LECTURE NOTE

COURSE CODE (TITLE): PSB 422 (ADVANCED PLANT ANATOMY)

LECTURER: F. I. NWAFOR

LESSON 1: THE ULTRASTRUCTURE OF FIBRE CELL WALL

1. INTRODUCTION Last updated: November 07, 2022

Wood is a complex composite material that fulfils a range of functions. The roles of wood, in a
living tree, are to support the aerial part, transport water and store carbohydrates. The bark is
external to the wood. Beneath the bark lies the vascular cambium, which is composed of a few
layers of living cells that actively divide and produce more wood, which results in an increase in
the girth of the tree. Wood may be sub-divided into sapwood and heartwood. Sapwood conducts
water and inorganic nutrients (sap), while heartwood is old sapwood that and has been chemically.

Wood is a complex composite material that fulfils a range of functions. The roles of wood, in a
living tree, are to support the aerial part, transport water and store carbohydrates. The wood of
softwood species is composed predominantly of tracheids. These cells are mainly longitudinal, or
axial—their long axis runs parallel to the axis of the trunk (vertical in the standing tree). Axial
parenchyma is present in certain softwood species, but radial parenchyma is always present
and constitutes the rays, sometimes together with radial tracheids. In hardwoods the proportion
of constituent cell types—vessel members, fibres, and parenchyma—depends mainly on species.
Vessel members and fibres are always present and axially oriented; axial parenchyma is seldom
absent. Rays in hardwoods are made entirely of radial parenchyma cells.

All the above cells are tubelike. Tracheids and fibres have closed ends. Vessel members have ends
wholly or partly open; in wood tissue, vessel members are connected end to end to form vertical
pipelike stacks (vessels) of indeterminate length. The characteristic pores visible in the transverse
section of hardwoods are actually vessel members. Axial tracheids in softwood species and vessel
members in hardwood species are the principal water-conducting cells. Although fibres in
hardwood trees may also participate in conduction, their main function is to provide mechanical
support.

A living plant cell consists of two primary domains: the protoplast and the cell wall. The protoplast
is the sum of the living contents that are bounded by the cell membrane. The cell wall is a non-
living, largely carbohydrate matrix extruded by the protoplast to the exterior of the cell membrane.
The plant cell wall protects the protoplast from osmotic lysis and often provides mechanical
support to the plant at large. For cells in wood, the situation is somewhat more complicated than
this highly generalized case. In many cases in wood, the ultimate function of the cell is borne solely
by the cell wall. This means that many mature wood cells not only do not require their protoplasts,
but indeed must completely remove their protoplasts prior to achieving functional maturity. For
this reason, a common convention in wood literature is to refer to a cell wall without a protoplast
as a cell. Although this is technically incorrect from a cell biological standpoint, this convention
is common in the literature. In the case of a mature cell in wood in which there is no protoplast,
the open portion of the cell where the protoplast would have existed is known as the lumen (plural:
lumina). Thus, in most cells in wood there are two domains; the cell wall and the lumen. The lumen
is a critical component of many cells, whether in the context of the amount of space available for
water conduction or in the context of a ratio between the width of the lumen and the thickness of
the cell wall. The lumen has no structure per se, as it is the void space in the interior of the cell.
Thus, wood is a substance that has two basic domains; air space (mostly in the lumina of the cells)
and the cell walls of the component cells.

2. FIBRE

The term fibre is used by many researchers using natural fibres in composites. However, ‘fibre’
(or fiber) has a strict botanical meaning, which describes a single elongated, thick-walled, plant
cell. The non-botanical definition also applies to the pulp and paper field, and the term ‘shive’
refers to a bundle of fibres. In non-woody plants (e.g. hemp/flax), ‘fibre’ may be used to refer to a
bundle of non-individualised cells adhering to each other in the same way as they were in the plant.
These are also generally known as bast/sisal fibres. In this context, ‘shive’ refers to a bundle
contaminated with non-fibrous plant material. For convenience however, the term ‘fibre’ in this
lecture note means a single elongated cell. Fibres that remain attached to each other in a long
fibrous strand are referred to as fibre bundles.

Various types of fibre. (a) Softwood shive, r = ray; (b) softwood fibre (tracheid); (c) bast
fibre; (d) bast fibre bundle; (e) hardwood fibre.

