The Tasmanian forest industry is increasingly applying intensive
management practices to produce logs suitable for sawing from plantation
eucalypts. Such practices include green pruning necessary for producing
knot-free wood as well as thinning to encourage rapid diameter growth in
retained trees (Dudzinski et al. 1992; Gerrand et al. 1997). However,
one of the potential consequences of thinning and pruning operations is
wounding to the retained trees, resulting in defects such as kino vein
formation. Concern has been raised about the potential problem of sawlog
downgrade due to a high incidence of kino veins (Gerrand et al. 1997;
Somervile and Davies-Colley 1998; Vasiliauskas 2001).
Kino veins or pockets are commercially regarded as characteristic
defects of some Eucalyptus spp. (Nelson and Hillis 1978; Bamber 1985).
Kino veins, more commonly known as 'gum veins', can be the
most severe forms of defect in the wood of some eucalypt species
(Tippett 1986). Somervile and Davies-Colley (1998) found that kino
pockets had the potential to seriously reduce recovery of veneer-grade
product in a stand of E. regnans. Susceptibility to kino vein formation
varies with species and environmental conditions (Doran 1975; Nelson and
Hillis 1978; Tippett 1986). For example, trees growing on dry sites have
apparently been observed to be severely affected by kino formation (Day
1959). The exact cause of the formation of kino veins is still not
clearly understood, but it has been generally associated with stresses
that apparently damage but do not kill the vascular cambium. The
stresses include branch shedding, damage by biological agents (e.g.
insects and fungi), physiological stress, fire and mechanical damage
(Jacobs 1986). Most authors agree that kino veins generally play a role
in wound defence as a tree resistance mechanism, acting as a
'barrier zone' (Wilkes 1986; Hillis 1987). Barrier zones have
been defined as specialised tissue produced by the cambium in a
non-specific response to infection as well as to mechanical wounding,
and they serve to compartmentalise necrotic sapwood from living cambium
(Tippett and Shigo 1981).
The present study investigated the causes of kino vein formation in
eucalypt tree defence by comparing the responses of two commercially
important Eucalyptus species (E. globulus and E. nitens) to various
mechanical, chemical and biological wounding treatments. Both E.
globulus and E. nitens are widely planted in Tasmania for pulp and
sawlog purposes (Beadle and Mohammed 1999). Previous surveys of E.
nitens sawlogs suggest that this species does not readily form kino
veins (Nicholls and Pederick 1979; Yang and Waugh 1996a; Eyles and
Mohammed 2002) while a small amount of kino has been observed in sawn
timber of E. globulus (Yang and Waugh 1996b).
Materials and methods
Experimental site and wounding treatments
Studies were conducted at two experimental plantation sites growing
both E. globulus and E. nitens. The first site was located at Esperance
(about 2 km south of Dover, Tasmania--latitude 43[degrees]15'S,
longitude 146[degrees]50'E). The altitude is 240 m a.s.l., and mean
annual rainfall is 869 mm. The second site was at Lewisham (about 50 km
east of Hobart, Tasmania--latitude 42[degrees]49'S, longitude
147[degrees]37'E). The altitude is 20 m a.s.l., and mean annual
rainfall is 512 mm. Details of site establishment and tending for the
Esperance and Lewisham sites are provided by Beadle et al. (1996) and
White et al. (1998), respectively. At the Lewisham site, an additional
study was carried out to explore the effect of drought stress on tree
wound responses. The site had been divided into six blocks (replicates),
three designated as 'irrigated' and three designated as
'rain-fed'. The experimental design and irrigation treatment
have been outlined elsewhere (Honeysett et al. 1996; Worledge et al.
