MICROFIBRIL ANGLE: MEASUREMENT, VARIATION AND RELATIONSHIPS
A REVIEW
Microfibril angle (MFA) is perhaps the easiest ultrastructural variable
to measure for wood cell walls, and certainly the only such variable that
has been measured on a large scale. Because cellulose is crystalline, the
MFA of the S
2 layer can be measured by X-ray diffraction. AutomatedX-ray scanning devices such as SilviScan have produced large datasets
for a range of timber species using increment core samples. In conifers,
microfibril angles are large in the juvenile wood and small in the mature
wood. MFA is larger at the base of the tree for a given ring number from
the pith, and decreases with height, increasing slightly at the top tree. In
hardwoods, similar patterns occur, but with much less variation and much
smaller microfibril angles in juvenile wood. MFA has significant heritability,
but is also influenced by environmental factors as shown by its
increased values in compression wood, decreased values in tension wood
and, often, increased values following nutrient or water supplementation.
Adjacent individual tracheids can show moderate differences in MFA
that may be related to tracheid length, but not to lumen diameter or cell
wall thickness. While there has been strong interest in the MFA of the
S
2 layer, which dominates the axial stiffness properties of tracheids andfibres, there has been little attention given to the microfibril angles of S
1and S
3 layers, which may influence collapse resistance and other lateralproperties. Such investigations have been limited by the much greater
difficulty of measuring angles for these wall layers. MFA, in combination
with basic density, shows a strong relationship to longitudinal modulus
of elasticity, and to longitudinal shrinkage, which are the main reasons
for interest in this cell wall property in conifers. In hardwoods, MFA is
of more interest in relation to growth stress and shrinkage behaviour
Key words:
Microfibril angle, cellulose microfibrils, X-ray diffraction,microscopy, wood properties.
Lloyd Donaldson
Cellwall Biotechnology Centre, Scion, Private Bag 3020, Rotorua, New Zealand
[E-mail: lloyd.donaldson@scionresearch.com]
IAWA Journal, Vol. 29 (4), 2008: 345–386
.