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Thinning intensity and growth of


Norway spruce stands in Finland
HARRI MKINEN* AND ANTTI ISOMKI
Finnish Forest Research Institute, PO Box 18, FIN-01301 Vantaa, Finland
*Corresponding author. E-mail: harri.makinen@metla.fi

The effects of thinning intensity on the growth and yield of Norway spruce (Picea abies (L.) Karst.)
were investigated in long-term thinning experiments on mineral soil sites in southern Finland. The
measurement period was on average 27 years, and the intensity of the thinnings from below ranged
from heavy thinning (45 per cent of basal area removed) to no thinning. Total stem volume
increment and merchantable volume produced per hectare were the highest on the unthinned plots,
but light and moderate thinning (<30 per cent removed) produced almost as much. Heavy thinning
(>30 per cent removed) reduced the volume increment by about 10 per cent. However, a part of the
total production of unthinned plots was lost through natural mortality. On the thinned plots, natural
mortality was considerably lower compared with the unthinned plots. The average diameter
increment of all the trees, as well as the diameter of the largest trees, clearly increased with
increasing thinning intensity. In contrast, dominant height increment was not affected by thinning.
The stand age at the time of establishment of the experiments had no major effect on the growth
reactions after thinning. Thus, heavy thinning results in earlier thinning yields and a higher
proportion of larger-sized stems at the expense of a somewhat lower total yield.

Introduction
Control of stand density by thinning has been the
major tool in regulating tree growth and improving timber quality. While thinning from below
may increase the merchantable volume of a
stand, usually it does not increase the total
volume increment per unit area (e.g. Assmann,
1954; Carbonnier, 1967; Hasenauer et al., 1997;
Zeide, 2001). Several studies have shown that
volume increment of many tree species does not
decline with decreasing stand density within a
wide range of stand density (e.g. Hamilton, 1981;
Horne et al., 1986). This indicates that thinning
Institute of Chartered Foresters, 2004

from below redistributes the increment from


smaller trees to larger ones, and a smaller number
of trees is able to produce the same volume increment per unit area. In order to increase the merchantable volume, or not to decrease the total
volume increment, the intensity, timing and frequency of thinnings have to be defined.
In Finland, Norway spruce (Picea abies (L.)
Karst.) is economically and ecologically one of
the most important tree species. The first growth
and yield tables for Norway spruce were published by Blomqvist in 1872 (see also Heikkil,
1914), but they were not widely applied. Systematic growth and yield studies began in the 1910s
Forestry, Vol. 77, No. 4, 2004

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Summary

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the establishment of experiments was continued


and the first results on Norway spruce were published on the basis of five experiments (Vuokila,
1975). In later publications, Vuokila (1980,
1985) described the further development of the
same experiments as in the first paper (see
Table 1; numbers Nyn1Nyn5). Mielikinen
(1978) demonstrated the effects of thinning on
the diameter increment of Norway spruce during
a 7-year period in one experiment (Table 1;
Nyn1). In addition, the results of the experiments
have been presented in seminars and excursions
(e.g. Isomki, 1993, 1995). However, no systematic presentation of the whole material exists.
In the other Fennoscandian countries, recent
results from Norway (Braastad and Tveite, 2000,
2001) and Sweden (Eriksson and Karlsson, 1997)
have shown that the volume increment in
Norway spruce stands stays constant or decreases
only slightly with increasing thinning intensity
over a wide range of stand density. Earlier
findings from central Europe by Assmann (1954),
Hamilton (1976), Schober (1979, 1980) and
Kramer and Jnemann (1985) have also shown
that the volume increment of Norway spruce
does not decrease proportionally with decreasing
stand density. However, the benefits and
disadvantages of thinnings are still the subject of
heated debate (Braastad, 2001; Elfving et al.,
2001).
The objective of this study was to relate
thinning intensity with diameter, height and
volume increment on the basis of permanent
long-term experiments with thinnings from
below in planted Norway spruce stands. This
study is a sequel to the reports of Vuokila (1975,
1980, 1985), which were based on the five
experiments also used in this study. In this study,
we expanded the database and used all the
thinning experiments of the HARKAS series,
established by the Finnish Forest Research Institute in Norway spruce stands (21 experiments).
Only two stands which were fertilized before the
establishment of the experiments were excluded
from the data set. Many experiments have
already been studied for over 30 years (Table 1)
and they approach maturity or are already
mature. This gives us an opportunity to investigate total stem volume production and thinning
removal, as well as stand structure, during the
whole stand rotation.