3. ORIGIN OF FIBRE

Primary plant growth produces leaves and non-woody material with the function of providing
mechanical support and form to the plant, while secondary plant growth mainly produces woody
material used in both sap conductance and mechanical support. Both types of growth can be used
to produce fibres. Primary plant growth produces bast fibres in the phloem (the ‘inner bark’,
sometimes called ‘skin’ or ‘bast’) surrounding the stem of certain dicotyledonous plants, such as
hemp and flax. Secondary plant growth increases the girth of a stem. Wood is a product of
secondary growth, as is bamboo. A comparatively small amount of bast fibre can also be produced
by secondary growth.
4. FIBRE SIZE

Softwood fibres (botanically called tracheids) can have lengths up to approximately 3.5 mm,
diameters of approximately 30 µm and wall thicknesses in the range of 2–3 µm. Hardwood fibres
are approximately 0.7–1.5 mm long, with diameters in the range of 13–20 µm and wall thicknesses
of 3–6 µm. Literature values for the dimensions of bast fibres vary widely. For example, the fibres
of kenaf tend to be in the range of 2–3 mm long (similar to wood), while flax and hemp fibres can
be much longer; in the range of 2–5 cm. Bast fibre diameters vary between approximately 8 and
25 µm (calculated from fibre areas), and wall thicknesses vary between 4 and 12 µm. Some of the
reasons for these differences among bast fibre dimensions relate to their ontogeny.

5. THE ULTRASTRUCTURE OF FIBRE CELL WALL

The mechanical properties of the fibres originate in the architecture of the cell wall. The cell wall
of land plants is characterized by a framework of polysaccharides, specifically semi-crystalline
cellulose microfibrils embedded in an amorphous matrix containing hemicellulose, pectin,
polyphenolic material (lignin) and proteins. The cell wall properties are controlled on many levels,
such as by the transcriptional regulation of genes, membrane trafficking and molecular interaction
between different components of the cell wall.

Observed microscopically, the cells of wood appear to be composed of cell wall and cell cavity;
in dead cells the cavity is empty. Gaps of various shapes, called pits, are often seen in great
numbers in the cell walls. Pits serve as passages of communication between neighbouring cells
and come in pairs—one in each of the adjoining cell walls—separated by a membrane. Other
microscopic features are tyloses, plugs comprising various plant materials that obstruct the vessel
members of hardwoods and that form mainly when sapwood is transformed to heartwood. Under
the microscope, the resin canals of softwoods are revealed to be not cells but tubular spaces
between cells, lined with specialized parenchyma; they also are plugged in heartwood.

Polarization microscopy, X rays, electron microscopy, and other techniques provide information
regarding the structure of cell walls and other features hidden to light microscopes. The wall is
organised with three layers: middle lamella, primary cell wall and secondary cell wall. During cell
division (cytokinesis), one cell is divided into two by the formation of the cell plate. This structure
will later become the middle lamella. It is a thin layer highly enriched in another type of
polysaccharide (pectins). Then, a primary wall is deposited on each side, as the cells expand
towards their final dimensions. The primary cell wall is a heavily hydrated gel-like composite
comprised mostly of cellulose, hemicelluloses, pectins, structural glycoproteins, some proteins and
small quantities of phenolic acids. It contains 3–4 layers of cellulose microfibrils with a dispersed
orientation. Nevertheless, some degree of microfibril alignment can be observed in tissues where
cell elongation is taking place. Middle lamellae and primary cell walls are two hydrophilic
contiguous structures ubiquitous in all land plants. Later, after the lignification process, they will
become indiscernible and might be grouped under the term compound middle lamella (CML).
Once cell enlargement has finished, the secondary cell walls are formed. The deposition of new
material results in the walls becoming thicker. This addition occurs on the inside of the primary
cell wall, so this process reduces the internal cell diameter (lumen). Secondary cell walls mainly
contain cellulose, hemicellulose and lignin in varying proportions.

