Article
Influence of Four Spacings between Trees and Four Samplings
Heights on Selected Wood Quality Attributes of White Spruce
(Picea glauca (Moench) Voss)
Cyriac S. Mvolo 1,2, *, Venceslas Goudiaby 3 , Ahmed Koubaa 2
Citation: Mvolo, C.S.; Goudiaby, V.;
Koubaa, A.; Stewart, J.D. Influence of
Four Spacings between Trees and
Four Samplings Heights on Selected
Wood Quality Attributes of White
Spruce (Picea glauca (Moench) Voss).
Forests 2022, 13, 1807. https://
doi.org/10.3390/f13111807
Academic Editor: Christian Brischke
Received: 6 October 2022
Accepted: 27 October 2022
Published: 30 October 2022
Publisher’s Note: MDPI stays neutral
with regard to jurisdictional claims in
published maps and institutional affil-
iations.
Copyright: © 2022 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Forests 2022, 13, 1807. https://doi.org/10.3390/f13111807
and James D. Stewart 1
1 Natural Resources Canada, Canadian Forest Service, Canadian Wood Fibre Centre,
Edmonton, AB T6H 3S5, Canada
2 Institut de Recherche sur les Forêts, Université du Québec en Abitibi-Témiscamingue,
445 boul. de l’Université, Rouyn-Noranda, QC J9X 5E4, Canada
3 Département des Sciences Biologiques, Université du Québec à Montréal, Montréal, QC H3C 3P8, Canada
* Correspondence: cyriac.mvolo@nrcan-rncan.gc.ca; Tel.: +1-825-510-1173
Abstract: Four Picea glauca (Moench) Voss trees grown at each of four square spacing intensities
between trees: 1.2 m, 1.8 m, 4.3 m, and 6.1 m in a plantation established in 1967 in the Petawawa
Research Forest, Ontario, Canada (lat. 45.59◦ N, long. 77.25◦ W, elev. 168 m) and sampled at
four different heights (1.3 m, 4.3 m, 7.3 m, 10.3 m) were used to study the impact of spacing between
trees and sampling height on nine wood quality attributes (ring width, ring density, tracheid length,
tracheid diameter, latewood proportion, intra-ring density variation, ring area, earlywood width,
and latewood width). In the juvenile wood, ring width was wider and ring density higher than
in the mature wood. Tracheid length was longer and tracheid diameter wider in the mature wood
compared to the juvenile wood. The variation of ring density between the two wood zones was
limited, and latewood proportion did not show any difference with wood zone. Sampling height
induced variation in more wood quality attributes than did spacing. Except for growth rate, spacing
between trees did not significantly impact wood quality attributes. Most of these variations were
registered between widely different spacings.
Keywords: silvicultural management; boreal forests; sampling height; wood properties; growth rate;
wood density; tracheid dimensions; juvenile wood; mature wood; latewood proportion
1. Introduction
Often called “circumpolar” because it circles the Northern hemisphere, forming a ring
around the North Pole, just south of the Arctic Circle, the boreal zone covers 1.9 billion ha,
14% of earth’s land, and 33% of earth’s forested area. Canada, Russia, the United States,
Norway, Sweden, China and a few others are the countries with forests and land in the
boreal zone. Most of Canadian forests and woodlands (75%) is located in the Canadian
boreal area, which represents 28% of the world’s boreal zone [1].
Silvicultural management such as spacing between trees is used to improve forests
productivity, shorten rotation age and meet the increasing demand in woody raw mate-
rial [2–5]. Although intensive studies have been done to determine the impact of spacing
between trees on tree growth and wood volume [6], studies of the effects of spacing on
wood properties have been limited to wood density (ring density (RD), all acronyms
used in this text are listed in Table 1) and some macroscopic characteristics, or, for many
species, absent [5]. As for many commercial tree species in the Canadian boreal forests,
impacts of spacing between trees on many Wood Quality Attributes (WQA) of white spruce
(Picea glauca (Moench) Voss) remain to be clarified.
https://www.mdpi.com/journal/forests
Forests 2022, 13, 1807 2 of 16

Table 1. Acronyms used in the text and their description.

Acronym Description
EWW Earlywood width
IDV Intra-ring density variation
JW Juvenile wood
LWP Latewood proportion
LWW Latewood width
MW Mature wood
RA Ring area
RD Ring density
RFP Ring from pith
RW Ring width
TD Tracheid diameter
TL Tracheid length
WQA Wood quality attributes