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and the first results for Norway spruce were published by Ilvessalo (1920). They described the
structure and development of fully stocked,
naturally regenerated pure Norway spruce
stands. A decade later, Cajander (1933) studied
the development of planted, unthinned Norway
spruce stands.
After World War II, selective thinnings, i.e.
removing only the largest trees, were prohibited
and thinnings from below were promoted (Anon.,
1948; Kalela, 1948). At the same time, a research
project was initiated to study the effects of thinnings on the structure and development of
naturally regenerated Norway spruce stands
(Vuokila, 1956; Kallio, 1957). However, the
intensity of the thinnings was low and the
removed trees were heavily suppressed or dying.
In the early 1960s, the thinning intensity in
practical forestry increased mainly due to the
mechanization of harvesting operations. Because
the short- and long-term growth effects of heavy
thinnings were not known, a new research project
was established and growth and yield tables, as
well as thinning guidelines, were prepared for
planted Norway spruce stands (Vuokila and
Vliaho, 1980). During the last decades, growth
models have been developed for forest management planning (e.g. Hynynen et al., 2002).
All the Finnish studies cited above were based
on temporary sample plots (apart from the recent
growth models that were based on remeasured
inventory growth plots) because there was an
urgent demand for growth and yield tables, but
no data available. However, permanent sample
plots provide more reliable results on long-term
growth and stand dynamics. In Finland, the first
thinning experiments were established in the
1920s and 1930s (Ilvessalo, 1932). Some of the
results of these experiments have been published
(Vuokila, 1960, 1977), but the experiments have,
in the main, not fulfilled their expectations due to
natural damage, changes in measurement
methods, subjective classification of many parameters, experimental design, etc. (Vuokila,
1965). In addition, the thinnings applied have
become out-of-date.
In the 1960s, new thinning experiments on
Norway spruce, Scots pine and silver birch (the
so-called HARKAS series) involving more intensive treatments and a better experimental design
were established (Vuokila, 1983). In the 1970s,

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T H I N N I N G I N T E N S I T Y A N D G R O W T H O F N O RWAY S P R U C E

Materials and methods


The experiments

different stages of stand development. The


material consisted of two separate sets: (1) thinning experiments based on stem number (13
experiments); and (2) thinning experiments based
on stand basal area (eight experiments). The
experiments based on stem number were
originally planned to include three different treatments with varying thinning intensity and a
number of repeated thinnings from below, as well
as an unthinned control plot. The final stem
number per hectare was planned to be the same
in all the treatments, and it was to be reached
using two heavy, three moderate or five light thinnings from below, based on a fixed number of
stems remaining after the thinnings. However, it
was soon noticed that, in many cases, the treatments resulted in only small differences among
the plots and the research plan was therefore
modified. The intensity of the later thinnings was
increased in order to maintain the differences in
stand density between the plots. In the experiments based on basal area, a constant basal area
ratio compared with the unthinned control plot
was maintained on the thinned plots, i.e. the plots
were thinned to basal areas of 90, 75 and 60 per
cent compared with the control plot.
Square or rectangular plots surrounded by a
5-m-wide buffer zone were established in each

Figure 1. Location of the experiments. For explanation of the onset stages, see Materials and methods.

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The material was collected from 21 thinning


experiments in southern Finland (Figure 1). The
experiments were established by the Finnish
Forest Research Institute in the 1960s and early
1970s, apart from three experiments which were
established in the late 1970s and early 1980s
(Table 1). The stands were even-aged, pure or
almost pure Norway spruce (Picea abies (L.)
Karst.) stands located on mineral soil, and were
established by planting with seed of local origin.
The sites were classified as the OxalisMyrtillus
or Myrtillus forest site type (Cajander, 1909),
which corresponds to highly fertile or fertile sites
typical for Norway spruce. The site index (H100,
dominant height at age 100 years in metres)
ranged from 29 m to 36 m. Site indices were
calculated using the functions of Vuokila and
Vliaho (1980). Due to changes in land use (e.g.
the construction of houses and roads), some of
the experiments had to be terminated earlier than
planned (Table 1).
The principal aim of the experiments was to
investigate the effects of thinning intensity on
growth and yield of Norway spruce stands at

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Table 1: Characteristics of the experiments at establishment


At establishment

No.