From tree to the molecular structure of wood fibre cell wall

The secondary cell wall is usually organized into three layers. Adjacent to the compound middle
lamellae is an outer layer called the S1, which has transversely-oriented microfibrils with a high
angle. The middle, S2, layer makes up the greatest proportion of the wall thickness and has the
lowest microfibril angle orientation (angle between microfibril and fibre axis). The inner, S3, layer
also has transversely-oriented microfibrils. The microfibril angle varies depending on the species,
maturity, position in the cell wall and growth rate; it dictates some of the characteristics of the
plant, such as stiffness. The mechanisms that control changes in cellulose microfibril orientation
among wall layers are not known. During the last stage of differentiation, when the lignin is
deposited, an additional warty layer that consists of spherical particles with a diameter of 0.1–0.3
µm is created, next to the lumen, on the innermost side. The mechanical properties of plant fibres
depend on the architecture of the secondary cell walls; an understanding of the cell wall
architecture is important for not only an understanding of how fibres perform, but how they can
be modified to improve their performance.

Cells with secondary cell walls are generally dead at maturity. However, the monocotyledon
bamboo has living fibres characterized by a multi-layered secondary cell wall structure. It consists
of an alternation of thin layers with high microfibril angles and high lignin content followed by
thick layers with low microfibril angles and low lignin content. This type of architecture allows
the bamboo to have high mechanical properties.
 Structural composition of a fibre cell

6. FUNCTIONS OF THE DIFFERENT ULTRASTRUCTURAL COMPONENTS

The middle lamella and primary cell wall are hypothesized to function in cell-to-cell adhesion, cell
expansion and the determination of cell shape. Primary walls have two conflicting functions: they
have to be rigid to withstand the internal and external stresses and, at the same time, have to be
pliant to allow cell wall expansion during growth. In contrast, the secondary cell wall structure
provides high axial stiffness while at the same time providing high collapse and burst resistance.
As it constitutes approximately 80% of the cell wall, the S2 layer has a profound effect on the
properties of wood. The characteristics of the S2 layer (such as low microfibril angle, thickness
and high cellulose content) are important in generating the wood stiffness. The S1 and S3 layers
are relatively thin in comparison, but nevertheless play a critical role in increasing the elastic
modulus of the cell in the transverse plane. The role of the S1 and S3 layers is to reduce the
deformation of the S2 layer under compression and tension. The S1 layer acts as a reinforcing
layer preventing excessive radial expansion and rotation of the cell, while the S3 layer helps to
avoid sideway collapse when under hydrostatic tension forces. In Booker and Sell’s model, a
sandwich structure is composed of the S3, S2, S1, CML, S1, S2, S3 layers. This structure acts as
a vibrational dampening system, and the angle of cellulose microfibrils changes the direction of
crack propagation during fracture. The fibres will slightly rotate under stress as a result of the
microfibril angle arrangement of the S2 layer. This puts the lignin-rich CML under shear stress,
and the rubbery nature of the lignin dissipates the stress energy. Overall, the functions of the cell
wall are to keep the plant upright, to take up and translocate water and nutrients and to defend from
external abiotic stress and biotic attack. It is also a system in charge of sensing, processing and
responding to internal and external cellular signals.

7. MOLECULAR COMPONENTS OF THE FIBRE CELL WALL

• CELLULOSES
Cellulose is a linear chain polysaccharide, consisting of repeating β-(1,4) linked glucose units,
with cellobiose (two glucose) being the repeat unit. Each glucose contains three free hydroxyl (-
OH) moieties that can interact to form hydrogen bonds. These bonds play a critical role in the
aggregation of cellulose chains and determine the crystal structure of the cellulose. These
crystalline chains form microfibrils in two arrangements, cellulose Iα and Iβ. The
thermodynamically more stable cellulose Iβ form is predominant in terrestrial plants.

Cellulose chain length or degree of polymerization (number of glucose monomers) is variable

(2000–6000) in primary walls and more homogenous (10,000) in secondary walls. The cellulose
chains are synthesized in the plasma membrane (the cell membrane) by a rosette-like CesA protein
complex consisting of six units that each have six sub-units. This has led to various
hypotheses regarding the number of cellulose chains in microfibrils. The initial ‘bundling’
hypothesis described the presence of 36 cellulose chains, matching the number of CesA proteins
per rosette. Such microfibrils would have a hexagonal arrangement. Other researchers have
suggested 18 or 24 chains per microfibril or variations in the number of cellulose chains per
microfibril with a variety of cross-sectional shapes. A further model of two 18-chain microfibrils
‘twinned’ to give an apparent 36-chain microfibril has been suggested. A microfibril of 18–24
chains should be 2–3 nm wide.
 • HEMICELLULOSES

