TRACHEID LENGTH - GROWTH RELATIONSHIPS OF YOUNG PINUS BRUTIA GROWN ON REFORESTATION SITES - BRILL
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IAWA Journal, Vol. 33 (1), 2012: 39– 49
Tracheid length – growth relationships of
young Pinus brutia grown on reforestation sites
Stergios Adamopoulos 1,*, Rupert Wimmer 2 and Elias Milios 3
SUMMARY
Brutia pine (Pinus brutia Ten.) reforestations have been successfully
used for decades in restoration of degraded forest ecosystems in Greece.
The future purpose of these reforestations might expand to include wood
utilisation. This study provides information on tracheid length of juvenile
brutia pine aged 14–22 years grown on good and medium sites in North-
eastern Greece. In addition, relationships among ring width, latewood
proportion, wood density, and tracheid length were evaluated by using
Causal Correlation Analysis. Similar mean tracheid length values were
found for good and medium sites. Radial variability of tracheid length
was similar on the good and medium sites, showing the typical increase
in the juvenile phase. On both site types, latewood proportion showed
a strong and positive relationship with wood density. Unexpectedly and
only on the good sites, a significant positive relationship was found be-
tween ring width and wood density. On the medium sites, tracheid length
was negatively related to fast growth and positively to high wood density.
Tracheid length on the good sites was correlated only with latewood
proportion with a weak positive relationship. The overall results may
provide opportunities to better understand the quality of small-dimension
timber of brutia pine and to better utilise it.
Key words: Pinus brutia Ten., tracheid length, site quality, radial varia-
tion, simple and partial correlation.
Introduction
One of the most serious problems in Greek forestry is the rehabilitation of severely
degraded forest land. Degradation has been going on for centuries due to unsuitable
environmental conditions, forest fires or improper forest management. Reforestation
activities started before the Second World War, mainly for aesthetic and conservation
purposes (Dafis & Hatzistathis 1984; Hatzistathis & Hatzistathis 2003).
1) Technological Education Institute of Larissa, Department of Forestry and Management of Natural
Environment, 43100 Karditsa, Greece.
2) Wood Technology and Wood-based Composites Unit, Faculty of Forest Sciences and Forest Ecol-
ogy, Georg-August-University Göttingen, Göttingen 37077, Germany.
3) Democritus University of Thrace, Department of Forestry and Management of the Environment
and Natural Resources, 68200 Nea Orestiada, Greece.
*) Corresponding author [E-mail: adamopoulos@teilar.gr].
Associate Editor: Steven Jansen
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Brutia pine (Pinus brutia Ten.) being an indigenous, drought resistant and soil
tolerant conifer species has been widely used for reforestation of depleted lands
(Dafis 1987; Hatzistathis et al. 1995). The main purpose of planting brutia pine was
to protect and improve soils, rather than grow trees for wood utilisation (Russodimos
& Petinarakis 1997). Previous research has drawn attention to the utilisation aspect of
brutia pine. The study presented selected wood characteristics of young trees (14–22
years), which originated from reforestation programmes established in Northeastern
Greece (Adamopoulos et al. 2009). Stem height variability and site quality effects were
investigated with respect to ring width, latewood proportion and wood density. Data
provided preliminary indications about the potential wood quality produced in these
reforestations. However, in order to improve our understanding of small-dimension
timber, additional information about wood properties is needed, including relationships
among selected properties. Tracheid length is another important variable mainly for
the paper industry, but also to some extent for the wood industry (Dinwoodie 1965;
Biermann 1996; Haygreen & Boyer 1996; Sirviö & Kärenlampi 1997).
Although not universally recognised, wood qualities from forests throughout the
world are changing. In many areas, forest stands tend to be harvested in shorter age
rotations leading to small diameter logs. As a result, trees contain higher proportions
of juvenile wood compared to earlier traditional harvesting (Zobel 1984; Kennedy
1995). Juvenile wood, currently widely disregarded by forest product manufactures
(Zobel & Sprague 1998), needs increased attention in terms of applicability for proc-
esses and products. Wood used by the paper industry is almost entirely juvenile; an
increasing amount of juvenile wood is also used by the wood panel industry, as well
as by solid wood processors (Wimmer et al. 2002, 2008; Downes et al. 2003). In this
context, knowledge of juvenile wood properties should be a required consideration in
processing operations of brutia pine stems from reforestation sites.