H100*

6
3
10
4
8
10
8
8
4
5
8
12
3
5
4
4
12
4
8
4
4

34.4
31.6
28.9
31.1
30.2
32.4
32.9
32.7
32.9
30.0
31.0
34.0
36.4
30.2
33.0
33.0
30.0
34.5
34.7
33.0
33.0

1918
1925
1931
1931
1914
1916
1932
1918
1936
1934
1934
1955
1930
1938
1934
1924
1922
1931
1926
1925
1930

1970
1970
1971
1973
1970
1970
1970
1971
1971
1973
1977
1981
1979
1965
1964
1964
1961
1962
1962
1962
1962

1998
1998
1994
1998
1999
1998
1998
1998
1985
1992
2001
1994
1994
1998
1999
1999
1998
1998
1999
1988
1992

3
2
1
2
3
3
2
3
2
2
2
1
3
1
1
2
1
1
1
1
1

4
5
5
5
5
6
6
5
3
4
5
4
5
6
6
6
8
8
8
6
7

Hdom
(m)

No. of
stems ha1

24.3
16.9
13.5
17.0
22.4
23.4
18.1
23.1
16.3
18.0
16.9
12.5
24.1
9.8
12.1
17.4
14.2
14.7
16.0
15.6
14.3

1072
1404
2212
1856
1114
1042
2013
935
2932
1526
2137
2973
1626
3335
1689
1168
2055
3247
2394
2295
3402

* H100 site index, dominant height at age 100 years (m).


1, early; 2, medium; and 3, late onset of the treatment based on the H
dom/H100 ratio, see Materials and
methods.
Dominant height based on 100 largest trees ha1.
Before the onset of the treatments.

stand. The average area of the plots was


10001600 m2 (range 5002500 m2). Most of
the experiments had four, eight or 12 plots, i.e.
the four treatments were replicated in a randomized block design in the experiments with eight or
12 plots (Table 1). In the experiments with five,
six or 10 plots, one or two of the treatments were
additionally replicated within the experiments or
blocks. In experiments Vh002 and Vh098,
having only three plots, the light thinning intensity was missing (see below).
Measurements and statistical analysis
Following establishment, the experiments were
measured between two and seven times (Table 1).
The measurement period was on average 27
years. At the time of the last measurement, stand

age ranged from 39 to 85 years. Tree species, stem


diameter at breast height (d1.3), and possible
damage (fallen, broken, decaying, needle loss,
etc.), as well as its cause (wind, snow, competition, insect or fungus species) and severity
(temporary, causing permanent defects, lethal),
were recorded for each tree on the plot. In the
selection of sample trees, the probability for a tree
to be selected was proportional to its diameter,
but the sample trees were randomly located on
the sample plots. Height, height to crown base,
and stem diameter at 6 m height (d6.0) were
measured on each sample tree (on average, 54 per
plot). The crown base was defined as the lowest
whorl with at least one living branch that was
separated from the other living whorls above it by
no more than one dead whorl.
Stand characteristics for individual plots were

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Vh001
Vh002
Vh005
Vh009
Vh011
Vh012
Vh013
Vh014
Vh017
Vh040
Vh048
Vh097
Vh098
Ha001
Pu041
Pu042
Nyn1
Nyn2
Nyn3
Nyn4
Nyn5

No. of
plots

Year
of
Establishment
Last
Treatment
No. of
planting
of plots
measured onset measurements

