
The diploid (2n) cell that results from the fusion of male and female gametes.Frost, F. H. 1930b. Specialization in secondary xylem in dicotyledons. II. Evolution of end wall of vessel segment. Bot. Gaz. 90: 198–212.
The diploid (2n) cell that results from the fusion of male and female gametes.Frost, F. H. 1931. Specialization in secondary xylem in dicotyledons III. Specialization of lateral wall of vessel segment. Bot. Gaz. 91: 88–96.
The diploid (2n) cell that results from the fusion of male and female gametes.Fukuda, H. 1996. Xylogenesis: initiation, progression, and cell death. Annu. Rev. Plant Physiol. Plant Mol. Biol. 47: 299–325.
The diploid (2n) cell that results from the fusion of male and female gametes.Fukuda, H. 1997. Tracheary element differentiation. Plant Cell 9: 1147–1156.
The diploid (2n) cell that results from the fusion of male and female gametes.Fukuda, H. 2000. Programmed cell death of tracheary elements as a paradigm in plants. Plant Mol. Biol. 44: 245–253.
The diploid (2n) cell that results from the fusion of male and female gametes.Funada, R. 2002. Immunolocalization and visualization of the cytoskeleton in gymnosperms using confocal laser scanning microscopy. In Chaffey, N. G., ed., Wood Formation in Trees: Cell and Molecular Biology Techniques. London: Taylor and Francis, pp. 143–257.
The diploid (2n) cell that results from the fusion of male and female gametes.Funada, R., Furusawa, O., Shibagaki, M.et al. 2000. The role of cytoskeleton in secondary xylem differentiation in conifers. In Savidge, R. A., Barnett, J. R., and Napier, R., eds., Cell and Molecular Biology of Wood Formation. Oxford: Bios Scientific Publishers, pp. 255–264.
The diploid (2n) cell that results from the fusion of male and female gametes.Giddings, T. H.. and Staehelin, L. A.. 1988. Microtubule-mediated control of microfibril deposition: a re-examination of the hypothesis. In Lloyd, C. W., ed., The Cytoskeletal Basis of Plant Growth and Form. London: Academic Press, pp. 85–99.
The diploid (2n) cell that results from the fusion of male and female gametes.Gray-Mitsumune, M., Mellerowicz, E. J., Abe, H.et al. 2004. Expansins abundant in secondary xylem belong to subgroup A of the α-expansin gene family. Plant Physiol. 135: 1552–1564.
The diploid (2n) cell that results from the fusion of male and female gametes.Groover, A., DeWitt, N., Heidel, A., and Jones, A.. 1997. Programmed cell death of plant tracheary elements differentiating in vitro. Protoplasma 196: 197–211.
The diploid (2n) cell that results from the fusion of male and female gametes.Hejnowicz, Z. 1980. Tensional stress in the cambium and its developmental significance. Am. J. Bot. 67: 1–5.
The diploid (2n) cell that results from the fusion of male and female gametes.Hepler, P. K. and Newcomb, E. H.. 1964. Microtubules and fibrils in the cytoplasm of Coleus cells undergoing secondary wall deposition. J. Cell Biol. 20: 529–533.
The diploid (2n) cell that results from the fusion of male and female gametes.Hillis, W. E. 1987. Heartwood and Tree Exudates. Berlin: Springer-Verlag.
The diploid (2n) cell that results from the fusion of male and female gametes.Hirakawa, Y. 1984. A SEM observation of microtubules in xylem cells forming secondary walls of trees. Res. Bull. Coll. Exp. Forestry, Hokkaido Univ. 41: 535–550.
The diploid (2n) cell that results from the fusion of male and female gametes.Hogetsu, T. 1991. Mechanism for formation of the secondary wall thickening in tracheary elements: microtubules and microfibrils of tracheary elements of Pisum sativum L. and Commelina communis L. and the effects of amiprophosmethyl. Planta 185: 190–200.