Ring width (RW) is often used to evaluate growing performances at tree level [7] and,
although easy to measure and often related to WQA, it is insufficient to evaluate wood
quality [8]. RD is probably the most studied WQA [3]. It is the key property affecting both
the stiffness and strength of structural timber [9], guides raw material selection in veneers
and appearance products manufacturing [9,10], determine the pulp yield and the quality of
paper products [11,12], and also the biomass calorific value and carbon sequestration [13,14].
Less investigated WQA include tracheid length (TL), tracheid diameter (TD), latewood
proportion (LWP), intra-ring density variation (IDV), ring area (RA), earlywood width
(EWW) and latewood width (LWW). Tracheids represent 92.5% of wood volume in white
spruce [15]. TL and TD have significant impacts on the quality of the pulp and paper and
fibre-based products [16].
The radial pattern of variation in wood properties is usually designated by the number
of annual rings counting from the pith outward (RFP), also known as the cambial age of the
ring. Changes in the radial trends are used to define the juvenile wood (JW) and mature
wood (MW) zones [17]. TD radial and longitudinal variations explain a tree’s adaptation
strategy to overcome hydraulic resistance [18,19]. Variations in TL and TD have not often
been addressed [20] because measuring anatomical properties is both time consuming
and expensive [21]. LWP is the key property determining RD value [5,22–25]. IDV is an
indication of the uniformity of wood density inside a growth ring, and is highly correlated
with RD and RW components, especially in the latewood [24]. A greater uniformity (lower
IDV) is valuable for veneer production [10]. RA tells us about tree growth in both the radial
and tangential directions [4]. Growth-related (RW, EWW, LWW, RA and LWP), anatomical
(TL and TD) and physical (RD and IDV) WQA are known to change from pith to bark [26],
with tree height [27], and in the tree’s juvenile and mature wood zones [26]. These changes
are mainly due to cambium aging and the proximity of the active living crown [2].
Within ring growth variation is due to transition from earlywood to latewood [28].
The earlywood zone is produced under the strong influence of growth hormones, especially
indole-3-acetic-acid (IAA, auxin) at the beginning of the growth season, when newly formed
buds are actively elongating, and earlywood production proceeds downward towards the
tree [3,29]. Latewood production coincides with the cessation of shoot elongation, proceeds
upward towards the crown, and has thicker cell walls, as a result of greater photosynthate
availability [3,29]. Earlywood width (EWW) is more correlated with RW than latewood
width (LWW) [23].
RW increases with increasing spacing in white spruce [30], black spruce (Picea mariana
(Mill.) B.S.P.) [4,6,31], and radiata pine (Pinus radiata D. Don) [5]. With increasing RW, a sig-
nificant decrease has been observed in TL for Norway spruce (Picea abies (L.) Karst) [32,33],
radiata pine [5], and Jack pine (Pinus banksiana Lamb.) [34]. RD is also negatively influenced
by increasing RW in white spruce [30,35], black spruce [6,23,31], Norway spruce [36] and
Jack pine [34]. However, other researchers rather found positive relationships between
Forests 2022, 13, 1807 3 of 16

increased RW and RD in balsam fir (Abies balsamea Mill) [24] and jack pine [37]. Similarly, a
positive relationship between RW and TL was documented for white spruce [30]. Bränd-
ström [38] reported an increase in TD with RW in Norway spruce. Yang and Hazenberg [31]
found both an increase and a decrease of TL with RW for black spruce depending on the
spacing. Similarly, Dinwoodie [39] found both an increase and a decrease of TL with
RW, for Sitka spruce (Picea sitchensis (Bong.) Carr.) depending on the position (juvenile,
transition and mature wood) in the tree core. Some of the discrepancies may be related to
species-specific differences, sampling strategies or age factor [40].
Sampling height and its relationship with living crown also affects changes in WQA [18].
Although an increased RW could be associated with significant variations in other WQA,
many researchers found that the variations in RD [2,3,41,42], TL [2,30,33,43], and TD [5]
with the growth rate is limited and without practical implications. Furthermore, even when
significant variations due to silvicultural management were found for RD and tracheid
dimensions, they were within the range of variations caused by age, height, genotype
and geographic locations [40]. These observations have led to the conclusion that stand
productivity may be improved without sensible loss of wood quality [2,43,44]. However,
one must keep in mind the highly relevant age effects and avoid generalizing results of
the fast-grown young trees with a high JW proportion to older trees with comparable
dimensions but lower JW proportion [40].
In comparison to MW, JW is composed of shorter, smaller tracheids with thinner walls,
larger microfibril angles, larger spiral grain angles, lower tangential and higher longitudinal
shrinkage, lower holocellulose and alpha cellulose content, higher lignin and hemicellulose
content, and lower strength properties [7]. JW is found near the pith and transitions to
MW toward the bark of older trees. It is formed under the strong regulatory influence of
the active living crown, where growth hormones, particularly auxin, are synthesized [45].
Therefore, isolated or very widely spaced trees with a deep crown as well as the top of all
trees are generally assumed to be composed entirely of JW [3,46]. The impact of spacing
between trees on WQA was found to change between JW and MW [30,47,48], with JW
being more sensitive to spacing than MW [30,31], supporting the suggestion that these
wood zones should be considered as distinct populations [49].
White spruce has a broad ecological niche and occurs in pure stands as well as in
mixed stands in association with a wide variety of both coniferous and deciduous species
in the boreal forests [50]. As such, it is widely distributed across North America, extending
from Newfoundland in the east to northwestern Alaska in the west. Its latitudinal range
is from the northern treeline across the continent, south (in the Western Cordillera) to
southern British Columbia and Alberta, following the southern boreal boundary in the
prairie provinces (Alberta, Saskatchewan, Manitoba) and east along the northern tier states
from Minnesota to Maine [51]. The impact of spacing on WQA may be species- specific
in nature and not showing any universal trend [30]. Therefore, establishing accurate
relationships between white spruce growth as affected by spacing between trees and WQA
in both JW and MW would contribute to better-informed decisions about spacing selection
for specific end uses.
The objectives of this study were: to evaluate the impacts of spacing intensities on
selected WQA of white spruce, to evaluate the impacts of sampling heights on these WQA,
and to compare trends in JW and MW zones. Radial variations of white spruce selected
WQA were also explored. To our knowledge, the influence of spacing between trees on
IDV in white spruce has not yet been assessed.