The objectives of this study are therefore 1) to characterise the variability of tracheid
length in young brutia pine trees grown on good and medium quality sites, and 2) to
explore the complex relationships between tracheid length, radial growth, latewood
proportion, and wood density. In growth rings the shortest fibres are usually found
in first-formed earlywood, while the longest fibres are laid down in the latewood at
or close to the end of a growth ring (Bisset & Dadswell 1949; Wimmer et al. 2002).
Since it is known that ring width is related to latewood proportion and wood density,
respectively, we hypothesise that tracheid length intervenes in these relationships,
which had so far not been investigated for wood coming from young brutia pine trees.
We further hypothesise that site quality differences may affect existing relationships.
Finally, special care was taken to avoid spurious conclusions from mixing cambial
growth effects and interrelationships between the investigated parameters (Larson
1969; Wimmer & Downes 2003).
MaterialS and MethodS
Overall, 16 dominant brutia pine (Pinus brutia Ten.) trees were harvested in 2005
from good and medium quality reforestation sites, all located in the central part of the
Evros Province, Northeastern Greece. The reforestations were established during the
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1980’s and early 1990’s, with seeds originating from nearby natural forest stands. Since
sampling sites were based on one seed origin, it was assumed that trees belonged to
the same genetic population. The density of planted trees varied throughout time and
the considered reforestations were not subject to thinning. Since trees were less than
23 years old the obtained wood samples are considered to be mostly juvenile.
To classify as good or as medium quality sites, a two-stage procedure was used (see
also Adamopoulos et al. 2009). As a first step, site distinction was made using site
quality surrogates such as the steepness and the exposure of the slope, shape of the ter-
rain (concave or convex) and soil depth (Dafis1986; Barnes et al. 1998; Milios 2004).
After tree harvest and age determination at breast height (1.3 m) so called “site index
curves” were used, available from reforestations established at an adjacent area north
of the study area (Hatzistathis et al. 1995). These site index curves confirmed our site
classification. In addition, stem analysis (Carmean 1972; Newberry 1991) was used
to verify that the selected dominant trees from the medium sites at age 14 had lower
tree heights than same-age trees coming from the good sites. The age of 14 years was
used since it was the age of the younger dominant tree that was cut. Good growth,
health, stem straightness and tree vigour were criteria of dominant tree selection. The
distance between the selected dominant trees and their neighbouring trees ranged be-
tween 2.2 and 2.6 m. Description of the study area as well as sample tree characteris-
tics are shown in Table 1.
Table 1. Description of the study area and characteristics of the selected dominant brutia
pine trees in good and medium sites.
Latitude 40° 59 ' – 41° 15 ' N
Longitude 26° 19 ' – 26° 36 ' E
Altitude (m) 70 – 400
Mean annual precipitation (mm) 548.5
Mean annual temperature (°C) 15
Parent material Mainly mica schist, slates, gneiss schist
Soil Sandy clay
Good sites
Number of trees used for the study 8
Height (m) 10.1 – 11.8
Age (years) 15 – 22
Stump diameter (cm) 16.55 – 23.80
Medium sites
Number of trees used for the study 8
Height (m) 7.7– 9.6
Age (years) 14 –19
Stump diameter (cm) 12.50 –22.90
The applied methods for measuring ring widths, latewood proportions and dry wood
densities followed standard procedures as described in Adamopoulos et al. (2009).
For the tracheid lengths, narrow radial strips from pith to bark were sawn from discs
taken at 0.3 m above ground. Ring width and dry wood density data measured directly
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adjacent to the tracheid lengths were already available (Adamopoulos et al. 2009). The
low sampling height (0.3 m above ground) had the risk to be associated with buttress
and reaction wood formation. Therefore, special attention was paid to avoid leaning
trees and any type of butt-swelling. For the small-diameter trees buttress influence was
also kept at a minimum. Annual ring boundaries, as well as the infrequent presence of
intra-annual density fluctuations, commonly known as “false rings”, were previously
determined (Adamopoulos et al. 2009). Each growth ring was separated into earlywood
and latewood, taking matchstick-sized subsamples. These subsamples were macerated
in glacial acetic acid and 30% hydrogen peroxide (1:1), following Tsoumis (1991).
The macerated fibres were placed onto microscope glass slides, embedded in distilled
water and covered with a glass slip. The lengths of fifty randomly selected unbroken
tracheids were measured using an Eclipse 50i light microscope, which was equipped
with a Sight DS-5M-L1 digital camera, a stand-alone control unit and image software
(all Nikon). Tracheid lengths were averaged for both earlywood (ETL) and latewood
(LTL). Latewood proportion data were used as a weighting factor, resulting in weighted
tracheid length calculated for each ring ( WTL).