1998). Briefly, the irrigated blocks received 1300-1500 mm [y.sup.-1]
while the rain-fed blocks received 900-1100 mm [y.sup.-1] including
supplementary irrigation. Trees at Esperance were 16 y of age with an
average stand diameter of 24 cm and 26 cm for E. nitens and E. globulus,
respectively. The trees at Lewisham were 9 y of age. The average stand
diameters of the irrigated trees were 19 cm and 19.5 cm for E. nitens
and E. globulus, respectively. The average stand diameters of the
rain-fed trees were 16 cm and 16.3 cm for E. nitens and E. globulus,
In late summer of 2000, twelve trees (six E. nitens and six E.
globulus at Esperance and ten trees each of E. nitens and E. globulus
(five from the irrigated block and five from the rainfed blocks at
Lewisham) were randomly selected for wounding treatments. The
circumference of each tree was divided into four equal quadrants at
about 1.3 m above ground-level. Four treatments were randomly assigned
to the quadrants, one per quadrant as follows:
1. the impact force of a 3 kg steel rod (5.7 cm diameter x 15 cm
long) attached to a rope 1 m long swinging through an arc of 90[degrees]
(Fig. 1). This stimulated the 'bruising' type wound commonly
inflicted on retained trees during thinning operations.
2. an injection of 0.5 mL of 0.2% CEPA aqueous solution
(2-choroethyl-phosphonic acid, SIGMA) into a small wound (1 [cm.sup.2])
created by a chisel. CEPA releases ethylene under physiological
conditions. The application of exogenous ethylene to induce production
of tree secretions has been well documented in many tree species
including Eucalyptus (Dowden and Foster 1973; Nelson and Hillis 1978;
Hillis 1987). Anatomical and histochemical responses to CEPA treatment
were recently reported for juvenile E. globulus and E. nitens (Eyles and
3. a hole 10 mm in diameter drilled 3 cm into the outer sapwood to
simulate the stem wounds commonly associated with stem-boring insects. A
second hole with identical dimensions to the first was drilled 30 cm
above the first hole and inoculated with a decay fungus previously
isolated from a white-rot decay column in E. nitens. Inoculation
involved inserting rectangular pieces, 2 cm x 1 cm, of a 2-week-old
culture growing on malt extract agar into the drill hole and sealing it
with a layer of lanolin wax. The decay fungus used for the inoculation
is an unidentified basidiomycete known as 'Isolate D'. The
fungus is described in Barry (2001).
4. a block of dry ice (approx. 7 [cm.sup.2]) pressed firmly onto
the bark for 5 min. This treatment attempted to damage the vascular
cambium without physically disrupting the outer bark so as to minimise
the exposure of injured tissue to the external environment (Robinson
About three months after treatment, three trees each of E. nitens
and E. globulus at each of the Esperance and Lewisham sites were
harvested. Billets ~50 cm in length that included the wounding
treatments were taken to the laboratory. Each wounding treatment was
excised from the billet by cutting longitudinally through the centre of
the wounds using a bandsaw. The production of dark extractives was
consistently observed as part of the wound response, to varying degrees,
amongst all treatments. Therefore, responses were classified according
to whether dark extractives (defined as non-structural components of
wood and bark) were produced within specialised secretory cells, that is
kino veins, or within new wound-associated wood formed directly adjacent
to the wounding site (Figs 3 and 4). For clarity, we use the term
'kino' to refer to the exudate produced in kino veins and we
refer to other exudates, not formed in kino veins, as 'dark
extractives'. In cases where kino veins had been formed, the length
and width of the kino veins were recorded.
[FIGURE 1 OMITTED]
Wood samples of each wound response (a cube with edge of about 15
mm) were fixed in FAA (formaldehyde:acetic acid:70% ethanol, 5:5:90
v/v/v) for a minimum of 24 h at 4[degrees]C, dehydrated with an ethanol
series and infiltrated in a LR White acrylic resin (Proscitech,
Brisbane) series. Samples were then polymerised in fresh 100% LR White
resin for 8-10 h at 60[degrees]C (Eyles and Mohammed 2002). Transverse
and longitudinal sections, 10 mm thick, were cut with a sledge
microtome, dried onto glass slides, stained with a 1% aqueous solution
of toluidine blue and permanently mounted in Cytoseal (ProSciTech,
Brisbane). Tissue structure and cell changes were noted on a Zeiss
This procedure was repeated with the remaining trees, about 13 mo
and 17 mo after setting up the experiments at Esperance and Lewisham,
[FIGURE 2a OMITTED]
[FIGURE 2b OMITTED]
[FIGURE 3a OMITTED]
[FIGURE 3b OMITTED]
Since this qualitative study aimed to investigate the type of
wounding treatments that might induce kino vein formation, the results
were interpreted simply as positive or negative. Therefore, the
formation of kino veins, regardless of size, was regarded as a positive
response to the wounding treatment. Only data from the CEPA treatment
was considered for statistical analysis, because kino veins were
generally formed only in response to this chemical treatment. A [chi
square] contingency analysis was used to test whether kino vein
incidence was species-dependent. Due to the small number of replicates,
the data obtained from trees harvested at 3 mo and those at 17 mo at
both Esperance and Lewisham (including trees from the irrigated
treatments only) were pooled.
