VARIATIONS INWOODANATOMY WITHIN SPECIES OF EUCALYPTUS* - Brill
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IAWA Bulletin n.s., Vol. 9 (1),1988: 13-23
VARIATIONS INWOODANATOMY WITHIN SPECIES OF EUCALYPTUS*
by
John Wilkes
Wood Technology and Forest Research Division, Forestry Commission ofNew South Wales,
P.O. Box 100, Beecroft, 2119, Australia
Summary
Eucalyptus species are characterised by 1978; Wilkes & Abbou 1983; Bamber
substantial genetically predetermined within- 1985):
and between-tree variations in wood anatom- Fibres (fibre-tracheids) - 10-20 ).l1ll in di-
ical features including fibre and vessel di- ameter, 0.8-1.3 mm long and occupying
mensions. This effect of genotype outweighs > 60% wood volume;
that of growing conditions; indeed the in- Vessels - solitary and diffuse, 80-180 11m
fluence on wood anatomy of rate of growth in diameter, 10-20% volume;
per se is minor. It must therefore be assumed Rays - 1-3-seriate, < 20 cells high, 10-20
that the reputed difference in wood quality %volume;
between regrowth/plantation stands and old Axial parenchyma - apotracheal and para-
growth forests is founded on non-anatomical tracheal, < 10% volume;
features, or reflects simply the greater pro- Vasicentric tracheids - < 2 % volume.
portion of juvenile wood in smalI, rapidly
Intraspecific variations in gross wood density
grown sterns.
closely reflect variations in the transverse di-
Key words: Eucalyptus, wood anatomy, ju-
mensions of fibres as indicated, for example,
venile wood, growth rate.
by the Runkel ratio (double wall thickness/
Introduction lumen diameter) (Chudnoff & Tischler 1963;
Native to Australia, Eucalyptus species are Sardinha 1977a). This relationship is here-
now grown extensively in many countries. after presumed; the majority of studies con-
The number of studies on variations in wood cerning the intrinsic wood quality of euca-
quality in the genus has correspondingly in- lypts deal not with wood anatomy per se, but
creased, and for certain species information with the more easily measured wood density
exists pertaining to the properties of wood which correlates weH with the usefulness of
produced under a variety of conditions. Sum- wood for a range of applications.
marised here are the variations in wood anat-
omy. Particular attention is focused on the Within-treevariation in wood anatomy
often reported differences between regrowth/ The most widely studied variations in
plantation ('new growth') stands and natural! wood anatomy of Eucalyptus are within-tree.
virgin Cold growth') forests (Dadswe1l1958; This age related variation, of obvious impor-
Knigge & Lewark 1976). The new growth tance in itself, must be thoroughly under-
forests are typically more vigorous than their stood if between-tree variations are to be in-
old growth counterparts at a similar age. terpreted accurately; not only may the mean
values of an anatomical feature differ between
Anatomy of Eucalypts trees, but the patterns of variation within
The eucal ypts generallylack distinct growth
trees mayaiso differ.
rings and have a relatively simple hardwood
structure (Fig. 1). The various cell types have Radial direction
the following common features in the majori- In the centre of older eucalypt sterns, a
ty of species (Dadswell 1972; Hillis 1972, zone of juvenile wood can be defined where
* This review formed the basis of a presentation to the XIV International Botanical Congress,
Berlin, July, 1987.
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via free access14 IAWA Bulletin n.s., Vol. 9 (1),1988
Fig.l.A.Eucalyptus regnans. Transverse section showingvessels in diffuse(tendingto oblique)
arrangement and the virtual absence ofaxial parenchyma. - B. E. tesselaris. Vessels often oc-
cur in short radial multiples, and paratracheal parenchyma is plentiful. These features charac-
terise only the subgenus Corymbia. x 35.
..... 20 1-5
N Z4
~
'e 150
o e
wz
.0
,a: w
~ ~
wu
15
\00
E
!1
%:
I-
-
>u
.;
e -
.....::L
a::
~
~~Wilkes - Wood anatomy of Eucalyptus 15
Fig. 4. Euca/yptus grandis. Transverse seetions of wood 2 (A) and 30 (B) annual increments
from the pith. x 35.
1·0 most anatomical features are changing rapid-
ly. Over the first 10-20 years of growth,
marked increases occur in the diameter, wall
~
.., 1·1 thickness and length of fibres (e.g. 10, 30
'eo and 50% respectively) (Santos & Nogueira
1971; Brasil & Ferreira 1972; Foelkel et a/.