2. Materials and Methods

2.1. Stand Description
The material used in this study comes from a spacing experiment. The study site
consisted of a plantation established in 1967 in the Petawawa Research Forest, Ontario,
Canada (lat. 45.59◦ N, long. 77.25◦ W, elev. 168 m) with four square spacing intensities
between trees: 1.2 m, 1.8 m, 4.3 m, and 6.1 m. Four apparently healthy trees were randomly
Forests 2022, 13, 1807 4 of 16

sampled within each spacing treatment. All 16 trees used in this study were felled and
pruned once on the ground in July 2008 [18].

2.2. Sample Collection, Preparation, and Wood Quality Attribute Assessments

Stem discs were collected from felled trees at 1.3 m, 4.3 m, 7.3 m, and 10.3 m. Two
1.86 mm (tangential) adjacent strips centred on the pith were sawn bark to bark from
each disc. One single radius per strip was carefully selected for the analyses. Direction
was random, avoiding compression wood and knots. The first strip per disc was used
for measurement of growth-related (RW, EWW, LWW, RA, and LWP) and physical prop-
erties (RD, maximum ring density and minimum ring density). These properties were
measured at a 25 µm linear resolution step size with an X-ray densitometer (Quintek Mea-
surements Systems QMS model QTRS-01X, Knoxville, TN, USA). The boundary between
earlywood and latewood was delineated using the intra-ring wood density profile and the
maximum derivative method [10]. Intra-ring density variation was computed as the differ-
ence between the maximum and minimum densities of the ring. Using the second wood
strip, wood sticks from earlywood and latewood were taken at 3, 6, 9, 12, 15, 20, 25, and
30 growth rings. This sampling was performed at a fixed growth ring and not at deter-
mined calendar years [18]. Wood sticks were macerated using a Franklin [52] solution
consisting of (1:1 v/v) hydrogen peroxide diluted to 30% and concentrated glacial acetic
acid. Each stick was placed in a separate test tube of Franklin solution, and the tube
immersed in hot distilled water (85–90 ◦ C) for 5 to 6 h until complete lignin dissolution.
Delignified wood sticks were gently shaken in water with a laboratory blender to obtain
a tracheid suspension. Earlywood and latewood TL were measured with a Fibre Quality
Analyzer, LDA02 FQA (Op Test Equipment Inc. Hawkesbury, Ontario, Canada). A total of
4000 tracheids were measured in every sample. The TL in each zone was measured as the
weight weighted length, LWW = ΣniLi3 /ΣniLi2 (where i = 1, 2, 3 . . . N categories; n = fibre
count in the (ith) category; L = contour length). Using this method, measurements were
similar to true TL measurements and controlled for the bias caused by the large number
of fines generated during the preparation process [21,32]. Average ring TL and TD were
computed by weighting TL and TD of each wood zone with the relative RW.

2.3. Statistical Analyses

Analyses of variance were performed using the lme function of the nlme package [53]
of R-4.2.1with R Studio version 4.1.0 [54,55]. Prior to analyses, conditions for linear model,
i.e., linearity, normality, and homogeneity of variances were tested by visual inspection
using the plot function of R. When those conditions were not met, a natural logarithm
transformation was performed to the concerned WQA, which were RW, LWP, RA, EWW,
and LWW. All fitted models aimed to explain how a given wood quality attribute (RW,
RD, IDV, LWP, TL, TD, RA, EWW, and LWW) varied: radially (i.e., inter-ring variation),
longitudinally (i.e., intra-ring and inter-height variation) restricted to the juvenile wood
section, from tree to tree (i.e., inter-tree variation) irrespective of the spacing treatment
applied, and following spacing treatment performed (i.e., spacing between trees impact).
Plot, i.e., spacing treatment, was the experimental unit, and all the analyses were performed
at breast height (1.3 m) except for the intra-ring and inter-height variation of juvenile wood
which included all the stem heights (1.3 m, 4.3 m, 7.3 m, and 10.3 m). Model 1 was used
to test inter-ring, intra-ring, and inter-tree effects (at breast height) on WQA. Model 2 was
used to test the effects of combined spacing treatment and height on WQA.

Yi = β0 + β1 Xi + γi + εi (1)

where Y is the response variable representing a wood quality attribute (WQA) and X the
predictor variable explaining a WQA, for the ith Tree. The predictor X was RFP for the
inter-ring effects, Height for the intra-ring effects, and Tree for the inter-tree effects. The
predictors RFP and Height were considered as categorical variables. The models also
included γ as a tree random effect (γi ~ N(0, σi2 )) and εi as an error term (εi ~ N(0, σi2 )). The
Forests 2022, 13, 1807 5 of 16

intercept is β0 and the coefficient of the predictor is β1 . For the inter-ring analysis, tree rings
occurred each year, so that we considered a first-order autoregressive AR(1) correlation.
Spacing between trees and height effects on WQA were tested using the
following model:
Yi = β0 + β1 Si + β2 Hi + β3 Si Hi + γi + εi , (2)
where Y is a WQA, S the spacing between trees, H tree height, along with their interaction
for the ith Tree, γ and εi are as previously described. β0 to β3 represent the intercept and
the coefficients for spacing, height, and their interaction, respectively. The predictors S and
H were considered as categorical variables.
For all models, predictors were considered significant at p < 0.05 based on F-test.
Significant responses always prompted us to check which levels of this predictor variable
were significant based on a t-test. t-tests were performed on estimated marginal means,
with p-value adjustments for multiple comparisons using Bonferroni correction.