Hemicelluloses are a group of heterogeneous polymers that play a supporting role in the fibre
wall. 20–30 % of the dry weight of wood consists of hemicelluloses. The hemicellulose polymers
are built up of several different monomers, such as mannose, arabinose, xylose, galactose, and
glucose. Some acidic sugars like galacturonic acid and glucuronic acid are also constituents of
hemicelluloses. One, two or several types of monomers usually build up the backbone of the
hemicellulose polymers. Most of the hemicelluloses also have short branches containing types of
sugars other than those of the backbone. The degree of polymerisation for the hemicelluloses is
between 100 and 200.

In softwood, the hemicellulose galactoglucomannan, makes up about 16 % of the dry weight of

the wood. Galactoglucomannans have a backbone of (1-4)-linked β-D-glucose and β-D-mannose
units, with α-D-galactose linked to the chain through (1-6)-bonds. An important structural feature
is that the hydroxyl groups at C(2) and C(3) positions in the chain units are partially substituted by
O-acetyl groups, on the average one group per 3–4 hexose units.
The hemicellulose arabinoglucuronoxylan constitutes about 8–9 % of the dry weight of softwood.
It has a backbone of (1-4)-β-D-xylose, where most of the xylose residues have an acetyl group at
C(2) or C(3). About every tenth xylose unit also has a 4-O-methyl-α-D-glucuronic acid residue
linked by a (1-2)-bond. The backbone substitution and degree of branching can vary considerably
between hemicelluloses of the same category.

• LIGNINS

Lignins are heterogeneous three-dimensional polymers that constitute approximately 30 % of the

dry weight of wood. Lignin limits the penetration of water into the wood cells and makes wood
very compact. Lignins are complex polymers based on the three monolignols: p-coumaryl alcohol,
coniferyl alcohol and sinapyl alcohol. The monolignols in various proportions are the building
blocks for the 3-D structure of native lignin in higher plants. The monolignols units are considered
to polymerise mainly according to a radical polymerization process.
Average composition of lignocellulosic material in different types of wood : (a) hardwood;
(b) softwood; (c) monocotyledon
LESSON 2: THE REACTION WOOD ANATOMY

1. INTRODUCTION
“The rings on the cross-section of the branch of a tree show the number of its years,
and the greater or smaller width of these rings show which years were wetter and
which drier. They also show the direction in which the branch was turned, for the part
that was turned towards the north grows thicker than that turned towards the south
so that the centre of the stem is nearer to the bark that faces south than to that on
the north side.” Leonardo da Vinci.

Leonardo published his observations of stem asymmetry in his notes for a treatise on
painting without any attempt at explanation. It must represent one of the earliest
references to reaction wood in the literature, although there can be no doubt that
carpenters and joiners had long been intuitively aware of its effects on the working
and mechanical properties of timber. With the passage of time our understanding of
why and how it is formed in the tree has increased, providing a scientific basis for
folk knowledge, but despite extensive research, much remains to be explained.

2. DEFINITION OF REACTION WOOD

Reaction wood has been defined by the Committee on Nomenclature of the

International Association of Wood Anatomists (IAWA, 1964) as: “Wood with more or less
distinctive anatomical characters, formed typically in parts of leaning or crooked stems and in
branches and tending to restore the original position, if this has been
disturbed. It is divided into two types: tension wood in dicotyledons and compression
wood in conifers”.

The Committee further defines compression wood as “Reaction

wood formed typically on the lower sides of branches and leaning or crooked stems
of coniferous trees and characterized anatomically by heavily lignified tracheids that
are rounded in transverse section and bear spiral wall checks; zones of compression
wood are typically denser and darker than the surrounding tissue”. Tension wood is: “Reaction
wood formed typically on the upper sides of branches and leaning or crooked stems of
dicotyledonous trees and characterized anatomically by lack of cell
wall lignification and often by the presence of an internal gelatinous layer in the
fibres”.

The Oxford English Dictionary provides several definitions of the word “reaction” several of
which encompass the nature and function of the term when used in conjunction with wood. Perhaps
the two most appropriate are: “The response made by a system or an organ to an external stimulus”
and “A movement towards a reversal of an existing tendency or state of things………or desire to
return, to a previous condition of affairs”. The first definition is appropriate to the formation of
reaction wood, while the second is appropriate to its function in the tree.