Results and discussion
Response to CEPA treatment
The incidence of kino veins in response to CEPA treatment was
significantly higher for E. globulus than for E. nitens (P < 0.001).
Kino veins were induced in the xylem of all 11 E. globulus treated with
CEPA at both Lewisham and Esperance (Fig. 2a and Table 1).
The kino veins were 15-177 cm in length and 4-7 cm in width.
Microscopic examinations showed that the general structure of kino veins
produced in E. globulus was similar to that described for other eucalypt
species including E. obliqua (Skene 1965), E. radiata (Dowden and Foster
1973), and E. wandoo and E. marginata (Tippett 1986). Typical kino veins
consisted of a tangential series of radial and transverse parenchyma
bridges that linked cavities filled with kino (Fig. 2b).
Kino veins were induced in only three out of the 11 E. nitens trees
treated. Furthermore, in two of the three trees, the kino veins were
less than 15 cm long and 4 cm wide. These results confirm findings from
previous studies that kino veins are not readily induced in E. nitens,
unlike other Eucalyptus species. Instead, in half of the E. nitens
replicates, the CEPA treatment induced the production of a thin layer of
dark extractives (ranging from 8 to 24 cm in length) within the phloem.
Microscopic sections showed that this layer was anatomically distinct
from kino veins. In contrast to the regular arrangement of kino veins,
the thin layer of extractives was composed of traumatised parenchyma of
varying shape and size that stained strongly for the presence of
polyphenolic compounds. Moreover, unlike kino veins, which are derived
from the initials of the vascular cambium, the traumatised parenchyma of
E. nitens was observed to originate from the de-differentiation of
pre-existing phloem parenchyma.
[FIGURE 4 OMITTED]
[FIGURE 5 OMITTED]
[FIGURE 6 OMITTED]
[FIGURE 7 OMITTED]
Nelson and Hillis (1978), studying the involvement of ethylene in
kino vein formation in E. regnans, found the area of kino veins produced
was a positive logarithmic function of the CEPA concentration. They
recommended that 1 mL of 0.1% ethrel (a commercial preparation of CEPA)
solution would be adequate for studies in the production of kino veins.
It could be argued that application of higher concentrations than those
used in the present study (0.5 mL of 0.2% CEPA) might have led to a
higher incidence of kino veins, particularly in E. nitens. However, in
another study, injection of even 0.3 mL of 1% CEPA solution failed to
induce kino vein formation in juvenile E. nitens trees (Eyles and
As mentioned earlier, in addition to observed genetic differences
in kino vein occurrence between species, environmental factors may also
contribute to the incidence of kino vein formation (Hillis 1987). Dry
sites have been observed to be severely affected by kino vein formation,
and Day (1959) suggested that water deficiency during periods of active
growth might explain this observation. The effect of water stress on the
incidence of kino vein formation has not been studied. In this study,
kino veins were not induced in the xylem of any of the five E. nitens
trees grown under simulated drought conditions. More significantly, kino
veins were induced in only three out of the five E. globulus trees.