Relationships between growth sites and tracheid length were related to the growth
rings of the most recent years (2001–2005). We selected a group of years as to avoid age
trends and to be able to evaluate statistical comparisons and relationships on the basis
of wood produced during the same period. All parameters were checked for normality
and found suitable for correlation analysis.
To explore relationships among wood characteristics, a procedure called Causal
Correlation Analysis (CCA) was employed. This method compares simple and partial
correlations into four cases, following Wimmer (1995), and Wimmer & Downes (2003).
Simple (Pearson) correlation (rs) is computed to describe the association between two
parameters. Simple correlation reflects both direct and indirect relationships and does
not imply causality. Simple correlation does not tell about the relationship between
two data sets if both are influenced by at least a third parameter. The effects of a third
parameter can be controlled by partial correlation (rp). This correlation describes the
linear relationship between two variables, while controlling the effects of one or more
additional variables. The number of controlled parameters gives the order of the partial
correlation coefficient (Snedecor & Cochran 1989; Weigl et al. 2007). With CCA four
cases can be produced: (Type 1) direct (causal) relationship: both correlation types are
significant with the same sign, (Type 2) indirect (non-causal, spurious) relationship:
significant simple correlation, non-significant partial correlation, (Type 3) obscured re-
lationship: non-significant simple correlation, significant partial correlation, (Type 4)
inverting relationship: both coefficients significant but sign changes. In this study the
two correlation coefficients (simple and partial) were applied to the wood variables
present in tree rings grown during a limited number of calendar years (2001 through
2005). Correlations were based on independent measurements, without possible age-
trend biases.
In addition to the Causal Correlation Analysis (CCA), we also performed ANOVA,
t-tests, and linear regressions. For all statistical analyses SPSS 17.0® software was
used.
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Table 2. Tracheid length (ETL, LTL, WTL) of brutia pine trees.
ETL
__________________ LTL
__________________ WTL
__________________
–x –x –x
s± s± s±
Good sites
Range 1.38–2.01 0.22–0.56 1.83–2.36 0.46–0.73 1.49–2.08 0.26–0.54
All trees 1.67 0.49 2.13 0.65 1.78 0.46
Medium sites
Range 1.44–2.00 0.27–0.47 1.90–2.41 0.43–0.66 1.55–2.08 0.27–0.44
All trees 1.65 0.41 2.10 0.59 1.74 0.40
ETL: earlywood tracheid length; LTL: latewood tracheid length; WTL: weighted tracheid length.
Results and discussion
Tracheid length and site conditions
Although trees had very different tracheid lengths, mean values for tracheid length
of the good (ETL 1.67 mm, LTL 2.13 mm, WTL 1.78 mm) and the medium (ETL
1.65 mm, LTL 2.10 mm, WTL 1.74 mm) sites, respectively, were more or less equal
(Table 2). The longest tracheids (3.38 mm) were measured on the good quality sites
in a 18-year-old tree. An average tracheid length of 2.76 mm was reported for a site in
Vartholomio in trees aged 23–28 years, and of 2.37 mm in 23–45-year-old trees in
Tripoli. Both sites were located in Southern Greece (Paraskevopoulou 1987). Accord-
ing to Tsoumis (1991), mature brutia pine wood has a mean tracheid length of 3.90
mm, but the age of trees was not mentioned. For mature brutia pine aged 80 years from
natural forests Papamichael (1970) reported even longer tracheid lengths, reaching
values of 4.81 mm.
For the growth rings formed during the 2001–2005 period only, tracheid lengths
(ETL, LTL, WTL) showed significant differences among trees within the sites (one-
way ANOVA, p < 0.05, Table 3). This finding is in agreement with the known large
tree-to-tree variation within a site, which usually results from environmental effects, or
existing genetic differences (McKimmy 1959). Overall, good to medium site differences
were found for ETL and LTL only (t-test, p < 0.05, Table 3). Although tracheid lengths
were slightly longer on the good sites, differences were of small magnitude with little
practical relevance. This agrees with the general observation that site quality differences
within a given geographic area might not result in major wood differences (Zobel &
Talbert 1984). The results of the present and the previous study (Adamopoulos et al.