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BA

n -1 T

i =1

i * _ BAr i

+ BAt i + 1i /2
T tot

(1)

where BAr is the basal area of the remaining


living trees at the beginning of each measurement
period, BAt the total basal area at the end of each
measurement period including dead trees and
trees to be thinned, Ti the length of each measurement period (years), Ttot the length of the total
measurement period (Ti), and n the number of
individual measurement periods. The plots were
classified as follows: (U) unthinned (average basal
area 95 per cent of that on the control plots)
note that some plots with light thinning intensity
were classified as unthinned because their basal
area exceeded 95 per cent; (L) light thinning
(8594 per cent); (M) moderate thinning (70
84 per cent); and (H) heavy thinning (70 per
cent). In class H, the minimum and average
relative basal areas were 55 per cent and 63 per
cent, respectively.
Based on the ratio between dominant height
(based on 100 by diameter of largest trees ha1)
at the time of establishment and the site index
(Hdom/H100), the experiments were further
divided into three thinning onset stages as
follows: early onset (Hdom/H100 0.49), medium
onset (0.50 Hdom/H100 0.60) and late onset
(Hdom/H100 0.61). The mean Hdom/H100 ratio
was 0.42, 0.54 and 0.71 for the early, medium
and late onset stages, respectively. At the early
onset stage, only light pre-commercial thinnings
were carried out before establishment of the
experiments. The stands of medium onset stage
had already passed the first commercial thinning
phase, but the thinnings carried out had been
light. Accordingly, only light thinnings were
carried out in the stands of late onset stage before
establishment.
Statistical significance of the differences among
the treatments was analysed using covariance
analysis including random experiment and block
effects. The model used to test the treatment
effects was:
Y = + D + Xsbp + us + usb + usbp

(2)

where Ysbp is a dependent variable, is the


overall mean, D the effect of stand density class,
the regression coefficient, and us, usb and usbp
the random effects for stand s, block b and plot
p. The initial differences among the plots were
removed by applying a continuous covariate
(Xsbp) measured before the onset of the treatment,
i.e. mean stem diameter, dominant height (100 by

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calculated using the KPL software developed at


the Finnish Forest Research Institute (Heinonen,
1994). Stem volumes of the sample trees were
calculated using volume functions based on the
measured stem diameters (d1.3, d6.0) and tree
height (Laasasenaho, 1982). The heights of the
other trees were predicted using Nslunds height
curve (Nslund, 1937) that was fitted for each plot
with the help of the tree heights measured on the
sample trees. The volume of the other trees was
calculated using smoothing functions fitted to the
sample tree data. The minimum length applied for
pulpwood boles was 3.0 m and the minimum top
diameter was 8.0 cm over bark. The stem wood
below this size was considered as wastewood. The
minimum length applied for timber logs was
3.7 m, and the minimum top diameter over bark
was 19.5 cm. The minimum top diameter progressively decreased with increasing log length,
being 16.0 cm when the log length was over 4.3 m.
Annual increments were calculated as the
difference between successive measurements
divided by the number of years between the
measurements. The periodic growth between
the measurements was corrected to correspond to
the average climatic conditions using annual
radial growth indices. The indices for Norway
spruce in southern Finland were based on the
increment cores measured in connection with the
National Forest Inventory. The indices published
by Henttonen (1986) were used for the period
19611979, and the indices by Henttonen
(unpublished data) for the period 19801999. As
no indices were available for the years 2000 and
2001, they were assumed to be average years.
The method for calculating the growth indices is
described in Henttonen (1986, 1990).
Because the number and intensity of the thinnings varied among the plots and experiments,
the plots could not be classified into different
groups in accordance with the original research
plan. Therefore, the plots were grouped on the
basis of the average basal area on the plot during
the whole measurement period _ BAi compared
with that on the control plots of each experiment.
The basal areas of the different measurement
periods were weighted by the period length, i.e.

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Figure 2. Mean annual diameter increment of all the trees (parts ac) and the 400 by diameter largest trees
ha1 (parts df) on plots of different thinning intensity (U, unthinned; L, light; M, moderate; H, heavy
thinning) and treatment onset stage (early, parts a and d; medium, parts b and e; late onset, parts c and f).
Treatments marked with the same letter are not significantly different (P 0.05). Stem diameter before the
onset of the treatment was used as the covariate in equation 2.

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355

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diameter of largest trees ha1) or stand volume.