The diploid (2n) cell that results from the fusion of male and female gametes.Ito, J. and Fukuda, H.. 2002. ZENI is a key enzyme in the degradation of nuclear DNA during programmed cell death of tracheary elements. Plant Cell 14: 3201–3211.
The diploid (2n) cell that results from the fusion of male and female gametes.Juniper, B. E. 1977. Some speculations on the possible role of the plasmodesmata in the control of differentiation. J. Theor. Biol. 66: 583–592.
The diploid (2n) cell that results from the fusion of male and female gametes.Kribs, D. A. 1935. Salient lines of structural specialization in the wood rays of dicotyledons. Bot. Gaz. 96: 547–557.
The diploid (2n) cell that results from the fusion of male and female gametes.Kuriyama, H. and Fukuda, H.. 2002. Developmental programmed cell death in plants. Curr. Opin. Plant Biol. 5: 568–573.
The diploid (2n) cell that results from the fusion of male and female gametes.Larson, P. R. 1994. The Vascular Cambium: Development and Structure. Berlin: Springer-Verlag.
The diploid (2n) cell that results from the fusion of male and female gametes.Ledbetter, M. C. and Porter, K. R.. 1963. A “microtubule” in plant cell fine structure. J. Cell Biol. 19: 239–250.
The diploid (2n) cell that results from the fusion of male and female gametes.Lockhart, J. A. 1965. An analysis of irreversible plant cell elongation. J. Theor. Biol. 8: 264–275.
The diploid (2n) cell that results from the fusion of male and female gametes.Metcalfe, C. R. and Chalk, L.. 1950. Anatomy of the Dicotyledons, 2 vols. Oxford: Clarendon Press.
The diploid (2n) cell that results from the fusion of male and female gametes.Milburn, J. A. 1996. Sap ascent in vascular plants: challenges to the cohesion theory ignore the significance of immature xylem and the recycling of Münch water. Ann. Bot. 78: 399–407.
The diploid (2n) cell that results from the fusion of male and female gametes.Ohdaira, Y., Kakegawa, K., Amino, S., Sugiyama, M., and Fukuda, H.. 2002. Activity of cell-wall degradation associated with differentiation of isolated mesophyll cells of Zinnia elegans into tracheary elements. Planta 215: 177–184.
The diploid (2n) cell that results from the fusion of male and female gametes.Paredez, A. R., Somerville, C. R. and Ehrhardt, D. W.. 2006. Visualization of cellulose synthase demonstrates functional association with microtubules. Science 312: 1491–1495.
The diploid (2n) cell that results from the fusion of male and female gametes.Peters, W. J. 1974. Tylosis formation in Pinus tracheids. Bot. Gaz. 135: 126–131.
The diploid (2n) cell that results from the fusion of male and female gametes.Preston, R. D. 1974. The Physical Biology of Plant Cell Walls. London: Chapman and Hall.
The diploid (2n) cell that results from the fusion of male and female gametes.Ray, P. M., Green, P. B., and Cleland, R. E.. 1972. Role of turgor in plant cell growth. Nature 239: 163–164.
The diploid (2n) cell that results from the fusion of male and female gametes.Rioux, D., Chamberland, H., Simard, M., and Ouellette, G. B.. 1995. Suberized tyloses in trees: an ultrastructural and cytochemical study. Planta 196: 125–140.
The diploid (2n) cell that results from the fusion of male and female gametes.Roberts, L. W., Gahan, P. B., and Aloni, R.. 1988. Vascular Differentiation and Plant Growth Regulators. Berlin: Springer-Verlag.
The diploid (2n) cell that results from the fusion of male and female gametes.Romberger, J. A., Hejnowicz, Z., and Hill, J. F.. 1993. Plant Structure: Function and Development. Berlin: Springer-Verlag.
The diploid (2n) cell that results from the fusion of male and female gametes.Sachs, T. 1981. The control of patterned differentiation of vascular tissues. Adv. Bot. Res. 9: 151–262.