While it is difficult to draw any firm conclusions from these results,
they appear contrary to Day's explanation, especially considering
that water deficits inhibit division of fusiform initials and xylem and
phloem mother cells (Kozlowski and Pallardy 1997). The relationship
between water stress and kino vein formation warrants further
investigations, as future plantations may have to be established on
Response to dry ice treatment
In a few trees, the dry ice treatment killed the underlying
vascular cambium, but in the majority of trees damage by the dry ice
treatment was restricted to the phloem. The severity of damage did not
appear to be directly related to bark thickness. Instead, differences in
surface area contact and pressure applied in the dry ice treatments
might explain the varying severity of damage--even though every effort
was made to ensure even contact between dry ice and bark. The responses
were generally characterised by the production of callus tissue with the
deposition of a layer of dark extractives at the interface of healthy
bark and bark that had been killed by the dry ice application. The
amount of deposition was variable, appearing either as a continuous
layer of variable width (Fig. 3a) or as irregular deposits along the
necrotic margin. Microscopic examinations showed that the lesion margin
was composed of a completely differentiated wound periderm comprising
multiple layers of thick-walled phellem in addition to traumatised
parenchyma cells often filled with polyphenols, which had been derived
from the de-differentiation of pre-existing phloem parenchyma cells
(Fig. 3b). Both E. globulus and E. nitens trees responded in this
manner. Some of the E. nitens trees at Lewisham, however, produced
different coloured extractives in the xylem and phloem. Instead of the
usual deposition of dark extractives, some E. nitens replicates produced
a distinctly pink band of extractives located in the ray and axial
Response to drill wounding with or without fungi inoculation
Three months after treatment, both species had responded to drill
wounding, with or without fungal inoculation, by producing callus tissue
along the margins of healthy sapwood. The exposed callus was usually
covered with a thin film of dark extractives. At Lewisham, the drill
holes of irrigated trees were fully closed over by callus, unlike those
trees growing under rainfed conditions. The slower rates of wound
closure in trees subjected to drought could increase the time during
which trees are exposed to microbial infection. Previous investigations
of seasonal responses of E. maculata after artificial wounding found
evidence that the rate of wound closure, whether by production of new
wood or deposition of exudates, could be important in restricting the
colonisation of wounds by decay fungi (Mireku and Wilkes 1989).
Furthermore, a clonal trial selecting for tolerance of E. grandis to
Cryphonectria canker found that clones with the fastest rate of wound
closure were less susceptible to infection (Van Zyl and Wingfield 1999).
Seventeen months after injury, all drill hole wounds (even those
trees exposed to drought at Lewisham) had fully occluded and the
vascular cambium had been re-established. In both species, varying
amounts of dark extractives were observed in the wound wood, whether
fungal colonisation had occurred or not (Fig. 4). Kino veins had formed
in response to drill wounding in four of the twenty-two E. globulus
trees. Kino veins were not associated with drill wounding in any E.
nitens trees. Drill holes inoculated with the unidentified decay fungus
had formed decay columns generally >10 cm in length in both species.
Fungal hyphae were visibly abundant in the drill hole and some were
observed to be in intimate contact with the inner side of the dark
extractives. Microscopic examination revealed that the hyphae had not
spread into the wood formed after wounding, suggesting that the
wound-associated wood may be an effective barrier against further fungal
colonisation of wood formed subsequent to damage.
Response to bruising treatment
In both species, the impact from the 3 kg steel rod generally
killed the bark as well as the underlying vascular cambium that was
located directly under the area of impact and surrounding margins.
Subsequent to the impact, the overlying bark separated from the vascular
cambium, causing it to become detached from the bole while remaining on
the stem (Fig. 5). Assessment of the level of damage was usually
possible only after removal of the dead bark. In these cases, the
bruising treatment was observed to cause extensive discolouration of the
underlying xylem (Fig. 6). Furthermore, kino veins (<5 cm long)
and/or dark extractives were common along the outer edges of the callus
margins, particularly in E. globulus trees. Another feature consistently
associated with this treatment was the occurrence of bark splitting,
which provided potential entry points for insect and fungi (Fig. 5).
Larvae and hyphae were observed in a number of trees in both species.