The values shown in the figures and tables are
adjusted by setting covariate effects to their mean
values.
The pairwise comparisons were performed by
computing generalized least-square means of the
treatment effects. Owing to the unbalanced
design, adjusted P-values for the multiple comparison were computed from the simulated distribution of a multivariate random vector. In the
figures and tables, the treatments marked with
the same letter are not significantly different (P
0.05). Restricted maximum likelihood (REML)
estimation in the MIXED procedure of SAS (SAS
Institute, 1999) was used in the analysis.

Results
The arithmetic mean annual diameter increment of
all the trees on a plot increased with decreasing
stand density (Figure 2ac). The differences
between the thinning intensities were statistically
significant at all treatment onset stages. Because
the differences of the mean increment may be
caused by different numbers of stems, mean
diameter increment was also calculated for the
400 (by diameter) largest trees ha1. The differences between the treatments in the diameter increment of the largest trees were, however, similar to
those for all the trees (Figure 2df). On the other
hand, thinning intensity had no clear effect on
annual dominant height increment (Figure 3).
On plots with light and moderate thinning
intensity, the annual volume increment per hectare
was about the same as on the unthinned plots at
all treatment onset stages (Figure 4). Compared
with the unthinned plots, a heavy thinning intensity decreased the volume increment per hectare
during the whole measurement period by 8 per
cent at the early onset stage. At the medium onset
stage, a heavy thinning intensity decreased the
volume increment even more (11 per cent), but the
differences between the treatments were not
statistically significant. At the late onset stage, the
volume increments among the individual plots
were very variable and, therefore, the differences
between the thinning intensities were not statistically significant.
In addition to the analysis according to fixed
treatment onset stages, the whole material was

Figure 3. Mean annual dominant height increment


on plots of different thinning intensity and treatment onset stage. Dominant height before the onset
of the treatment was used as the covariate in
equation 2. For explanation of the symbols, see
Figure 2.

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Figure 4. Mean annual volume increment on plots


of different thinning intensity and treatment onset
stage. Total volume before the onset of the treatment was used as the covariate in equation 2. For
explanation of the symbols, see Figure 2.

also combined, i.e. the volume increments


between the successive measurements were
analysed according to the dominant height in the
middle of each measurement period (Figure 5).
The unthinned plots and heavy thinning intensity
differed from each other at all dominant height
stages. The annual volume increment of the light
and moderate thinning intensity lay between
those of the unthinned and heavy thinning plots,
excluding the lowest and highest dominant height
stages. However, the mutual order of the light
and moderate intensities was variable at different
dominant heights and it was not possible to distinguish between them.
Volume increments and basal areas on the
thinned plots during the whole measurement
period were also examined in relation to the
mean volume increment and basal area of the
unthinned plots in each experiment (Figure 6).
Because the experiments had several unthinned
plots, and their basal areas were also related to
the mean value, some relative basal areas
exceeded 100 per cent. Figure 6 clearly shows
that the variation of relative volume increment
was high among individual plots, but that the
mean relative volume increment decreased only
slightly with decreasing mean relative basal area.
As expected, the current stem volume per
hectare decreased with increasing thinning
intensity at all treatment onset stages (Table 2).
However, total yield, calculated by summing up
the current volumes of living trees and the
volumes of thinned and dead trees, did not differ
significantly between the unthinned plot and the
light and moderate thinning intensities (Table 2).
The heavy thinning intensity decreased the total
yield by 7 per cent, 4 per cent and 7 per cent
compared with the unthinned plots at the early,
medium and late onset stages, respectively, but
the differences at the medium and late onset
stages were not statistically significant. On the
unthinned plots, some of the trees had also been
removed in connection with the thinnings
(Table 2), because some of the thinned plots were
placed in the unthinned group if their basal area
did not differ from the control plots >5 per cent.
In addition, light sanitary thinnings were carried
out in some experiments, i.e. dying trees were
removed at the request of the landowner.
The current log volume was highest with the
unthinned or light thinning intensity and lowest

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13

10

7
15

U
L
M
H
17

19

21

23

25

27

29

31

Dominant height (m)


Figure 5. Mean annual volume increment between the successive measurements plotted against mean
dominant height during each measurement period on plots of different thinning intensity. For explanation
of the symbols, see Figure 2.