3. Results
3.1. Inter-Ring Radial Variation at Breast Height
Inter-ring radial variation at breast height was significantly different for RW, RD, TL,
TD, IDV, RA, EWW, and LWW whereas it did not vary for LWP (Table 2—Part 1). RW (and
EWW) slightly increased from pith to the 7th ring (1.2 and 1.8 m spacing) and to the 15th
ring (4.3 and 6.1 m spacing) before decreasing toward the bark. LWW decreased from pith
to bark (1.2 and 1.8 m spacing), while no clear pattern was exhibited in wider spacings
(4.3 and 6.1 m). RA slightly increased from pith to the 10th ring (1.2 and 1.8 m spacing)
before levelling off. RA increase lasted until the 17th ring (4.3 and 6.1 m spacing) before
decreasing toward the bark. While RD increased toward the bark in the 1.2 and 1.8 m
spacing, a decreasing trend was observed at 4.3 and 6.1 m spacing. Increasing RFP was
associated with longer and wider tracheids. IDV sharply increased for the first 10 rings
and tended to level off thereafter (graph not shown).

Table 2. Analyses of variance.

Part 1: Inter-Ring Radial Variations at Breast Height.

WQA Predictor Num-DF Den-DF F-Value
log (RW) ‡ 32 406 23.56 ***
RD 32 406 1.978 *
IDV 32 398 10.6268 ***
log (LWP) ‡ 32 397 1.17 ns
TL Ring from pith 7 76 116.92 ***
TD 7 75 13.10 ***
log (RA) ‡ 32 405 20.93 ***
log (EWW) ‡ 32 397 16.67 ***
log (LWW) ‡ 32 397 6.62 ***
Part 2: Inter-tree variation at breast height
log (RW) ‡ 15 434 5.66 ***
RD 15 435 2.351 *
IDV 15 427 6.74 ***
log (LWP) ‡ 15 427 2.21 *
TL Tree 15 80 1.36 ns
TD 15 79 2.19 *
log (RA) ‡ 15 434 8.92 ***
log (EWW) ‡ 15 426 4.82 ***
log (LWW) ‡ 15 426 2.77 **
Forests 2022, 13, 1807 6 of 16

Table 2. Cont.

Part 3: Intra-ring longitudinal variation of juvenile wood

log (RW) ‡ 3 691 12.67 ***
RD 3 692 2.38 ns
IDV 3 687 7.741 ***
log (LWP) ‡ 3 686 0.16 ns
TL Height 3 112 3.74 *
TD 3 111 0.28 ns
log (RA) ‡ 3 691 15.24 ***
log (EWW) ‡ 3 686 8.64 ***
log (LWW) ‡ 3 686 4.08 *
Part 4: Inter-tree variation for different spacings between trees (I_Spacing) and heights
I_Spacing 3 12 11.85 **
log (RW) ‡ Height 3 1061 11.19 ***
I_Spacing × Height 9 1061 1.69 ns
I_Spacing 3 12 1.24 ns
RD Height 3 1062 1.33 ns
I_Spacing × Height 9 1062 1.25 ns
I_Spacing 3 12 1.73 ns
IDV Height 3 1045 12.18 ***
I_Spacing × Height 9 1045 5.13 ***
I_Spacing 3 12 0.48 ns
log (LWP) ‡ Height 3 1044 0.14 ns
I_Spacing × Height 9 1044 1.08ns
I_Spacing 3 12 0.16 ns
TL Height 2 134 0.21 ns
I_Spacing × Height 6 134 0.92 ns
I_Spacing 3 12 0.28 ns
TD Height 2 133 0.14 ns
I_Spacing × Height 6 133 1.05 ns
I_Spacing 3 12 11.56 ***
log (RA) ‡ Height 3 1061 27.63 ***
I_Spacing × Height 9 1061 3.42 **
I_Spacing 3 12 10.39 *
log (EWW) ‡ Height 3 1044 7.21 **
I_Spacing × Height 9 1044 1.05 ns
I_Spacing 3 12 14.98 **
log (LWW) ‡ Height 3 1044 2.91 **
I_Spacing × Height 9 1044 1.70 *
All intercepts are highly significant (p-value < 0.0001) with Num-DF =1 and are not presented, WQA: Wood
Quality Attributes, RW: ring width, RD: ring density, IDV: intra-ring density variation, LWP: Latewood Proportion,
TL: tracheid length, TD: tracheid diameter, RA: ring area, EWW: earlywood width, and LWW: latewood width,
‡ : natural logarithm transformation, Num-DF: numerator degree of freedom, Den-DF: denominator degree of

freedom, *, **, and ***, indicate significance at p < 0.05, p < 0.001, and p < 0.0001, respectively. ns indicates not
significant at p ≤ 0.05.