Despite the preliminary nature of this study, the results suggest
that assessment of damage levels in retained trees after thinning
operations may be underestimated since bark-covered wounds are not as
visible as bark-torn wounds. This finding may have important
implications to industry, particularly as decay and discolouration have
been reported to occur at a greater rate in bark-covered than open
wounds over a two-year study period (Dudzinski et al. 1992; White and
Role of kino and dark extractives in wound repair in eucalypts
In previous studies of eucalypt host-pathogen interactions, any
exudate produced by the tree was simply referred to as kino (Mireku and
Wilkes 1989; Wardlaw 1999). There, kino was defined as 'a red-brown
aqueous polyphenolic exudate with the main constituent being polymerised
forms of proanthocyanidins otherwise known as condensed tannins'
(Hillis and Carle 1962; Hillis and Yazaki 1975). Recent studies using
modern analytical techniques including High Performance Liquid
Chromatography coupled with Electrospray Ionisation Mass Spectrometry
(HPLC with ESI-MS) have shown the chemical composition of kino to be
markedly different from that of dark extractives detected in wound wood
(Eyles et al. 2002, 2003; Eyles unpublished data). Given that there are
clear differences in the chemistry and origin of these extractives, we
suggest that future studies investigating the role of exudates in a
eucalypt's defence should fully detail the nature of the exudates,
particularly as these differences may reflect different roles in wound
healing and defence. For example, in the present study, the production
of dark extractives was shown to occur readily in the xylem and/or the
phloem in both species--even in E. nitens, a species that does not
usually form kino veins.
The production of tree exudates within specialised intercellular
secretory spaces in the xylem and/or phloem is found in a range of woody
tree species (e.g. latex from Hevea species, gum from Prunus species and
resin from Pinus species; Hillis 1987). While they are generally
implicated in protection against pests, diseases and damage, the precise
nature of their involvement remains unclear (Pearce 1996). There is
conflicting evidence of the role of kino vein formation as a defence
mechanism in host-pathogen interactions. In one study, the kino veins
were significantly larger in trees inoculated with various canker fungi
than in control trees (Old et al. 1986). In another report, kino vein
formation was not correlated with size of lesions formed in response to
the root rot, Phytophthora cinnamomi in E. marginata (Tippett et al.
1983, 1985). Given that fungal hyphae were observed to grow intimately
with dark extractives--as also seen by Wilkes (1986)--it would seem that
the induction of exudates in eucalypts is primarily in response to
wounding and not specifically against biotic agents such as fungi. Even
other forms of wounding not considered in this experiment, such as
pruning, were shown to induce kino veins in the absence of fungal decay
(Fig. 7). Hanks et al. (1999) found no evidence to suggest that kino
served to defend E. rudis trees against Phoracantha semipunctata
(colonising phloem-boring larvae), particularly as production of kino
veins usually requires days or even weeks whereas beetle larvae can
penetrate the bark and reach the cambium within 24 h. Investigations of
the wound reactions of Scots pine (Pinus sylvestris L.) to attacks by
two bark beetles (Lieutier et al. 1995) similarly concluded that the
induced production of resin was a wound reaction arising from mechanical
stress due to insect boring, and not solely a defence reaction against
biological agents such as fungal pathogens. This does not mean, however,
that these dark extractives may not have a role in impeding the spread
of fungi or insects. For example, kino veins have been observed to
effectively prevent the outward spread of wood decay (White and Kile
The production of exudates, either within specialised kino veins or
in less organised wound-associated wood or bark, was consistently
observed as part of host response to dry ice, bruising and drill
wounding treatments, regardless of colonisation by fungi. The integrity
of the vascular cambium appeared to be the key factor in kino vein
production, as has been found in previous studies. Any stress, whether
physiological, mechanical or chemical, that damages but does not kill
the vascular cambium has the potential to cause kino veins to be
produced in a susceptible species. Kino veins are not readily induced in
E. nitens--unlike other Eucalyptus species--providing some evidence that
this species is suitable for the commercial production of kinofree
timber. In contrast, as kino veins were readily induced in E. globulus,
kino vein defect may be a problem when trees of this species are grown
We thank Dr Chris Beadle for access to CSIRO field sites, Messrs
Malcolm Hall, Dale Worledge and Jean Richmond for their assistance with
fieldwork, Dr David Ratkowsky for statistical analysis and Dr Bill
Neilsen for devising the 'bruising' wounding treatment. We
thank Mr Tim Wardlaw, Dr Ken Old and Mr Peter Bennet for reviewing
drafts of the manuscript. A.E. was supported by a University of Tasmania
Strategic Scholarship with additional funding from the Co-operative
Research Centre for Sustainable Production Forestry.
Revised manuscript received 11 December 2002
Bamber, R.K. (1985) The wood anatomy of eucalypts and papermaking.
Appita 38, 210-216.