The diploid (2n) cell that results from the fusion of male and female gametes.Schrader, J., Baba, K., May, S. T.et al. 2003. Polar auxin transport in the wood-forming tissues of hybrid aspen is under simultaneous control of developmental and environmental signals. Proc. Natl. Acad. Sci. USA 100: 10096–10101.
The diploid (2n) cell that results from the fusion of male and female gametes.Stiller, V. and Sperry, J. S.. 1999. Canny's compensating pressure theory fails a test. Am. J. Bot. 86: 1082–1086.
The diploid (2n) cell that results from the fusion of male and female gametes.Sugimoto-Shirasu, K., Carpita, N. C. and McCann, M. C.. 2004. The cell wall: a sensory panel for signal transduction. In Hussey, P. J., ed., The Plant Cytoskeleton in Cell Differentiation and Development. Oxford: CRC Press, pp. 176–203.
The diploid (2n) cell that results from the fusion of male and female gametes.Turner, S., Gallois, P. and Brown, D.. 2007. Tracheary element differentiation. Annu. Rev. Plant Biol. 58: 407–433.
The diploid (2n) cell that results from the fusion of male and female gametes.Uehara, K. and Hogetsu, T.. 1993. Arrangement of cortical microtubules during formation of bordered pit in the tracheids of Taxus. Protoplasma 172: 145–153.
The diploid (2n) cell that results from the fusion of male and female gametes.Uggla, C., Mellerowicz, E. J. and Sundberg, B.. 1998. Indole-3 acetic acid controls cambial growth in Scots pine by positional signaling. Plant Physiol. 117: 113--121
The diploid (2n) cell that results from the fusion of male and female gametes.Bel, A. J. E. 1990. Xylem–phloem exchange via rays: the undervalued route of transport. J. Exp. Bot. 41: 631–644.
The diploid (2n) cell that results from the fusion of male and female gametes.Bel, A. J. E. and Ehlers, K.. 2000. Symplasmic organization of the transport phloem and the implications for photosynthate transfer to the cambium. In Savidge, R. A., Barnett, J. R., and Napier, R., eds., Cell and Molecular Biology of Wood Formation. Oxford: Bios Scientific Publishers, pp. 85–99.
The diploid (2n) cell that results from the fusion of male and female gametes.Werker, E. and Fahn, A.. 1969. Resin ducts of Pinus halepensis Mill.: their structure, development and pattern of arrangement. Bot. J. Linn. Soc. 62: 379–411.
The diploid (2n) cell that results from the fusion of male and female gametes.Wooding, F. B. P. and Northcote, D. H.. 1964. The development of the secondary wall of the xylem in Acer pseudoplatanus. J. Cell Biol. 23: 327–337.
The diploid (2n) cell that results from the fusion of male and female gametes.Zimmermann, U., Meinzer, F., and Bentrup, F.-W.. 1995. How does water ascend in tall trees and other vascular plants? Ann. Bot. 76: 545–551.
FURTHER READING
The diploid (2n) cell that results from the fusion of male and female gametes.Botosso, P. C. and Gomes, A. V.. 1982. Radial vessels and series of perforated ray cells in Annonaceae. Int. Ass. Wood Anatomists Bull. 3: 39–44.
The diploid (2n) cell that results from the fusion of male and female gametes.Braun, H. J. 1970. Handbuch der Pflanzenanatomie, vol. 9, part 1, Funktionelle Histologie der sekundären Sprossachse. I. Das Holz. Berlin: Bornträger.
The diploid (2n) cell that results from the fusion of male and female gametes.Carlquist, S. 1961. Comparative Plant Anatomy. New York, NY: Holt, Rinehart and Winston.
The diploid (2n) cell that results from the fusion of male and female gametes.Carlquist, S. 1975. Ecological Strategies of Xylem Evolution. Berkeley, CA: University of California Press.
