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Xylem Structure and Function: Alexander A Myburg, Ronald R Sederoff
Xylem Structure and Function: Alexander A Myburg, Ronald R Sederoff
Function . Introduction
Article Contents
Alexander A Myburg, North Carolina State University, Raleigh, North Carolina, USA . Xylem Structure and Variability
Ronald R Sederoff, North Carolina State University, Raleigh, North Carolina, USA . Xylem Functions: Water Transport and Structural
Support
. Xylem Differentiation and Cell Wall Biosynthesis
Vascular plants have evolved a highly specialized tissue, called xylem, which provides . Origin and Evolution of Xylem in Plants
mechanical support and transports water, mineral nutrients and phytohormonal signals in . Genetic Manipulation of Xylem Formation
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.
Perforation plate
Bordered Simple pit
pits Perforation
(e)
(d)
(c)
(b)
(a)
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).
Border
Secondary growth
The secondary xylem of woody plants constitutes the
major part of the stem, i.e. the wood. This section will focus
Inner aperture
on various aspects of wood structure that are directly
Torus related to the development and organization of secondary
Margo
xylem.
co-ordinated with cell divisions in the adjacent interfasci- these circles are true annual rings and the age of the stem
cular region to produce a continuous cylinder of vascular can be deduced from the number of rings. In many regions
cambium. Usually, secondary xylem is formed on the inner of the world, particularly the tropics, growth rings do not
side and secondary phloem on the outer side of the always represent annual increments. More than one (or less
cambium (Figure 4). than one) growth ring can be formed per year, for example
when several dry and wet periods occur within a year.
Fusiform and ray initials give rise to the axial and radial
components of xylem
Heartwood and sapwood
The vascular cambium consists of fusiform and ray initials.
Wood cells have a limited lifetime in which they can
Fusiform initials divide longitudinally to give rise to the
actively transport water. After a variable number of years,
axial components of secondary growth, i.e. tracheary
cavitation occurs in most of the vessels and tracheids and
elements, bres and axial parenchyma towards the inside
the rest of the xylem cells in the growth ring die. These cells
and phloem cells towards the outside of the stem. Ray
are then lled with resinous materials and polyphenols,
initials divide to form ray cells that run radially across the
and constitute the inner, often darker part of the woody
secondary vascular tissue. Rays serve to transport water,
stem called heartwood. The outer, water-conducting part
dissolved gases and organic nutrients radially in wood. As
of the stem is called sapwood. In many species, as sapwood
secondary growth proceeds, the cambial cylinder increases
is converted to heartwood, air-lled vessels in the sapwood
in diameter through lateral division of fusiform initials.
are often sealed o by the intrusive growth of surrounding
parenchyma cells. These intrusions are called tyloses and,
Earlywood, latewood and growth rings together with the resinous materials, serve to prevent
The cambium of many woody plants exhibits periodic fungal growth in the empty vessel lumens. The outer,
activity. In the spring and early summer (in temperate conducting part of the stem is called sapwood.
regions), conditions are conducive to active growth and
relatively wide tracheary elements with thin walls are
produced. Later in the summer and autumn, relatively Dicot versus conifer wood
narrow tracheary elements with thick walls are formed Woods are commonly classied as either hardwoods or
(Figure 4). These two types of xylem, called earlywood and softwoods. Hardwoods are angiosperm (dicotyledonous)
latewood, are most commonly observed as concentric trees, while softwoods are gymnosperm (conifer) trees.
circles on the transverse section of the stem and are formed These two terms do not accurately express dierences in
as a result of changes in the activity of the vascular the hardness or density of the wood, but are useful for the
cambium. When the activity of the vascular cambium is description of the basic structural dierences between dicot
controlled by annual seasons (one ring is formed per year), and conifer wood.
Latewood vessel
Earlywood vessel
Ray
(a)
Earlywood Latewood M M C M
Pith
Primary xylem
Secondary xylem
Cambial
(b) Phloem
zone
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.
Dicot wood contains vessels, fibres, parenchyma and Secondary thickening in monocots
tracheids The majority of monocots are herbaceous, which means
Dicot wood contains a greater number of cell types than that the primary xylem has to full all the requirements of
conifer wood and the structure of dicot wood, therefore, water transport that the plant may encounter throughout
varies more than that of conifer wood. The cell types of its lifetime. However, some monocots do undergo thicken-
dicot wood include vessel elements, bres and parenchyma ing of the primary stem. In bamboos and other monocot
cells. Tracheids are rare in dicots, but occur in some species species with wide stems, a broad region of mitotic activity,
such as oaks and chestnuts. The cell types in dicot wood are called the primary thickening meristem, is responsible for
also more diversied in function; vessels and tracheids radial and tangential expansion of the primary stem. Very
transport water, bres provide structural support and few examples exist of truly woody monocots. In woody
parenchyma cells perform storage and regeneration func- monocot genera such as Yucca and Dracaena, the activity
tions. The feature that most distinguishes dicot wood from of a secondary thickening meristem in the outer cortex of
conifer wood is the presence of large-diameter vessel the stem is responsible for anomalous secondary growth.