Figure 1 gives an easy and quick overview of the paired comparisons between the
9 studied WQA. This figure also shows the relationships between juvenile and mature
wood. In this figure, RFP is the label in both the x-axis and the y-axis, like the matrix
representation of WQA one can find in a table. By doing so, we were able to compare all
possible variations of a given WQA between rings. Only significant comparisons were
represented with a dot in Figure 1, meaning that all blank spaces (and missing rings) are
non-significant comparisons.
Forests 2022, 13, x FOR PEER REVIEW 8 of 17

Forests 2022, 13, 1807 7 of 16

Figure 1. Outcomes of the models comparing a given ring with, respectively all other rings at breast height. The blue line represents best fit and shaded region
Figure 1. Outcomes of the models comparing a given ring with, respectively all other rings at breast height. The blue line represents best fit and shaded region the
the 95% confidence level interval for predictions. Only significant differences (black dotes, p > 0.05) were shown (p-value adjustments for multiple comparisons
95% confidence level interval for predictions. Only significant differences (black dotes, p > 0.05) were shown (p-value adjustments for multiple comparisons using
using Bonferroni correction). RW: ring width, RD: ring density, IDV: intra-ring density variation, TL: tracheid length, TD: tracheid diameter, RA: ring area,
Bonferroni correction). RW: ring width, RD: ring density, IDV: intra-ring density variation, TL: tracheid length, TD: tracheid diameter, RA: ring area, EWW:
EWW: earlywood
earlywood width,width, and LWW:
and LWW: latewood
latewood width.width.
Forests 2022, 13, 1807 8 of 16

At a first glance, patterns of paired rings comparisons were different between wood
quality attributes (Figure 1). For RW, no significant difference was noted between the first
23 rings (except for ring 9 vs. ring 19 and consecutive rings), while those later were generally
different from the 25th ring onward. For EWW, rings between 3 and 22 were not different
amongst them (except for ring 15 vs. ring 21). These rings were all different from ring
22 onwards. Ring 3 was significantly different from rings 26–30 and ring 22 was significantly
different from only ring 30. For TL, there was no difference in the mature wood zone. There
were few significant paired comparisons among TL in rings of the juvenile wood, but more
were found between TL in the juvenile wood vs. TL in the mature wood. Except for a
single significant paired comparison in the MW, TD behaves similarly to TL, but with
fewer significant paired comparisons among TD in rings of the JW section. For IDV, with
very few exceptions, most of the variations involved the first 7 rings against values in ring
10 onward. For RA, ring 3 was significantly different from ring 12 to 27, and all other rings
between 3 and 12 were significantly different from rings 12 to 26, at the most. For these
comparisons, the higher the rank of the ring, the fewer the differences from other rings.
Few other RA differences were found between rings 15–18 vs. rings 28–30. For RD, the
handful of significant paired comparisons all involved ring 3 (vs. rings 9, 10, 16, 17, 23 and
25). For LWW, only ring 8 was noted to be significantly different from rings 26, 27, and
28. LWP was purposely dropped from Figure 1 due to consistent patterns between rings
(totally blank figure).

3.2. Variations between Wood Type

Mean values of all selected WQA are presented in Table 3—Part 1 for the four heights,
Table 3—Part 2 for the four spacing between trees and Table 3—Part 3 for the JW and the MW
zones. Diagnostics of WQA were performed by analyzing the data dispersion (Figure 2). In
the following descriptions, range will refer to the interquartile range (25th–75th percentile)
rather than the minimum and maximum values. For RW and EWW, radial variation was
similar in the two wood types; however, the values of RW were higher in JW compared to
MW (Figure 2). RD and LWP approximately had same ranges and median between juvenile
and mature wood. More variability for TL and TD was observed in JW compared to MW.
TL and TD were greater in the mature compared to the juvenile wood. The ranges of IDV
were similar between wood types, but IDV values were higher in the MW compared to the
JW. The range of RA was approximately twice as large for MW compared to JW, whereas
the median of JW was lower than median of MW. LWW had a lower median value in MW.

Table 3. Mean values (and standard error) of wood quality attributes.

Part 1—with Height

3 3 LWW
Height (m) RD (kg/m ) IDV (kg/m ) LWP (%) RW (mm) TD (µm) TL (mm) RA (mm2 ) EWW (mm)
(mm)
1.3 452.4 (1.6) 507.9 (6.7) 21.8 (0.6) 3.9 (0.1) 32.3 (0.2) 2.2 (0.0) 784.7 (37.8) 3.0 (0.1) 0.9 (0.0)
4.3 447.9 (1.9) 555.2 (7.4) 22.0 (0.8) 4.1 (0.1) 33.4 (0.2) 2.5 (0.0) 855.6 (39.9) 3.1 (0.1) 0.9 (0.1)
7.3 444.3 (2.6) 537.1 (9.3) 22.2 (1.0) 3.5 (0.1) 33.3 (0.2) 2.4 (0.0) 662.3 (38.7) 2.7 (0.1) 0.8 (0.1)
10.3 443.5 (2.5) 501.9 (13.0) 22.4 (1.3) 3.6 (0.1) - - 517.9 (34.5) 2.9 (0.1) 0.8 (0.1)
Part 2—with spacing
LWW
Spacing (m) RD (kg/m3 ) IDV (kg/m3 ) LWP (%) RW (mm) TD (µm) TL (mm) RA (mm2 ) EWW (mm)
(mm)
1.2 445.6 (2.6) 537.1 (9.9) 21.6 (0.9) 2.9 (0.1) 31.9 (0.3) 2.3 (0.0) 429.7 (19.7) 2.3 (0.1) 0.6 (0.0)
1.8 445.9 (2.0) 473.5 (10.1) 22.4 (0.9) 2.9 (0.1) 32.7 (0.2) 2.3 (0.0) 394.2 (15.7) 2.3 (0.1) 0.6 (0.0)
4.3 453.4 (2.8) 534.4 (10.0) 20.1 (1.0) 4.2 (0.1) 33.5 (0.3) 2.3 (0.0) 812.9 (43.4) 3.3 (0.1) 0.9 (0.1)
6.1 446.2 (2.7) 568.7 (9.0) 24.1 (1.2) 5.4 (0.1) 32.9 (0.4) 2.2 (0.0) 1323.0 (66.4) 4.0 (0.1) 1.4 (0.1)
Part 3—in the juvenile and mature wood zones