Figure 6. Volume increment on plot i during the whole measurement period in relation to the mean volume
increment on the unthinned plots of the same experiment (ivi/ivu) plotted against the relative average basal
area during the whole measurement period (related to that of the unthinned plots, BAi/BAu, see equation 1).
The thick continuous line is a non-linear curve fitted to the data [ivi/ivu = (BAi/BAu)2/{(0.08 + 0.91BAi/
BAu)2}, R2 = 0.16].

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Volume increment (m2 ha1 a1)

16

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Table 2: Total yield and its structure on plots of different thinning intensity (U, unthinned; L, light; M,
moderate; H, heavy thinning) and treatment onset stages
Early onset
U

Late onset

374a
172b
29a
575b

377a
140c
28a
545b

365a
182a
20a
567a

408a
161a
20a
589a

396a
145a
17a
558a

374a
150a
18a
542a

560a
109a
10a
679a

556a
99a
9a
664a

552a
101a
9a
662a

541a
84a
7a
632a

339ab
94b
8b
441b

314b
45c
4c
363c

349a
145a
12a
506a

354a
103a
8ab
465ab

335a
50b
5b
390bc

286a
46b
4b
336c

526a
80a
7a
613a

480ab
55ab
5ab
540ab

442b
42b
4ab
488b

419b
25b
3b
447b

25a
72bc
20bc
117c

56b
93c
24c
173d

2a
4a
1a
7a

38ab
44ab
6ab
88ab

44ab
90bc
12b
146b

65b
101c
14b
180b

8a
3a
1a
12a

46ab
28b
3a
77ab

90b
52c
5b
147b

108b
62c
5b
175b

8a
5b
1b
14b

7a
2b
1b
10b

10a
35a
7a
52a

15a
19ab
6a
40a

14a
7b
1b
22a

12a
8b
1b
21a

28a
20a
2a
50a

29a
14ab
1ab
44ab

17a
6bc
1ab
24b

13a
0c
0b
13b

Total volume before the onset of the treatment was used as the covariate in equation 2.
Treatments marked with the same letter (in the same line and within the onset stages) are not significantly
different (P 0.05).

on the heavy thinned plots, even though the


differences at the medium onset stage were not
statistically significant (Table 2). Accordingly, the
current volume of pulpwood and wastewood
decreased with increasing thinning intensity at all
treatment onset stages. In contrast, total volume
of merchantable-sized timber, i.e. the current
volume of logs and pulpwood summed up with
that removed in thinnings, was about the same in
all thinning intensities.
On the unthinned plots, volume of dead trees
was higher than that on the thinned plots,
especially in the later phases of the experiments
(Figure 7). On the thinned plots, the differences
in dead wood volume were small among the
thinning intensities (Figure 7 and Table 2). At the
late onset stage, dead wood volume was,
however, about the same on the unthinned and
light thinned plots. The initiation of the treatment did not result in a dramatic change in mortality. Instead, the mortality rate increased

slightly in the later phases of the measurement


period. On the other hand, the volume of logsized dead trees was relatively similar in all
thinning treatments, i.e. the higher dead wood
volume on the unthinned plots was mainly
caused by the mortality of small-sized trees
(Table 2).

Discussion
Total volume production per unit area was
highest on the unthinned or lightly thinned plots,
irrespective of the treatment onset stage.
However, light and moderate thinning had
almost no effect on volume increment, or total
volume produced during the long measurement
period. Only heavy thinning resulted in a clear
reduction in volume increment. Even though the
basal area of the heaviest treatments was, on
average, 5570 per cent below that on the

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Total volume (m3 ha1)


Log
349a 384a
Pulp
206a 178b
Wastewood
29a
27a
Sum
584a 589b
Current volume (m3 ha1)
Log
331ab 370a
Pulp
165a 119b
Wastewood
15a
9b
Sum
511a 498a
Thinned volume (m3 ha1)
Log
7a
7a
Pulp
13a
51b
Wastewood
5a
16b
Sum
25a
74b
Dead volume (m3 ha1)
Log
8a
4a
Pulp
25a
7b
Wastewood
9a
1b
Sum
42a
12b

Medium onset

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359

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Figure 7. Cumulative volume of dead trees on plots of different thinning intensity: (a) early; (b) medium;
(c) late onset of the treatment. For explanation of the symbols, see Figure 2.