Barry, K.M. (2001) Antimicrobial defence in Eucalyptus nitens
sapwood. PhD dissertation, University of Tasmania, Australia.
Beadle, C. and Mohammed, C. (1999) Pruning for plantation sawlogs.
Onwood Research Updates No. 24. CSIRO Forestry and Forestry Products,
Beadle, C., Turnbull, C.R.A. and Dean, G.H. (1996) Environmental
effects on growth and kraft pulp yield of Eucalyptus globulus and E.
nitens. Appita 49, 239-242.
Day, W.R. (1959) Observations of eucalypts in Cyprus; the character
of gum veins and anatomical indications for their origin. Empire
Forestry Review 38, 35-44.
Doran, J.C. (1975) Occurrence of kino veins in two provenance
trials of Eucalyptus regnans. Australian Forest Research 7, 21-27.
Dowden, H. and Foster, R.C. (1973) Kino veins, their formation,
anatomy and fine structure. Proceedings of 16th Forest Products
Conference. CSIRO, Victoria, pp. 1-5.
Dudzinski, M.J., Old, K.M. and Gibbs, R.J. (1992) Minor stem damage
to silvertop ash from thinning operations. A report to the Department of
Conservation and Environment, Victoria. CSIRO. 33 pp.
Eyles, A. and Mohammed, C.L. (2002) Comparison of CEPA
(2-chloroethyl phosphonic acid) induced responses in juvenile Eucalyptus
nitens, E. globulus and E. obliqua: a histochemical and anatomical
study. International Association of Wood Anatomists Journal 23, 419-430.
Eyles, A., Davies, N.W. and Mohammed, C. (2002) Novel detection of
formylated phloroglucinol compounds (FPCs) in the wound-associated wood
of Eucalyptus globulus and E. nitens. Journal of Chemical Ecology 29,
Eyles, A., Davies, N.W. and Mohammed, C. (2003) Wound wood
formation in Eucalyptus globulus and E. nitens: anatomy and chemistry.
Canadian Journal of Forest Research. In press.
Gerrand, A.M., Neilsen, W.A. and Medhurst, J.L. (1997) Thinning and
pruning eucalypt plantations for sawlog production in Tasmania.
Tasforests 9, 15-33.
Hanks, L.M., Paine, T.D., Miller, J.G., Campbell, C.D. and Schuch,
U.K. (1999) Water relations of host trees and resistance to the
phloem-boring beetle Phoracantha semipunctata F. (Coleoptera:
Cermabycidae). Oecologia 119, 400-407.
Hillis, W.E. (1987) Heartwood and Tree Exudates. Springer-Verlag,
Berlin, 268 pp.
Hillis, W.E. and Carle, A. (1962) The chemistry of kinos IV.
Eucalyptus hemiphloia kino. Australian Journal of Chemistry 16, 147-159.
Hillis, W.E. and Yazaki, Y. (1975) Kinos of Eucalyptus species and
their acid degradation products. Phytochemistry 13, 495-498.
Honeysett, J.L., White, D.A., Worledge, D. and Beadle, C.L. (1996)
Growth and water use of Eucalyptus globulus and E. nitens in irrigated
and rainfed plantations. Australian Forestry 59, 64-73.
Jacobs, M.R. (1986) Growth Habits of the Eucalypts. Forestry and
Timber Bureau, Canberra. 1955. Reprint. 262 pp.
Kozlowski, T.T. and Pallardy, S.G. (1997) Growth Control in Woody
Plants. Physiological Ecology Series. Academic Press, San Diego, 641 pp.
Lieutier, F., Garcia, J., Yart, A. and Romary, P. (1995) Wound
reactions of Scots pine (Pinus sylvestris L.) to attacks by Tomica
piniperda L. and Ips sexdentatus Boern. (Col., Scolytidae). Journal of
Applied Entomology 119, 591-600.
Mireku, E. and Wilkes, J. (1989) Seasonal variation in the ability
of the sapwood of Eucalyptus maculata to compartmentalize discolouration
and decay. Forest Ecology Management 28, 131-140.
Nelson, N.D. and Hillis, W.E. (1978) Genetic and biochemical
aspects of kino veins formation in Eucalyptus. II Hormonal influence on
kino formation in E. regnans. Australian Forest Research 8, 83-91.