3 3 LWW
Wood zone RD (kg/m ) IDV (kg/m ) LWP (%) RW (mm) TD (µm) TL (mm) RA (mm2 ) EWW (mm)
(mm)
JW 447.5 (1.3) 527.6 (5.2) 22.1 (0.5) 3.8 (0.1) 32.8 (0.2) 2.3 (0.0) 727.4 (24.4) 2.96 (0.0) 0.84 (0.0)
MW 442.0 (2.2) 616.0 (6.9) 21.9 (0.7) 2.3 (0.1) 36.0 (0.2) 3.1 (0.0) 1206.6 (58.0) 1.86 (0.1) 0.50 (0.0)
RD: ring density, IDV: intra-ring density variation, LWP: Latewood Proportion, RW: ring width, TD: tracheid
diameter, TL: tracheid length, RA: ring area, EWW: earlywood width, LWW: latewood width, JW: juvenile wood,
and MW: mature wood.
Forests 2022, 13, x FOR PEER REVIEW 10 of 17

Forests 2022, 13, 1807 9 of 16

Figure 2. Radial variation of wood type (Juv: Juvenile, Mat: Mature), at the stem scale, of wood quality attributes of white spruce. For each wood type, the dispersion
Figure 2. Radial variation of wood type (Juv: Juvenile, Mat: Mature), at the stem scale, of wood quality attributes of white spruce. For each wood type, the
of data for a given wood quality attribute is represented by the interquartile range (distance between the upper and lower quartiles) and also the minimum and the
dispersion of data for a given wood quality attribute is represented by the interquartile range (distance between the upper and lower quartiles) and also the
maximum values. The wood quality attributes are: RW: ring width, RD: ring density, IDV: intra-ring density variation, LWP: latewood proportion, TL: tracheid
minimum and the maximum values. The wood quality attributes are: RW: ring width, RD: ring density, IDV: intra-ring density variation, LWP: latewood propor-
length,
tion, TL:TD: tracheid
tracheid diameter,
length, RA: ringdiameter,
TD: tracheid area, EWW:RA:earlywood width, earlywood
ring area, EWW: and LWW: width,
latewoodandwidth.
LWW: latewood width.
Forests 2022, 13, 1807 10 of 16

3.3. Inter-Tree Variation of Wood Quality Attributes at Breast Height

Inter-tree variation at breast height of WQA were assessed according to their intercept
or initial value and to the slope of the Tree predictor (Table 2—Part 2). Overall, the intercepts
for inter-tree initial variations were significant for all WQA. As shown by their slopes, inter-
tree effects were also significant for RW, RD, IDV, TD, RA, EWW, LWW, and LWP. By
contrast, there were no significant tree effects on TL.

3.4. Effects of Spacing between Trees and Height on Juvenile Wood Quality Attributes
For RW, the ranges of intra-ring longitudinal variation of juvenile wood of WQA were
quite similar between 4.3, 7.3 and 10.3 m, and lower at 1.3 m, with the median higher in
4.3 m compared to other tree heights (Figure 3). The two lower heights (1.3 m and 4.3 m)
had comparable mean RW, and the two upper heights (7.3 and 10.3 m) comparable mean
RW. RW was greater at lower heights compared to upper heights (Table 3—Part 1). Ranges
(and medians) of RD were similar along the stem. Although a slight decreasing trend could
be observed with increasing height, mean RD were comparable along the stem. Range
of IDV was lower at 4.3 m and comparable for other heights. Their medians displayed a
decreasing trend between 4.3 and 10.3 m height, while the value at 1.3 m was lower than its
counterparts at 4.3 and 7.3 m, and higher at 10.3 m. Mean IDV were comparable between
1.3 and 10.3 m, and decreased with height starting at 4.3 m. For LWP, the largest range
was observed at 4.3 m while it was comparable for the other heights. For all heights, LWP
was right skewed due to higher frequency of high values above the median, especially at
4.3 m. Mean LWP were comparable along the stem. Ranges of TL hardly differed with
height along the stem, with a left skew at 4.3 m, contrasting with small right skew at
1.3 m and 7.3 m. Mean TL was comparable along the stem. TD was left skewed at 1.3 m
and 4.3 m height, contrasting with right skew at 7.3 m height. Mean TD was comparable
along the stem. Ranges of RA were greater at 4.3 m followed by 1.3 m, 7.3 m, and 10.3 m,
respectively. It was right skewed at all heights. Mean RA decreased with height starting at
4.3 m. Although smaller than the highest value (at 4.3 m), the value at 1.3 m was higher
than values at 7.3 and 10.3 m. The range of LWW was greater at 4.3 m compared to the
other heights, which were similar. It was right skewed at all height (Figure 3). Mean LWW
were comparable along the stem (Table 3—Part 1).
Intra-ring longitudinal variation of juvenile wood was significant for RW, IDV, TL,
RA, EWW, and LWW, whereas LWP, TD, and RD did not change, (Table 2—Part 3). Breast
height RW did not show a linear trend, as it was significantly higher only compared to
height 7.3 m. Similarly, RW was significantly higher for height 4.3 m when contrasted with
height 7.3 m and 10.3 m, respectively. TL was significantly lower at breast height compared
to height 4.3 m. There was also significant lower IDV in 1.3 m compared to 4.3 m. By contrast,
IDV was significantly higher in 4.3 m compared to 10.3 m, and also 7.3 m compared to 10.3 m.
The effects of spacing and sampling height interacted for RA, IDV and LWW
(Table 2—Part 4). While spacing did not show any significant effect on IDV, height and its
interaction with spacing did significantly affect IDV. Both spacing and height independently
affected the three components of radial growth (RW, EWW and LWW). By contrast, no
significant effects of spacing between trees, height, or their interaction were found for RD,
LWP, TL, and TD.
Contrast analyses for different spacings were performed on wood quality attributes
significantly affected by spacing, height or their interaction. Hence, a pairwise comparison
per spacing was conducted for RW for different heights. In summary, RW significantly
increased at all heights between the spacings: 1.2 vs. 6.1, 1.8 vs. 6.1 and 1.2 vs. 4.3 (with
the exception of height 10.3 m). The contrast 1.8 vs. 4.3 was also significant at heights
1.3 m and 4.3 m. Except for breast height, RW did not significantly increase between the
spacings 4.3 vs. 6.1. No difference was found between the spacings 1.2 vs. 1.8. Except for
the significantly (0.05) lower IDV in the spacing 1.8 compared to 6.1 at 7.3 m height, all
other contrasts between spacings between trees and heights were not significant.
Forests 2022, 13, x FOR PEER REVIEW 12 of 17
Forests 2022, 13, 1807 11 of 16