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lated by the nutrients released from logging


residue. On the other hand, reduced height
growth on Norway spruce has also been reported
after heavy thinning (Abetz, 1976; Abetz and
Unfried, 1984).
In this study, the sites of the experiments were
fertile and this may contribute to the fast growth
of the remaining trees after the release. On less
fertile sites, the recovery of the remaining trees
may take a longer time and heavy thinnings
probably result in larger growth reductions
(Mkinen and Isomki, 2004a). On the other
hand, the treatment onset stage had no major
effect on the recovery after the thinning, i.e.
slower growing older trees could utilize the free
growing space as fast as the trees in younger
stands. Even though the medium and late onset
stands were dense at the time when the experiments were established, they were not overstocked and the tree crowns were vigorous
(Mkinen and Isomki, 2004b). Therefore, they
were able to rapidly respond to the treatments.
The experiments used in this study were
originally planned to study the effects on growth
and yield of heavy and seldomly repeated thinnings (Vuokila, 1983). Even though the heaviest
thinnings were considered exceptionally intensive
at the time of establishment, the results showed
that heavier treatments are needed to find out
thinning intensities that clearly reduce the total
stem volume production. In Norway spruce
stands growing on fertile soils, removing even
50 per cent of the growing stock has not resulted
in marked reductions in volume increment
(Mller, 1954; Carbonnier, 1967; Abetz, 1976;
Eriksson and Karlsson, 1997). New thinning
experiments that cover the whole stand density
range from free-growing trees to unthinned
stands are needed in order to define the complete
relationship between stand density and volume
production.
As was the case for total volume production,
merchantable volume produced up to the last
measurement was highest on the unthinned and
light thinned plots. Only the heavy thinning
resulted in a reduced merchantable volume. High
stand density may, however, keep tree size under
the merchantable limit. Therefore, after a certain
stand density level, increasing the stand density
may also decrease the total merchantable volume
(e.g. Spellmann and Nagel, 1996; Zeide, 2001).

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unthinned plots, the volume production was only


reduced by 47 per cent. The results of this study
are similar to the results reported in thinning
experiments of Norway spruce in other
Fennoscandian countries (Mller, 1954; Carbonnier, 1957, 1974; Bryndum, 1967, 1969;
Eriksson, 1987; Pettersson, 1993; Eriksson and
Karlsson, 1997; Braastad and Tveite, 2000,
2001) and in central Europe (Assmann, 1954;
Hamilton, 1976; Schober, 1979, 1980; Kramer
and Jnemann, 1985; Spellmann, 1986). In
addition, according to the growth and yield
tables for Norway spruce developed in Norway
(Braastad, 1975), Sweden (Eriksson, 1976),
Finland (Vuokila and Vliaho, 1980) and
Germany (Assmann and Franz, 1965), total
volume production during a stand rotation is not
closely related to the thinning intensity. The
volume of the trees possibly removed in precommercial thinnings and commercial thinnings
carried out before the study was not known.
However, the trees removed have no effect on the
differences among the treatments.
The results clearly demonstrate that the
remaining trees can rapidly occupy the growing
space released by the removed trees. The average
diameter increment of all trees, as well as that of
the largest trees clearly increased (cf. Hamilton,
1976; Abetz and Unfried, 1984; Kramer and
Jnemann, 1985; Eriksson, 1987). According to
Braastad and Eikeland (1986) and Braastad and
Tveite (2001), thinning either did not or only
slightly accelerated the growth of the largest
trees. Their results were based on delayed
and light thinnings and, most probably, the differences in thinning intensity explain these
differences.
The treatments had no effect on dominant
height increment. On the heavily thinned plots,
height increment was slightly faster than that
with the other treatments. Thus, the results of
this study are in accordance with most studies on
Norway spruce, i.e. the height increment of
dominant trees is not affected by stand density
(cf. Bryndum, 1967; Kramer and Jnemann,
1985; Eriksson, 1987; Handler, 1990; Eriksson
and Karlsson, 1997). According to Mller
(1954), Bryndum (1969) and Hamilton (1976),
height increment may, however, increase with
increasing thinning intensity. Those authors have
speculated that height growth could be stimu-