1983; Tomazello Filho 1985a; Fig. 2). Simi-
>'"'
I- lar increases (e.g. of 50%) occur in the
CI)
Z diameter and length of vessel elements, while
W
Q 0,7 vessel frequency declines (e. g. by 50 %)
(Chudnoff & Tischler1963; Ranatunga 1964;
CI)
4( Arulchelvam 1971; Santos & Nogueira 1977;
1110>1 Nicholls & Griffin 1978; Figs. 3 & 4). The
proportions of the various cell types change
relatively little (Nicholls & Phillips 1970;
0' 5 L-..o...-...:..._"--_ _'--_---'
10 15
Sardinha & Hughes 1979; Malan 1985). In
RADIUS (cm) relation to the total amount of wood sub-
stance per unit volume, the centrifugal in-
Fig. 5. Radial gradients in density within the crease in fibre wall thickness more than com-
heartwood of 40-50-year-old stems of Euca- pensates for the increment in fibre diameter
lyptus (Sourees: Gerhards 1965; Wilkes & such that a rise in density of 10-50% nor-
Heather 1982; Wilkes 1984). mally occurs with distance from the pith in
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via free access16 IAWA Bulletin n.s., Vol. 9 (1),1988
.
BASIC DENSITY (g em- 3 ) FIBRE LENGTH (mm)
• 0.7
"
3•
.§
,..
J:
"iii
..
J:
• __J -_ _ _ _---:-'
12
RADIUS (ern)
• RADIUS (ern)
Fig. 6. Disnibution of density in Eucalyptus Fig. 7. Disnibution of fibre length in one
robusta (Sourees: Skolmen 1975; Wilkes, stern of Eucalyptus regnans (Souree: Bisset
unpublished). & DadsweIl1949).
Eucalyptus sterns (Gerhards 1965; Skolmen Between-tree variation in wood anat-
1974; Hans 1976; Nahuz et al. 1980; Chafe omy
1985a; Fig. 5). Variations in wood anatomy may be as
great between as within sterns of Eucalyp-
Axial direction tus (GreenhiII & Dadswell 1940; Sardinha
Variations in wood anatomy along the stern 1977b). This is the ease even for one uniform
are apparently less eonsistent than those in site, i.e., the effeet is largely genetie. Varia-
the radial direetion (e. g. see Barriehelo et al. tionsin fibrelengthandbasiedensity between
1983). Density eommonly inereases with neighbouring trees of the same age ean easily
height, sometimes after an initial decIine exeeed 25 and 50% respeetively (Table 1).
(Crawford et al. 1972; Ferreira 1972; Frank- The importanee of tree genotype in ex-
!in & Meskimen 1975; Harris & Young 1980; plaining tree-to-tree differenees in wood anat-
Chafe 1978; Fig. 6). Fibre length more often omy is echoed in studies showing signifieant
inereases to a point weIl up the bole, then de- variation between clones or famiIies (Rudman
cIines at higher levels (Ranatunga 1964; Bam- 1970; Waugh 1975; Nieholls & Matheson
ber et al. 1969; Santos & Nogueira 1974; 1980; Wang et al. 1984) and between prove-
Fig. 7). These ehanges can oeeur primarily in nances (Sesbou & Nepveu 1978; Vital &
wood formed by eambia of the same age, Lucia 1980; Tomazello Filho 1985a). The
i. e., at a given inerement from the pith, or lack of appreciable interprovenance variation
additionally, within individual increments, in the densitometric eharacteristics of E.
i.e., at a set inerement from the cambium nitens (Deane and Maid.) Maid. (Nicholls &
(Figs 6 & 7). Pederick 1979; McKimm 1985) is apparent1y
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via free accessWilkes - Wood anatomy of Eucalyptus 17
2-3
....
~
=
High 8Ir . . .
~
2·1
W
Z
~
UH l=1-1
% CI
~ Z
W
jH
~
...J 1.0
C W
~ a:
11
~H ii: 0"
11
ii:
0 · . L - - - - - - - ' -_ _ _ _ _....
1-3
0 3S 70 o so 100
HEIGHT (") RADIUS (,,)
Fig. 8. Variation in fibre wall thickness with Fig. 9. Contrasting radial gradients in fibre
height in two sterns of Eucalyptus saligna length in 27-year-old sterns of Eucalyptus
(Source: Sardinha & Hughes 1979). grandis of differing growth stress intensity
(Source: Malan 1985).
exceptional; however even in this species, 1977b; Bamber 1985). Differences in density
other features such as the length and lumen and fibre length, even between markedly
diameter of fibres, vary significantly with dissimilar sites, are mostly weIl below 20%
seed source (McKimm & nic 1987). (Table 2).