Figure 3. Intra-ring longitudinal variation of wood quality attributes of white spruce at different heights up the stem. The plots depict the interquartile range (the
Figure 3. Intra-ring longitudinal variation of wood quality attributes of white spruce at different heights up the stem. The plots depict the interquartile range (the
‘box’) and individual data points in the first and fourth quartiles, including the minimum and the maximum values (the ‘whiskers’). The wood quality attributes
‘box’) and individual data points in the first and fourth quartiles, including the minimum and the maximum values (the ‘whiskers’). The wood quality attributes
are: RW: ring width, RD: ring density, IDV: intra-ring density variation, LWP: latewood proportion, TL: tracheid length, TD: tracheid diameter, RA = ring area,
are: RW: ring width, RD: ring density, IDV: intra-ring density variation, LWP: latewood proportion, TL: tracheid length, TD: tracheid diameter, RA = ring area,
EWW
EWW = = earlywood
earlywood width,
width, and
and LWW
LWW == latewood
latewood width.
width.
Forests 2022, 13, 1807 12 of 16

4. Discussion
4.1. Impact of Spacing on the Inter-Ring Radial Variation at Breast Height
The initial increase followed by a steady decrease of RW is in agreement with known
pattern in white spruce [26,30] and black spruce [4,31]). The longer initial increase pattern
with wider spacing agreed with Yang’s [30] study, which found an initial increase for the
first 6 rings at square spacings of 1.8 m and 2.7 m, comparable to the increase during the
first 7 rings we found at square spacings of 1.2 m and 1.8 m. At wider spacings, the phase
of RW increase lasted longer, up to 10 rings at 3.6 m spacing (Yang’s [30] widest spacing),
and up to 15 rings at square spacings of 4.3 m and 6.1 m of our study, both of which are
wider than Yang’s [30] widest spacing. This longer period of increase may be caused by the
elongated crown and delayed self-pruning in the wider spacings, which favours prolonged
auxin production [3,40].
In closer spacings, RD presented the type II pattern (RD is high at the pith, decreases
with increasing RFP to a minimum, before again increasing) as described by Panshin and
de Zeuuw [49] and Schimleck, Dahlen and Auty [22], and reported for white spruce [26,30].
That pattern does not hold in wider spacings, which instead presented the Type III trend
(monotonic decrease from pith to bark) [22,49]. It is not uncommon to find different RD
patterns with different species, but the type II variation is the one most often reported for
spruces. This differences between radial RD patterns of even-aged white spruce growing in
the same environment is an interesting result of this study. It may explain why predicting
wood density is challenging, compared to other WQA such as tracheid dimensions (TL
and TD), RW, microfibril angle, and modulus of elasticity [56–58]. In western hemlock
(Tsuga heterophylla (Raf.) Sarg.), Fabris [59] found the same type II pattern at different
spacings, but age at minimum RD increased with spacing between trees, suggesting that
even if spacing intensity does not change the RD pattern type as seen in this study, it may
change the location of the inflection point of the radial trend, as seen with RW in this study.
Earlywood TL was found to behave similarly to ring TL [26], allowing comparing
our ring TL to Yang’s [30] earlywood TL. As expected, TL and TD followed Sanio’s first
law [60] with values increasing from pith to bark, as was also reported in other studies of
white spruce [26,30].
LWP and IDV exhibited the same pith-to-bark radial patterns as described for another
white spruce study by Mvolo et al. [26]. These consistent patterns suggest that cambium
maturation had the same impact on these WQA irrespective of the spacing between trees.
The significant impact of cambial age in most of the WQA was expected, as age (ring
number from pith) is the first factor explaining WQA variation in the radial direction [26].
The fact that LWP did not vary with age suggest that the relative amounts of EWW and
LWW produced over the year did not change significantly.