Briefly, reaction wood is formed in response to mechanical stress experienced by a

tree. Its formation can work to restore vertical growth (gravitropy) in main stems,
providing the stem is not already too thick to make this possible. It can be used also
to incline stems in order to move the canopy in towards light (heliotropy). In the case
of a branch, reaction wood formation is carefully controlled to balance its
continuously increasing weight, either as a buttress in the case of compression wood
in gymnosperms, or as a cantilever, in the case of tension wood in angiosperm
dicotyledons, thereby maintaining the branches pre-ordained orientation and the
architecture of the tree. It is noteworthy that reaction wood in a branch does not tend
to force the branch into a vertical alignment unless the dominance of the apical shoot
is lost.

However, reaction wood is required to change the orientation of a lateral

branch to the vertical in the event of damage to or loss of the leading shoot.
Compression wood and tension wood sectors in the stem are always associated with
local growth stresses which are very different from the normal tensile stress state
common to gymnosperms and angiosperms: compressive stress in the case of
compression wood, very high tensile stress in the case of tension wood.
There are, however, as will become apparent, variations on the theme. For example,
compression wood may form around the entire growth ring in straight vertical fast-growing conifer
stems. This may be a result of almost continual movement in the
wind of the long, recently formed apical internodes, which are highly flexible owing to
the high microfibril angle in the S2 layer in the juvenile tracheids. Also, gelatinous
fibres of the type normally associated with tension wood are sometimes found
distributed randomly in vertical stems of fast-growing hybrid aspen. These
phenomena might be explained by the existence of extraordinary growth stresses in
fast growing trees or by some variation in the level of growth regulators.
Some workers have observed cases in which the “normal” pattern of reaction wood
formation was not found. For instance, Höster and Liese (1966) described
compression wood in angiosperm species whose main axial elements were
tracheids, observations confirmed by Yoshizawa et al. (1993) and Baillères et al.
(1977). In contrast, Jacquiot and Trenard (1974) described gelatinous fibres in
coniferous wood.

The reason why branches and many tree stems are elliptical in cross section, with
growth rings having different widths on opposite sides, and pith located to the side of
the narrower growth rings, was already a subject of investigation in the nineteenth
century. It was noted that in conifers growing on mountain slopes more growth
occurred on the side of the stem facing the slope. Attempts were made to explain
this in terms of nutrient distribution to the cambium, in that nutrients moved
preferentially to areas to stimulate growth. Büsgen and Münch (1929), pointed out
that in fact the opposite was the case, as suggested by Cotta (1806), in that growth
stimulates the movement of nutrients to where they are required. They suggested
that this process was set in motion by stimuli which at that time were unknown. They
also noted that in Germany, where south-west and west winds predominate, conifer
stems take on an elliptical form with the long axis of the ellipse parallel to the wind
direction and greatest growth on the leeward side of the stem. Similarly, they noted
that in leaning conifer stems, greatest growth occurred on the lower side. Thus the
tree presents its least flexible profile to the prevailing stress. It was also noted that
those roots aligned with the direction of the stress, whether wind or gravitational pull,
also developed an elliptical profile. They proposed that this helped to prevent the stem from falling
over.
The facts that in conifers, reaction wood is produced on the underside of leaning
stems and branches under compressive stress, and that it has a reddish hue, led to
its being referred to in the German literature as Druckholz (pressure wood) or
Rotholz (red wood). These terms were supplanted by the name compression wood
as it was believed to be formed as a result of the tissue being under a compressive
load. In contrast, reaction wood produced in angiosperm dicots, which is formed in
tissues under tensile stress, and which is light in colour was referred to as tension
wood or Weissholz (white wood). As Dadswell and Wardrop (1949) pointed out, the
latter name is confusing as it was also used to describe wood formed in conifers on
the opposite side of the stem to Rotholz. The terms compression wood and tension
wood eventually acquired universal acceptance as reflecting the stress conditions under which they
are usually formed. However, there are circumstances in which tension wood can form in tissues
under compressive stress and vice versa.