Within-tree gradients in wood anatomy also Wood property variations in eucalypts ap-
differ between trees. Sardinha and Hughes pear not to correlate reliably with changes in
(1979) for example, found that two trees of specific factors of the environment, such as
E. saligna Sm. from Angola showed reverse temperature I altitudellatitude, rainfall and
trends of fibre wall thickness with height in soil properties (Taylor 1974; Skolmen 1975;
the bole (Fig. 8). Malan (1985) has demon- Sardinha 1977b). Correspondingly, a range
strated that in E. grandis Hill ex Maid., an as- of forest management practices inc1uding
sociation exists between wood anatomy and fertilisation (Higgs&Rudman 1973; Chauhanet
levels of longitudinal growth stress in sterns, al. 1983),irrigation (Tischler&Heth 1985),
e.g. in the outer regions of highly stressed prescribed buming (Nicholls 1974) and thin-
boles the length (Fig. 9) and wall thickness ning/espacement (Brasil & Ferreira 1971;
of fibres, wood density and vessel diameter Higgs & Rudman 1973; Schonau 1974; Fer-
are likely to be higher than in boles of low rari & Scaramuzzi 1982; Chauhan et al.
stress. Such wood anatomy-stress interrela- 1983) all have re1atively weak effects on eu-
tions may prove an important link in explain- calypt wood structure.
ing the considerable within- and between-tree The infiuence of site on within-tree varia-
variation in wood properties (Nicholson et al. tions in wood anatomy has yet to receive
1975; Boyd 1980). adequate attention, although such an effect
has been reported for the density of E. gran-
Environmental variation in wood anat- dis (Villiers 1968).
omy
Variations in wood anatomy between ge- New and old growth forests
netically similar trees growing under differ- The relatively rninor role of growing con-
ent conditions are usually smaller than the ditions in determining the wood structure of
effects due to age and genotype (Bamber & Eucalyptus calls into question the reputation
Humphreys 1963; Tay lor 1973; Sardinha of regrowth and plantation stands as produc-
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Table 1. Examples of variations in wood anatomy between Eucalyptus trees.
Wood Species Age* Variation Difference Reference
property (yrs) Min. Max. (% ofmin.)
Basic E. alba 5 0.44 0.67 52 Ferreira 1970
density E. citriodora 8 0.51 0.75 47 Shukla & Rajput 1981
(g cm-3 ) E. grandis 3 0.30 0.48 60 Brasil et al. 1979
E.grandis 15 0.34 0.53 56 Taylor 1973
E.grandis 15 0.40 0.63 58 Taylor 1974
E. nitens 8 0.39 0.54 38 Chafe 1985b
E. regnans 27 0.39 0.49 26 Dargavel 1968
E. robusta 5 0.50 0.68 36 Arulchelvam 1971
E. saligna 3 0.40 0.58 45 Ferreira et al. 1979
E. saligna 3 0.35 0.51 46 King 1980
Fibre E. grandis 20 0.91 1.11 22 Bamber et al. 1969
length E. grandis 25-30 0.84 1.02 21 Ranatunga 1964
(mm) E. grandis 20 0.86 1.10 28 Taylor 1973
E. grandis 18 0.93 1.19 28 Taylor 1974
E. robusta 3 0.83 1.04 25 Arulchelvam 1971
E. tereticornis 8 0.66 0.88 33 Purkayastha et al. 1984
* Either age of tree or increment from pith.
Data represents either the whole tree or the lower stern, e. g. breast height.
ing wood very different from virgin forest While in E. delegarensis R.T. Bak. a elose
material, e. g. the new growth wood is be- relationship appears to exist between cambial
lieved to have a relatively low basic density activity durlng the growing season and the
and short fibres (Hillis 1981). The scenario structure of maturing xylem (Amos et al.
can be largely explained in tenns of the pro- 1950), more general analyses ofthe effects of
portion of juvenile wood in the sterns. The vigour on eucalypt anatomy are fraught with
period of fonnation of juvenile wood is large- complexities. Perhaps most importantly, dif-
ly predetennined (Dadswell 1958; Wilkes ferent stimuli of growth may be associated
1984); thus in trees harvested at a given size, with different variations in a wood property,
a larger proportion of juvenile wood is pres- e.g. fertilisation and thinning may have op-
ent in those more rapidly grown, as in plan- posite effects on density (Higgs & Rudman
tations. 1973). Further, increment in height and diam-
A depressive effect of accelerated growth eter may be associated differently with a
on the density and fibre length of tissues of a wood feature (Taylor 1974). It is also note-
given fonnative age is also widely assumed. worthy that the extent to which vigour and
However,supportive evidence (SusmeI1953; wood anatomy are linked genetically in euca-
Hans et al. 1972; Schonau 1974) is limited, lypts remains largely unknown. Were a sub-
and other results are more often obtained stantial inverse relationship to exist for den-
(e.g. Taylor 1973; Schonau 1980; Frederick sity or fibre length, much of the so-called rate
er al. 1982; Bhat & Bhat 1984; Wilkes 1984). of growth effect could be an artifact of ge-
This might be expected in view of the weak netic differences between trees. Sesbou and
relationship between growing conditions and Nepveu (1978) have in fact reported such an
xylem anatomy in Eucalyptus. association between density and growth rate
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via free accessWilkes - Wood anatomy of Eucalyptus 19
Table 2. Examples of variations in wood anatomy between Eucalyptus between locations.