elements that disrupt the regular organization of the radial Arborescent monocots such as palms undergo diuse
cell les derived from the cambial initials (Figure 4a). Two secondary growth through the division of cells in the
types of bres are common in dicot wood: bre tracheids ground parenchyma of the stem.
and libriform bres (Figure 2b and c). Fibre tracheids have
thick walls with bordered pits. Libriform bres have simple
pits. Dicot wood generally contains larger rays than conifer Xylem Functions: Water Transport and
wood and in most dicot species the rays consist only of ray
parenchyma. Structural Support
Water transport
Conifer wood consists mostly of regular files of tracheids
Conifer wood is relatively simple in structure. The most A gradient of water potential drives water transport
distinctive features of conifer wood include: the regular Despite a large amount of research on this topic, the precise
organization of the radial les of tracheids, the absence of mechanism of water transport in plants is still debated. The
vessels and bres and the small amount of wood experimental evidence strongly suggests that water trans-
parenchyma (Figure 4b). The long, tapered tracheids form port in plants is driven by a gradient of water potential that
the predominant cell type and full both the mechanical exists between the air surrounding the leaves at one end and
and conductive functions of conifer wood. The majority of the water that surrounds the roots at the other. These two
parenchyma cells in conifer wood are present in rays and, in extremes are connected by the xylem, which supports a
some conifers such as the Pinaceae, in axial and radial resin water column that extends from the roots to the leaves. Air
ducts. Conifer rays consist primarily of ray parenchyma usually has a very negative water potential (even when the
and, in some conifers, a smaller amount of ray tracheids. humidity is very high). As the leaves of the plant lose water
Resin ducts are large intercellular spaces surrounded by to the air, the water potential becomes more negative inside
thin-walled parenchyma cells that excrete resin into the leaf cells. This causes water to gradually move from xylem
duct. The resin is believed to seal wounds and protect the cells to leaf cells. The water molecules inside the water
plant against fungi and herbivores. columns of the capillary xylem elements are pulled
upwards by cohesion forces when water molecules at the
top of the columns move out into the aerial parts of the
Reaction wood plants. This is known as the cohesiontension theory of sap
Woody plants respond to bending induced by external ascent.
forces, such as wind and gravity, by making reaction wood.
Conifers produce reaction wood on the side of the branch Adhesion, cohesion and tension forces act on the water
or stem where the tissues are compressed (usually the column
underside) and it is therefore called compression wood. In The upward movement of the water column is counter-
dicots, reaction wood forms on the side under tension acted by three forces: (1) the weight of the water column,
(usually the upper side) and it is called tension wood. (2) adhesion of water to the cell walls of tracheary elements
Compression wood has thicker cell walls, higher lignin and (3) adhesion of the water to soil particles. The upward
content and is darker than normal conifer wood. Tension movement of the water molecules in each tracheary
wood is characterized by the presence of gelatinous bres, element will cause tension in the water column, causing it
low lignin content and high cellulose content. The purpose to become narrower. During times of high transpiration,
of reaction wood is to reorient bent stems and branches to the negative pressure inside tracheary elements can become
allow optimal light exposure of the tree canopy. strong enough to cause these cells to collapse inward.
in primary xylem and from 1421 days in secondary xylem. delivered to the cell wall via Golgi and endoplasmic
Xylogenesis typically includes the following phases: (1) cell reticulum-derived vesicles and polymerized into lignin by
division and enlargement, (2) cell wall thickening, (3) wall-bound enzymes. The aromatic nature of the lignin
lignication and (4) programmed cell death. Cell wall monomers makes lignin hydrophobic. Lignin, therefore,
biosynthesis is an integral part of xylem formation. The provides a hydrophobic inner surface to the cell wall that
basic chemical components and organization of xylem cell facilitates water transport. The three-dimensional nature
walls are known, but little is known of the mechanisms by of the lignin polymer provides rigidity and compressive
which they are synthesized and organized to form the strength to the cell wall, while the chemical stability of
highly complex cell wall. This section will outline the lignin provides protection against pathogens.
phases of xylem dierentiation and the formation of xylem
cell walls, which form the major part of mature xylem cells. Tracheary elements undergo programmed
cell death
Xylem cells are derived from apical and lateral
meristems At the completion of secondary wall deposition and
lignication, tracheary elements undergo autolysis, an
Tracheary elements are derived from either the procam- example of programmed cell death in higher plants. Soon
bium (primary xylem) or vascular cambium (secondary after the initiation of secondary thickening, hydrolytic
xylem). The dierentiation of cambial initials into xylem enzymes (DNAases, RNAases and proteases) start accu-
elements is thought to be initiated by plant hormones. The mulating in the vacuole. The autolytic process is initiated
immature xylem cells have dense protoplasm, small when the tonoplast ruptures, causing the hydrolytic
vacuoles and thin primary walls (Figure 4). Soon after cell enzymes to spill out into the cytoplasm. This leads to the
division, these cells undergo cell elongation and an increase complete degradation of the cell contents and partial
in the size of the vacuole and nuclei. digestion of the unprotected regions of the primary wall.