The diploid (2n) cell that results from the fusion of male and female gametes.Chaffey, N. (ed.) 2002. Wood Formation in Trees: Cell and Molecular Biology Techniques. London: Taylor and Francis.
The diploid (2n) cell that results from the fusion of male and female gametes.Chattaway, M. M. 1949. The development of tyloses and secretion of gum in heartwood formation. Austral. J. Sci. Res. B, Biol. Sci. 2: 227–240.
The diploid (2n) cell that results from the fusion of male and female gametes.Esau, K. 1965a. Plant Anatomy, 2nd edn. New York, NY: John Wiley and Sons.
The diploid (2n) cell that results from the fusion of male and female gametes.Esau, K. 1965b. Vascular Differentiation in Plants. New York, NY: Holt, Rinehart and Winston.
The diploid (2n) cell that results from the fusion of male and female gametes.Funada, R., Abe, H., Furusawa, O.et al. 1997. The orientation and localization of cortical microtubules in differentiating conifer fracheids during cell expansion. Plant Cell Physiol. 38: 210–212.
The diploid (2n) cell that results from the fusion of male and female gametes.Greenidge, K. N. H. 1952. An approach to the study of vessel length in hardwood species. Am. J. Bot. 39: 570–574.
The diploid (2n) cell that results from the fusion of male and female gametes.Gregory, S. G. and Petty, J. A.. 1973. Valve action of bordered pits in conifers. J. Exp. Bot. 24: 763–767.
The diploid (2n) cell that results from the fusion of male and female gametes.Gregory, R. A. and Romberger, J. A.. 1975. Cambial activity and height of uniseriate vascular rays in conifers. Bot. Gaz. 136: 246–253.
The diploid (2n) cell that results from the fusion of male and female gametes.Jane, F. W. 1970. The Structure of Wood, 2nd edn. London: A. and C. Black.
The diploid (2n) cell that results from the fusion of male and female gametes.Meylan, B. A. and Butterfield, B. G.. 1972. Perforation plate development in Knightia excelsa R. Br.: a scanning electron microscope study. Austral. J. Bot. 20: 79–86.
The diploid (2n) cell that results from the fusion of male and female gametes.O'Brien, T. P. 1970. Further observations on hydrolysis of the cell wall in the xylem. Protoplasma 69: 1–14.
The diploid (2n) cell that results from the fusion of male and female gametes.Panshin, A. J. and Zeeuw, C.. 1970. Textbook of Wood Technology, vol. 1. New York, NY: McGraw-Hill.
The diploid (2n) cell that results from the fusion of male and female gametes.Picket-Heaps, J. D. 1968. Xylem wall deposition: radioautographic investigations using lignin precursors. Protoplasma 65: 181–205.
The diploid (2n) cell that results from the fusion of male and female gametes.Raven, J. A. 1977. The evolution of vascular land plants in relation to supracellular transport processes. Adv. Bot. Res. 5: 153–219.
The diploid (2n) cell that results from the fusion of male and female gametes.Robards, A. W. (ed.) 1974. Dynamic Aspects of Plant Ultrastructure. London: McGraw-Hill.
The diploid (2n) cell that results from the fusion of male and female gametes.Tyree, M. T. and Ewers, F. W.. 1991. The hydraulic architecture of trees and other woody plants. New Phytol. 119: 345–360.
The diploid (2n) cell that results from the fusion of male and female gametes.Tyree, M. T. and Zimmermann, M. H.. 2002. Xylem Structure and the Ascent of Sap. Berlin: Springer-Verlag.
The diploid (2n) cell that results from the fusion of male and female gametes.Wenham, M. W. and Cusik, F.. 1975. The growth of secondary wood fibres. New Phytol. 74: 247–261.
The diploid (2n) cell that results from the fusion of male and female gametes.Zimmermann, M. H. and Brown, C. L.. 1971. Trees: Structure and Function. New York, NY: Springer-Verlag.