General overview of the phenomenon of reaction wood

 3. STRUCTURE OF REACTION WOOD

The anatomical and structural features of reaction wood are usually described by
comparison with so-called normal wood. The latter term actually refers to wood
which has those properties most desired by the timber industry, for example, straight
grain, high density, high bending strength and uniform shrinkage without distortion
on drying. Interestingly there appears to be no definition of normal wood and it is
noteworthy that the IAWA glossary in its definitions of reaction wood, tension wood
and compression wood avoids the use of comparatives. Those features usually associated with
what authors refer to as normal wood and which are responsible for the desirable characteristics
of good timber are determined by the anatomical structure of the wood and the structure of fibre
(hardwoods) and tracheid (softwood) walls.

In particular, tracheids and fibres have the classical three-layered secondary wall with the
microfibril angle in the S2 layer being small, resulting in a high modulus of elasticity. In reacting
to gravity-induced displacement of the stem or branch, the tree produces fibres or tracheids whose
structure differs to a greater or lesser extent from this “ideal”.

3.1 CHANGES IN MACROSCOPIC APPERANCE

The macroscopic appearance of compression wood is often described as darker in

colour, varying in different species from brown to dark reddish brown. Its occurrence is associated
with eccentricity of the stem, the pith being further away from the side containing compression
wood. The growth ring is therefore normally wider on the lower side of stems and branches and in
species with distinct growth rings the latewood in the compression wood is wider and more
marked. This coupled with the greater proportion of lignin in the cell wall makes the wood denser,
more impermeable and stronger in compression.

Tension wood in angiosperms is not always as conspicuous as compression

wood in gymnosperms. They have been found to contain a lower proportion of lignin than normal
wood, giving it a whiter appearance. It is normally also associated with eccentricity of the
stem or branch with the wider rings normally being on the upper side of the stem
or branches, i.e. on the tension wood side. However, some authors demonstrated a lack of
eccentricity with tension wood occurrence or eccentricity opposite to the tension wood.
 Macroscopic features of compression wood (CW) and tension wood (TW)

3.2 CHANGES IN MICROSCOPIC FEATURES

The tracheids are more rounded in cross-section with the result that intercellular spaces are formed.
They are also shorter than normal wood tracheids in the same tree. Cell walls of
compression wood tracheids are normally thicker than those of normal wood
tracheids in the same part of the tree.

The structure of tension wood fibres is less clear-cut than that of compression wood
tracheids. Tension wood fibres are longer than those in normal wood.

Comparative anatomy of compression wood and normal wood for the conifer species
CW: compression wood; NW: normal wood
3.3 CHANGES IN THE ULTRASTRUCTURE

The microfibril angle in the S2 layer is larger, which reduces the tensile
strength, modulus of elasticity and increases the brittleness of the wood, making it
unsuitable for uses in which it is likely to experience high stresses. The larger
microfibril angle also means that the wood has a higher longitudinal shrinkage on
drying, but a lower transverse shrinkage. This explains the distortion on drying of
pieces of wood containing both normal and compression wood. In severe cases the
S3 wall layer is absent and the S2 layer contains splits which lie parallel to the
microfibril angle.

Tension woods are most commonly described as lacking an S3 layer and having
variable amounts of the S2 remaining, inside which is a gelatinous layer composed
mainly of hydrated cellulose microfibrils oriented almost parallel to the long axis of the fibre. This
gives the wood a glistening gelatinous appearance when wet. Variations on this theme have been
reported, however. For example Faruya et al. (1970) reported the presence of a gelatinous layer in
fibres of Populus euroamericana which had retained their S3 layer, while Côté et al. (1969)
reported an S3 layer inside a gelatinous layer. More and more variations on the structure of tension
wood fibres are being reported, with numerous species apparently lacking a gelatinous layer in
their fibre walls. However, trees as we know them could not have
evolved without reaction wood, a fact which needs to be borne in mind by those working to
improve wood quality. The formation of reaction wood demonstrates that the tree is capable of
fine tuning the structure of its fibre or tracheid walls to generate growth stress. It does this by
adjusting the proportions and arrangement of its major wall components, cellulose,
hemicellulose and lignin.
 Softwood Hardwood

Normal and reaction wood cell wall structures in gymnosperms (A,B) and angiosperms
(C,D). (A) tracheid in normal wood, (B) tracheid in compression wood, (C) fiber in normal
wood, (D) fiber in tension wood. G refers to the gelatinous layer.