2009) suggest that young brutia pines grow better on good sites in terms of ring width
but with little to no significant differences in latewood proportion, wood density and
tracheid length. Several suggestions have been made regarding effects of the growth
site on wood formation and properties. Site quality is seen as difficult to be related to
wood properties due to interactions with many other factors, apart from large between
tree differences (Zobel & Van Buijtenen 1989). No information was found about effects
of site on tracheid length for brutia pine. Site quality had an important impact on the
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Table 3. Tracheid length (ETL, LTL, WTL) of the last five annual rings (growth period
2001–2005) in brutia pine trees.
ETL
__________________ LTL
__________________ WTL
__________________
–x –x –x
s± s± s±
Good sites
Range 1.61–2.54 0.12–0.44 2.37–3.08 0.31–0.55 1.83–2.67 0.10–0.20
F 156.427* 58.852* 23.991*
All trees 2.09 0.29 2.77 0.49 2.24 0.35
Medium sites
Range 1.68–2.45 0.19–0.42 2.33–3.04 0.31–0.42 1.86–2.58 0.08–0.20
F 82.705* 66.868* 16.689*
All trees 1.99 0.36 2.64 0.45 2.13 0.25
t 4.317* 6.262* 1.577 ns
ETL: earlywood tracheid length; LTL: latewood tracheid length; WTL: weighted tracheid length.
Note: F values for statistical comparison of means among trees in each site type (anova) and
t values for comparison of means between sites (t-test).
* = differences statistically significant at p < 0.05; ns = differences not statistically significant.
physical and mechanical properties of naturally grown brutia pine (Bektas et al. 2003;
Gundogan et al. 2005). The large tree-to-tree variation and the expressed juvenility
allowed only conclusions with relevance to young brutia pine trees.
Tracheid length radial profiles
The pattern of tracheid length variation relative to cambium age was similar for all
trees, showing a rapid increase during the first 14–22 years of radial growth. The rela-
tionship between tracheid length (ETL, LTL, WTL) and cambium age was modelled
by linear regressions (Table 4). The mean tracheid length followed the general age
trend for conifers (Fig. 1; Anderson 1951; Dinwoodie 1961; Tsoumis 1991). The radial
variability of tracheid length was similar for the investigated site qualities, showing a
Table 4. Coefficients of linear regression formulas (Y = AX + B) for the relationships between
tracheid length (Y: ETL, LTL, WTL) and cambium age (X) in good and medium sites.
Y A B R2 F value
Good sites
ETL 0.057 1.095 0.45 106.2*
LTL 0.100 1.107 0.78 459.0*
WTL 0.069 1.080 0.56 165.2*
Medium sites
ETL 0.068 0.963 0.75 355.8*
LTL 0.106 1.030 0.84 617.7*
WTL 0.079 0.946 0.81 484.0*
ETL: earlywood tracheid length; LTL: latewood tracheid length; WTL: weighted tracheid length.
R2 = coefficient of determination; * = significant at p< 0.05.
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3.0 Good sites
2.5
Tracheid length (mm)
2.0
ETL
1.5 LTL
WTL
1.0
0.5
0 5 10 15 20
Age
3.0 Medium sites
2.5
Tracheid length (mm)
2.0
ETL
1.5 LTL
WTL Figure 1. Radial variation of mean tracheid
1.0 length (ETL, LTL, WTL) in good and medium
sites. ETL: earlywood tracheid length, LTL:
0.5 latewood tracheid length, WTL: weighted
0 5 10 15 20 tracheid length.
Age
continuous increase for mean ETL, LTL and WTL. Paraskevopoulou (1987) reported
a continuous increase in tracheid length for brutia pine trees aged 23 years, grown on
reforestation sites in Southern Greece. Since no other reports exist for this species,
older trees need to be investigated to complete the picture on pith-to-bark patterns for
this species.
Tracheid length relationships
There has always been considerable interest in the relationships between ring width,
wood density and tracheid length. For softwoods, it is usually assumed that the wider
the growth ring, the lower the wood density, and the shorter the tracheids (Dinwoodie
1965; Tsoumis 1991; DeBell 1994; Dutilluel et al. 1998; Fujiwara & Yang 2000;
Mäkinen et al. 2000). However, controversial results have been reported in literature
(Echols 1955; Zobel & Van Buijtenen 1989; Zhang 1995; Koga & Zhang 2002; Lin &
Chiu 2007). Possible explanations as described by Dutilluel et al. (1998) or Wimmer
& Downes (2003) include tree age effects, environmental influences, or management
practices.