Wood Species Age Variation Difference Reference
property (yrs) Site 1 Site 2 (% ofmin.)
Basic E. camaldulensis 9 0.55 0.59 7 Sesbou & Nepveu 1978
density E. citriodora 8-9 0.65 0.72 11 Shulda & Rajput 1981
(g cm- 3 ) E. grandis 15 0.43 0.46 7 Taylor 1973
E.grandis 18 0.45 0.48 7 Taylor 1974
E. robusta 34 0.54 0.63 17 Skolmen, 1975
E. saligna ca. 25 0.61 0.69 13 Sardinha & Hughes 1979
Fibre E. gomphocephala 3 0.59 0.74 25 Stern-Cohen & Fahn 1964
length E.grandis 15 0.85 0.98 15 Taylor 1973
(mm) E.grandis 15 0.92 1.02 11 Taylor 1974
E. tereticornis 8 0.75 0.82 9 Chauhan et al. 1983
E. tereticornis 8 0.74 0.80 8 Purkayastha et al. 1984
Data represent either the whole tree or the lower stem, e.g. stump height.
In most cases variation may be partly genetic, i.e. genotype was not held constant across the
sites.
In cases where more than two sites or seed sources were tested, the largest variation is pre-
sented.
Fig. 10. Eucalyptus grandis. Transverse sections from control (A) and intensively managed (B)
stems at 5 years of age. x 50.
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via free access20 IAWA Bulletin n.s., Vol. 9 (1),1988
aeross 25 provenanees of E. camaldulensis - & F. R. Humphreys. 1963. A prelirninary
Dehnh. Where tree genotype is held eonstant, study of some wood properties of Euca-
using either clonal or eoppiee material, direct lyptus grandis (Hill) Maiden. J. Inst.
'effeets' of vigour on wood strueture are Wood Sei. 11: 63-70.
found to be small and/or ineonsistent (King Barriehelo, L. E. G., J. O. Brito & A J. Migli-
1980; Wilkes & Abbott 1983; Wilkes 1984). orini. 1983. Estudo da variayiio longitu-
It may be signifieant that the majority of dinal da densidade basiea de Eucalyptus
work in this area has dealt only with density spp. Silvicultura 28: 726-731.
and fibre length (in effeet, fibre dimensions). Bhat, K.M. & K.V. Bhat.1984. Wood prop-
A study ofE.grandis by Bamber et al. (1982) erties of l-year-old Eucalyptus tereticor-
has shown a larger effeet of vigour on the nis Sm. Aust. For. Res. 14: 129-133.
physiologieally aetive eells; intensive eultural Bisset, I.J.W. & H.E. Dadswell. 1949. The
treatment was associated with an inerease in variation of fibre length within one tree of
the volume of parenehyma and a deerease in Eucalyptus regnans, F.v.M. Aust. For. 13:
vessel sizeand frequeney (Fig.lO). Clearly, in- 86-96.
suffieient is known regarding the physiology Boyd, J. D. 1980. Relationships between
ofwood formation inEucalyptus. Neverthe- fibre morphology, growth strains and
less, there ean be little doubt that the eritieal physieal properties of wood. Aust. For.
faetors governing wood anatomy are eambial Res. 10: 337-360.
age when tissues are formed and tree geno- Brasil, M. AM. & M. Ferreira. 1971. Varia-
type; growing eonditions are less important. yiio da densidade basica da madeira de
Eucalyptus alba Reinw., E. saligna Smith
Conc1usion
e E. grandis Hill ex Maiden aos 5 anos de
It appears that environmental variation in
idade, em funyiio do local e do espaya-
the anatomy of euealypts is generally over-
mento. IPEF 2/3: 129-149.
shadowed by both natural genetie diversity
- & - 1972. Variayiio da densidade basiea
and within-tree (age) effeets. Thus, the widely
e das earaeteristieas das fibras em Euca-
held view that wood properties vary mark-
/yptus grandis Hill ex Maiden ao nivel do
edly between new and old growth forests re-
DAP. IPEF 5: 81-90.
quires substantial qualifieation; the propor-
- , R.A.A.Veiga& H.A.Mello.1979.Den-
tion of juvenile wood is the critieal faetor.
sidade basiea de madeira de Eucalyptus
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