Only regions covered by lignied secondary wall material
Most xylem cell walls undergo secondary are protected from degradation. The end walls of
dierentiating vessel elements are degraded at the perfora-
thickening tion sites to allow direct cell-to-cell movement of water and
The deposition of secondary walls begins sometime before nutrients. Only regions covered by lignied secondary wall
tracheary elements and bres reach their full size. The material are protected from degradation. Pit membranes
cellulose, lignin, hemicellulose and protein components of are often partially degraded to leave mats of cellulose brils
the secondary wall are synthesized and deposited coopera- (Figure 3). This enhances the movement of water through
tively during secondary wall thickening. The onset of pit-pairs, which is the only way water can enter and leave
secondary wall thickening is associated with the formation tracheids.
of arrays of microtubules under those regions of the plasma
membrane where active secondary wall deposition will
take place. Microtubules may play a role in dening the
pattern of secondary walls by guiding dictyosome-derived Origin and Evolution of Xylem in Plants
vesicles with cell wall material to the sites of deposition on
the cell membrane. Cellulose microbrils are produced at Vascular plants (Tracheophyta) are characterized by the
the membrane surface of the cell by complex rosette presence of xylem tissue with lignied cell walls. Modern
structures, which consist of several dierent proteins. The vascular plants are ferns, gymnosperms and angiosperms.
movement of these rosette complexes in the plasma Mosses, liverworts and hornworts (Bryophyta) do not
membrane may also be directed by microtubules. contain xylem. Tracheid-like cells, called hydroids, are
present in certain bryophytes, but lignied cell wall
thickenings are absent in these plants. This section will
The cell walls and intercellular regions of outline the major trends of xylem evolution in vascular
xylem cells are lignified plants.
Following secondary thickening of the xylem cell walls,
lignin is deposited between the newly formed tracheary Evolution of primary xylem
elements and within their walls. The area between the cells,
called the middle lamella, and the primary walls are rapidly Tracheids were present in the first vascular land plants
lignied, followed by a more gradual lignication of the It is widely accepted that the rst land plants evolved from
secondary walls. Lignin is a very complex, crosslinked, green algae (Chlorophyta) and that these plants were
three-dimensional polymer of aromatic phenolic mono- adapted to aquatic or semiaquatic environments. The
mers, called cinnamyl alcohols. The lignin monomers are evolution of conducting tissue was closely associated with
Secondary xylem increased the lifespan of plants New technologies allow rapid progress in the
The ability to produce secondary xylem had profound genetic manipulation of xylem
consequences for early vascular plants. It greatly increased
the lifespan of plants by allowing plants to essentially form Studies of model plant systems, such as Arabidopsis,
a new water-conducting system each year that replaced the Zinnia, tobacco and maize have been important in
non-functional xylem elements from previous years. The identifying specic genes and proteins involved in cell wall
increase in lifespan enabled the existence of taller plants formation. Genetic and biochemical studies of cotton and
and increased the need for long-distance conductance and forest trees have identied some important genes for the
mechanical support. The major trends of xylem evolution formation of cellulose and lignin. Most recently, many
(the shift towards vessel elements, simple perforations and laboratories have decided to use high-throughput auto-
libriform bres) are thought to be associated with the mated techniques to identify all of the expressed genes of
increased eciency of water transport in xylem and, to a higher plants and to learn their function. This approach,
lesser degree, the increased demand for mechanical called genomics, is expected to rapidly advance the
support in plants. knowledge of the genes and proteins forming the primary
and secondary cell walls of xylem. With this knowledge, the Fukuda H (1996) Xylogenesis: initiation, progression and cell death.
modication of xylem in commercially important plants Annual Review of Plant Physiology and Molecular Biology 47: 299325.
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Phytologist 129: 203236. understanding lignin biosynthesis. Annual Review of Plant Physiology
Carlquist JS (1975) Ecological Strategies of Xylem Evolution. Berkeley, and Plant Molecular Biology 49: 585609.
CA: University of California Press. Zimmermann MH (1983) Xylem Structure and the Ascent of Sap. Berlin:
Delmer DP and Amor Y (1995) Cellulose biosynthesis. Plant Cell 7: 987 Springer-Verlag.
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Fahn A (1990) Plant Anatomy, 4th ed. New York: Pergamon Press.