For both types of sites, a strong positive direct relationship (CCA Type 1) was found
between wood density and latewood proportion (Fig. 2). Wimmer and Downes (2003)
found even stronger correlations (rs = 0.83, rp = 0.64) for Norway spruce. The good
quality plots exhibited a strong positive relationship between ring width and density
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Good sites Medium sites
rs = 0.66** rs = 0.62**
rp = 0.68** rp = 0.64**
Latewood Wood Latewood Wood
% density % density
rs = 0.31** rs = 0.21*
rp = 0.31** rp = 0.31*
Tracheid Tracheid
rs = ns length rs = 0.55** rs = ns
length rs = ns
rp = -0.26* rp = 0.57** rp = -0.28* rp = 0.30*
rs = -053**
rp = -0.57**
Ring Ring
width width
Figure 2. Relationships among weighted tracheid length (WTL), ring width, latewood propor-
tion and wood density calculated across tree rings of growth period 2001–2005 in good and
medium sites. Thick solid lines: strong significant simple (rs) and partial (rp) correlations
(strong direct relationship); light solid lines: weak significant rs and rp correlations (weak direct
relationship); dotted lines: non-significant rs correlation but significant rp (hidden relationship).
* = p < 0.05, ** = p < 0.01.
(rs = 0.55, rp = 0.57), and a direct positive relationship also existed between tracheid
length and latewood proportion, though weaker (rs = 0.31, rp = 0.31, Fig. 2). All Type 1
correlations are seen as causal in the first place. A weak but significant negative par-
tial correlation was found between latewood proportion and ring width. Since the
simple correlation between these two parameters was not significant, a CCA type 3 is
present. The significant partial correlation is interpreted as being obscured, and it might
be explained by the strong and positive linkages between ring width, and wood den-
sity and latewood proportion. These positive correlations superimposed the negative
correlation between ring width and latewood proportion, as revealed through partial
correlation.
A difference between the two site types was seen in the relationship between ring
width and tracheid length. No relationship existed for the good quality site trees. The
medium quality trees, however, showed a Type 1 negative relationship between ring
width and tracheid length (Fig. 2). This strong linkage seems to be the reason why no
significant simple correlation existed between wood density and ring width for the
medium quality trees. It is worth to mention that according to Guller (2007), there was
no clear evidence showing an effect of increased radial growths caused by thinning
treatments on fibre length of planted brutia pine.
The most unexpected result, however, was the significant positive relationship be-
tween ring width and wood density for the good quality plot trees (Type 1 for the good
sites, Type 3 for the medium sites; Fig. 2). For the ring width-wood density relationship
in conifers, literature mostly reports negative but weak relationships (Dutilleul et al.
1998; Wimmer & Downes 2003), while others found no significant relationships at
all (e.g. Petrik 1968). Dutilleul et al. (1998) reported that a negative ring width-wood
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density relationship found in slow-grown Norway spruce trees was absent in fast grown
trees. Increase of annual ring width after thinning of brutia pine plantations did not
affect wood density due to the insignificant effect of thinning on latewood percentage
(Guller 2007). However, our findings suggest that fast growth of brutia pine leads to
higher wood density. On sites with moderate growth and medium site quality, growth
rates seem to have a close link to tracheid length. On these sites fast growth was
linked to short tracheids (Type 1). It should be noted that the relationships found are
independent of cambial age. There is an inherent increase in latewood proportion with
cambial age that is related to the changing development and structure of the crown.
Larson (1969), who considered crown development as the major factor, demonstrated
that with increasing age, tree canopies become more closed and the lower bole of the
tree produces more latewood. This increase in latewood proportion runs parallel with
higher density and declining ring width, mimicking a negative relationship between
ring width and wood density.
The commercial value of brutia pine, besides being important as a nature conserva-
tion species, has been undervalued. Causal correlation analysis was employed here
for the first time to understand better direct and indirect relationships among wood
characteristics of juvenile wood of brutia pine. The obtained findings should help forest
growers as well as the wood industry to understand better the intrinsic relationship of
wood properties in this species and suggest improved utilisation strategies for brutia
pine grown on reforestation areas.
Conclusions
In young brutia pine, the effect of site on tracheid length was of little practical impor-
tance. On both sites, latewood proportion had a strong positive and direct effect on
wood density. Faster growth of brutia pine resulted in higher wood density, but only
on the good sites. Tracheids were influenced only on the medium sites, positively by
wood density and negatively by ring width. Since results are limited to young age trees,
future research should focus on mature brutia pine trees and reconsider the relationships
by comparing juvenile and mature wood trends.
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