Table 1: Summary of the differences between reaction wood and normal wood

Parameters Compression Wood Tension Wood

Macroscopic Darker colour; wider growth
appearance rings; more latewood
Physical properties High compression strength;
Higher wood density; Brittle
failure; Greater longitudinal
shrinkage; Lower pulp yield
Microscopic Shorter tracheids; Rounded cells
features with intercellular spaces
Ultrastructural Larger S2 angle; absence of S3
features
Chemical More lignin content; Less
properties cellulose
Lighter colour; Wider growth rings
Higher wood density; Lower
Compression strength; Seasoning
defects; Higher pulp yields; Poor
workability
Only fibres affected; Vessels not
affected; Shorter fibres
Presence of gelatinous layer which
could come after S3 or replace either
S3 or both S2 and S3. Microfibrils
less closely packed; Lower G-layer
angle; Higher S1 angle
More cellulose; Less lignin
 4. THE MECHANISMS OF REACTION WOOD

It is now well known that reaction wood effects stem reorientation by generating a
long lasting flexure momentum (Alméras et al 2005). This is linked to asymmetry
between wood production (new ring) on the two sides of the wooden axis:
asymmetry of generated growth stress and often asymmetry of ring width. But it is
not yet well understood how growth stress level is tuned during cell differentiation.
Various theories have been put forward, but none has so far proved satisfactory.

Brodski (1972) proposed that the water in the developing S2 layer of the
compression wood tracheid wall was replaced during maturation by a compound,
laricinan, and that this insertion provided the force needed to push the stem upright.
Boyd (1978) published a convincing argument against such a role for laricinan. The
tensional effect of gelatinous layer of tension wood fibres is also difficult to explain.
When gelatinous fibres are severed, the gelatinous layer may be observed to detach
itself from the other wall layers and contract. This is despite the fact that its major
component is cellulose microfibrils with their major axis parallel to the direction of
shrinkage. As the microfibrils are highly crystalline they would not be expected to
shrink in this way.

5. ECONOMIC IMPORTANCE OF REACTION WOOD

It is precisely those properties which enable reaction wood to carry out its function in
the tree that render it a problem for the timber industry. When Jaccard (1938) cut
loops in which reaction wood had formed, the curvature immediately changed as
internal stresses were released. In the case of the gymnosperm loop, in
which compression wood had formed on the convex side of the lower curve, and the
concave side of the upper curve, the effect of cutting the loop was to increase the
radius of curvature of the lower section, and decrease the radius of curvature of the
upper section. Thus, the compression wood appeared to have been exerting a
positive pressure prior to release. In the case of the angiosperm loop, where tension
wood formed on the upper sides of the loop, the effect was the opposite, with release
prompting a shortening of the tension wood side. These movements illustrate the
phenomena associated with growth stress release during wood processing, further
amplified by drying-induced deformations.

The commercial significance of such effects is enormous, since timber containing reaction wood
cannot be used where dimensional stability is required. Other problems associated with
compression wood include the difficulty of working the hard timber. Büsgen and Münch (1929)
commented that it is very difficult to drive a nail into it. It is also brittle and prone to brash
compression failure. The higher levels of lignin also increase costs for the pulp and paper industry
since lignin is expensive to remove and was hitherto difficult to dispose of. However, increasing
pressure on non-renewable natural resources mean that research is now looking into
utilisation of this abundant waste product.

Tension wood suffers from similar problems of dimensional stability and is also
difficult to work. The gelatinous fibres tear rather than cut, giving a wood surface
containing tension wood a woolly appearance. On the other hand, the lower levels of
lignin might be considered advantageous for the pulp industry.

If reaction wood were present in wood in only small amounts, these problems might
be ameliorated. However, the fact that tree growth takes place subject to
environmental pressures means that trees constantly have to make some reaction
wood to brace themselves against the wind, to correct for windthrow and to support
branches at the optimum angle. Even those trees which are perfect in form (from the
forester’s and timber merchant’s point of view), with straight and vertical stems, may
contain significant amounts of reaction wood. Vertical growth may only be achieved
by constant corrections of tendencies to lean under the influence of wind, whose
direction may change from day to day.