Xylem Structure and Function

Vascular plants have evolved a highly specialized tissue, called xylem, which provides mechanical support and transports water, mineral nutrients and phytohormonal signals in the plant. Although it is the most abundant biological tissue on earth, much remains to be learned about the structure, function, development and evolution of xylem and of the genes that regulate the processes.

Keywords: evolution; water transport; structural support; differentiation; xylogenesis

Figure 1. Drawing of the secondary thickened wall of a mature tracheary element showing the orientation of cellulose microfibrils in the different layers of the wall. Note the designation of the secondary wall layers and the average microfibril angle of each layer: S1 is the outermost layer, S2 is the middle layer and S3 the innermost layer. Most of the wall thickness is determined by the thickness of the S2 layer (the relative thicknesses are: primary wall, 1%; S1,10 to 20%; S2, 40 to 90% and S3, 2 to 8%). Modified after Côté WA (1967) Wood Ultrastructure: An Atlas of Electron Micrographs. Seattle: University of Washington Press.
Figure 2. Drawing showing the relative sizes and shapes of some xylem cell types: (a) conifer tracheid with circular bordered pits, (b) fibre tracheid with bordered pits, (c) libriform fibre with simple pitting, (d) vessel element with scalariform perforations and (e) vessel element with a simple perforation. Note that conifer tracheids (3 to 5 mm) are usually much longer in relationship to fibres (0.8 to 2.3 mm) and vessel elements (0.2 to 1.3 mm).
Figure 3. Structure of a bordered pit in the secondary wall of a conifer tracheid showing the modification of the pit membrane to a torus and margo. Note the loose network of cellulose fibrils that forms the margo and the secondary thickening of the central region to form the torus. In angiosperms, the pit membranes of bordered pits are usually not modified.
Figure 4. Drawing of cross sections of young woody stems showing the cambial zone and secondary xylem development. (a) Dicot wood. (b) Conifer wood. Note the abrupt change in the size of tracheids from earlywood to latewood. M, differentiating xylem and phloem mother cells; C, cambial initial.
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 Further Reading
    Boudet AM, Lapierre C and Grima-Pettenati J (1995) Tansley Review no 80: Biochemistry and molecular biology of lignification. New Phytologist 129: 203–236.
    book Carlquist JS (1975) Ecological Strategies of Xylem Evolution. Berkeley, CA: University of California Press.
    Delmer DP and Amor Y (1995) Cellulose biosynthesis. Plant Cell 7: 987–1000.
    book Fahn A (1990) Plant Anatomy, 4th ed. New York: Pergamon Press.
    Fukuda H (1996) Xylogenesis: initiation, progression and cell death. Annual Review of Plant Physiology and Molecular Biology 47: 299–325.
    book Higuchi T (1997) Biochemistry and Molecular Biology of Wood. Berlin: Springer-Verlag.
    book Ingrouille M (1992) Diversity and Evolution of Land Plants. London: Chapman & Hall.
    book Mauseth JD (1988) Plant Anatomy. Menlo Park, CA: Benjamin/Cummings.
    Whetten RW, MacKay JJ and Sederoff RR (1998) Recent advances in understanding lignin biosynthesis. Annual Review of Plant Physiology and Plant Molecular Biology 49: 585–609.
    book Zimmermann MH (1983) Xylem Structure and the Ascent of Sap. Berlin: Springer-Verlag.
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Myburg, Alexander A, and Sederoff, Ronald R(Apr 2001) Xylem Structure and Function. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1038/npg.els.0001302]