Nicholls, J.W.P. and Pederick, L.A. (1979) Variation in some wood
characteristics of Eucalyptus nitens. Australian Forest Research 9,
Old, K.M., Murray, D.I.L., Kile, G.A., Simpson, J. and Malafant,
K.W.J. (1986) The pathology of fungi isolated from eucalypt cankers in
south-eastern Australia. Australian Forest Research 16, 21-36.
Pearce, R.B. (1996) Tansley Review No. 87. Antimicrobial defences
in the wood of living trees. New Phytologist 132, 203-233.
Robinson, R.M. (1997) Response of western larch and Douglas-fir to
infection by Armillaria ostoyae. PhD thesis, Murdoch University,
Skene, D.S. (1965) The development of kino veins in Eucalyptus
obliqua L'Herit. Australian Journal of Botany 13, 367-378.
Somervile, A. and Davies-Colley, D. (1998) Growth and utilisation
of 22 years old managed Eucalyptus regnans. Tree Grower 19, 21-23.
Tippett, J.T. (1986) Formation and fate of kino veins in Eucalyptus
L'Herit. International Association of Wood Anatomists Bulletin 7,
Tippett, J.T. and Shigo, A.L. (1981) Barrier zone formation: a
mechanism of tree defense against vascular pathogens. International
Association of Wood Anatomists Bulletin 2, 163-168.
Tippett, J.T., Shea, S.R., Hill, T.C. and Shearer, B.L. (1983)
Development of lesions caused by Phytophthora cinnamomi in the secondary
phloem of Eucalyptus marginata. Australian Journal of Botany 31,
Tippett, J.T., Hill, T.C. and Shearer, B.L. (1985) Resistance of
Eucalyptus spp. to invasion by Phytophthora cinnamomi. Australian
Journal of Botany 33, 409-418.
Van Zyl, L.M. and Wingfield, M.J. (1999) Wound response of
Eucalyptus clones after inoculation with Cryphonectria cubensis.
European Journal of Forest Pathology 29, 161-167.
Vasiliasuskas, R. (2001) Damage to trees due to forestry operations
and its pathological significance in temperate forests: a literature
review. Forestry 74, 319-336.
Wardlaw, T. (1999) Endothia gyrosa associated with severe stem
cankers on plantation grown Eucalyptus nitens in Tasmania, Australia.
European Journal of Forest Pathology 29, 199-208.
White, D.A. and Kile, G.A. (1993) Discolouration and decay from
artificial wounds in 20-year-old Eucalyptus regnans. European Journal of
Forest Pathology 23, 431-440.
White, D.A., Beadle, C.L., Worledge, D., Honeysett, J.L. and
Cherry, M.L. (1998) The influence of drought on the relationship between
leaf and conducting sapwood area in Eucalyptus globulus and Eucalyptus
nitens. Trees--Structure and Function 12, 406-414.
Wilkes, J. (1986) Host attributes affecting patterns of decay in a
regrowth eucalypt. V. Barrier zones. Holzforschung 40, 37-42.
Worledge, D., Honeysett, J.L., White, D.A., Beadle, C.L. and
Hetherington, S.J. (1998) Scheduling irrigation in plantations of
Eucalyptus globulus and E. nitens: A practical guide. Tasforests 10,
Yang, J.L. and Waugh, G. (1996a) Potential of plantation-grown
eucalypts for structural sawn products. II. Eucalyptus nitens (Dean and
Maiden) Maiden and E. regnans F.Muell. Australian Forestry 59, 99-107.
Yang, J.L. and Waugh, G. (1996b) Potential of plantation-grown
eucalypts for structural sawn products. I. Eucalyptus globulus Labil.
ssp. globulus. Australian Forestry 59, 90-98.
Alieta Eyles (1,2) and Caroline Mohammed (1,2,3)
(1) Co-operative Research Centre for Sustainable Production
Forestry, Private Bag 12, Hobart, Tasmania 7001, Australia
(2) School of Agricultural Science, University of Tasmania, Private
Bag 54, Hobart, Tasmania 7001, Australia
(3) CSIRO Forestry and Forest Products, Private Bag 12, Hobart,
Tasmania 7001, Australia