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because all the treatments were included in


almost all of the experiments.
In those experiments where thinnings were
based on the number of stems per hectare, it was
soon noticed that the original research plan
resulted in a rather similar basal area per hectare,
even though the differences in stem number were
high. Thus, the original plan was modified and
the thinnings were intensified. Furthermore, in
some experiments the original schedule could not
be completely followed because of high measurement costs, low thinning removals that were not
economically viable for the land owner, etc. The
same average basal areas were classified into the
same group regardless of the thinning schedule,
i.e. whether the basal area level was achieved
with one intensive thinning or with several light
thinnings was not taken into account. All these
changes during the long measurement period
resulted in different intensities and intervals
between successive thinnings, as well as a
different number of thinnings. These irregularities may diminish the potential differences
between the thinning intensities. At least, the
results cannot shed any light on the profitability
of few intensive thinnings compared with several
light thinnings.
The results are based on experiments on fertile
sites in southern Finland. The experimental
stands were even-aged and almost pure planted
Norway spruce stands with a homogeneous
spatial structure. In addition, strip roads were
located outside the plots. In southern Finland, the
total production loss in Norway spruce stands
caused by strip roads at a spacing of 30 m was,
on average, 10 m3 ha1 during the 15-year period
after the first commercial thinning (Isomki and
Niemist, 1990). In practical forestry, the
distance between strip roads is nowadays 20 m
and most probably results in higher production
losses. In this study, the trees were manually
felled in random directions and the tree crowns
of the felled trees were not directed towards the
strip roads as in current mechanized thinnings.
Therefore, the nutrients released from logging
residues were spatially more evenly distributed.
Thus, the results do not completely represent the
development of normal commercial forests.
As described above, the material was rather
heterogeneous and it had some shortcomings.
The long duration of the experiments has caused

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At establishment, the experiments of this study


were not completely unthinned. Stand densities
on the unthinned plots were probably not high
enough to reduce the merchantable volume per
hectare.
Even though the differences in total volume
production between the unthinned plots and
lightly and moderately thinned plots were small,
a part of the total production was lost through
natural mortality on the unthinned plots. The
volume of dead trees was not taken into account
when calculating the merchantable volume. In
practical forestry, a part of natural mortality may
be collected in sanitary cuttings or is still usable
at the time of normal harvesting and, thus, the
differences in merchantable volume may be
somewhat larger. In the earlier measurements, the
cause of mortality was not defined and it often
remained unclear in the later ones. On the
unthinned plots, the main reason was most
probably the competition among trees. According to Abetz and Unfried (1984) and Laiho
(1987), thinning increases the risk of wind and
snow damage for a number of years after the
treatment. In contrast, the results of this study
indicate no dramatically increased damage risk
on thinned plots. However, the plots were rather
small and located within a closed stand and,
therefore, they do not represent conditions on
larger thinned areas.
The differences in basal area per hectare
between the thinned stands and their unthinned
counterparts have commonly been used as an
approximation of competition and suppression in
thinned stands (e.g. Pienaar, 1979). Growth after
thinning has been related to the basal area
remaining in the stand immediately after
thinning. In this study, the growth responses to
thinning were evaluated against the differences in
average basal area between the thinned and
unthinned plots during the whole measurement
period. The thinning intensity was therefore
calculated not only on the basis of the basal area
removed in thinnings, but also on the growth
response after the thinnings. Thus, the treatments
and possible differences in site fertility among the
plots may have an effect on average basal area
level. In addition, the length of the measurement
period varied among the experiments. Period
length may not, however, introduce any large bias
in the relative differences between the treatments

361

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Acknowledgements
We are greatly indebted to many technicians, especially
to Timo Siitonen, who have maintained and measured
the experiments. We also thank Professor Jari Hynynen
for his support and advice, Pentti Niemist and Dr
Anssi Ahtikoski for valuable comments on the manuscript, Dr Helena Henttonen for the annual increment
indices, John Derome for revision of the language and
Sointu Nenola for drawing Figure 1.

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