4.2. Differences between Juvenile Wood and Mature Wood Zones

The mean values presented in Table 3—Part 3, the comparisons summarized in Figure 1
for breast height data and the radial variations in Figure 2 for the whole stem highlight the
differences between JW and MW. The higher RW (and EWW) found in the JW compared
to the MW (Figure 2) agrees with earlier studies [30,31]. It is explained by the vigorous
growth in early growth stages, caused by crown proximity and high production of growth
hormones [3]. Figure 1 shows that growth rate (RW, EWW, and LWW) was comparable
within the first 20 rings, and that these rings grew larger than rings starting at age 25 from
pith (RW and EWW). Alteyrac, Zhang, Cloutier and Ruel [4] preferred RA over RW for
transition age determination, and found clearly different maturation ages (i.e., significant
differences between a WQA value along the radial profile) with RW and RA, as one can see
in Figure 1. The comparable means in both zones and the similar range (Figure 2), as well
as the absence of any significant difference in Figure 1, suggest that LWP does not change
from JW to MW.
The mean values for RD were comparable in both zones, agreeing with earlier stud-
ies [30,31]. Only a handful of significant comparisons were found, all involving the very
Forests 2022, 13, 1807 13 of 16

juvenile third ring. This agrees with an earlier finding that most of the negative impact
of growth rate on RD happens in the JW zone [61]. The smaller IDV value near the pith
suggests that higher growth rate favors wood uniformity.
Our observations of longer and wider tracheids in the mature wood agrees with earlier
studies (Yang [30], Yang and Hazenberg [31]). It is known to be caused by maturation
of cambium initial and rate of anticlinal divisions [62]. Most of the variation in TL and
TD in Figure 1 involved the same samples. As a rule, the more the rank from pith of a
ring increased, the less it showed significant difference with other rings (the opposite was
also true).

4.3. Variations Caused by Tree, Sampling Height and Spacing between Trees
Both spacing intensity and sampling height significantly affected the four WQA that
were growth rate proxies (RW, RA, EWW, and LWW). Spacing affected none of the other
five WQA (RD, IDV, LWP, TL, and TD), although IDV was affected by height. This result is
in agreement with findings on white spruce [30], black spruce [4,6] and radiata pine [5].
However, even for instances of significant growth rate (RW) variation, they were registered
mostly between widely different spacing (1.2 vs. 6.1, 1.8 vs. 6.1 and 1.2 vs. 4.3), in
agreement with finding in white spruce [30] and black spruce [6,31] of comparable age. In
agreement with our study, Lasserre, Mason, Watt and Moore [5] also found no difference in
tracheid width with spacing in juvenile radiata pine grown in New Zealand. On the other
hand, Lasserre, Mason, Watt and Moore [5] observed significant change in TL and LWP,
and Lindström [25] showed an increase in TD with ring width in mature Norway spruce
grown in Sweden, which contradicted our results. Similar to Lasserre, Mason, Watt and
Moore [5] who analyzed only juvenile wood, Yang [30] also found a significant impact of
spacing on white spruce juvenile wood RD and TL when contrasting the widest and the
narrowest spacings, but no spacing impact was found for their mature wood RD and TL.
This suggests that Yang’s [30] significant impact may disappear if the whole pith-to-bark
data are aggregated. Overall, this advocates for earlier findings that, except for growth
features, stand density management does not significantly impact WQA [2,43,44].
The decreasing of RW with sampling height is in agreement with Alteyrac, Zhang,
Cloutier and Ruel’s [4] demonstration that growth rate decreases from the base to the top of
the stem. This was explained by reduced activity as cambium matures and, more probably,
the reduction of crown proportion (and growth rate), together with increased competition
associated with stand age [4]. Alteyrac, Zhang, Cloutier and Ruel [4] also found a significant
variation of maximum ring density with sampling height, which can explain the significant
variation we found for IDV. However, the lack of variation of RD with sampling height in this
study contrasts with Alteyrac, Zhang, Cloutier and Ruel’s [4] finding.
Sampling height affected more WQA than did spacing intensity in our study. Alteyrac,
Zhang, Cloutier and Ruel [4] found that variation due to sampling height is larger than
that due to stand density. Although it varied less than EWW, the significant impact of both
spacing between trees and height on LWW contradicts Zhang and Chauret’s [6] finding in
black spruce that most of the variation in ring width is observed in EWW only, while no
variation is observed in LWW. Given the reduced significance of the interactions between
spacing intensity and sampling height, our study agreed with Alteyrac, Zhang, Cloutier
and Ruel [4] in concluding that the influence of stand density on wood traits (mainly growth
features) is additive and remains the same regardless of sampling height. By contrast, in
our study, variations between moderately different spacing (1.8 vs. 4.3 and 4.3 vs. 6.1)
applied only at given heights and may not represent a meaningful rationale to base decision
on if increasing the whole tree/stand production is the main goal. This support Echols’ [47]
and Yang’s [30] conclusion that optimal spacing for growth rate and volume production lies
somewhere between wide (greatest diameter growth) and narrow (maximum tree number
without inducing mortality due to competition) spacings.
Forests 2022, 13, 1807 14 of 16

5. Conclusions
The objective of this study was to improve our understanding of the relationship
between growth rate in four square spacing intensities between trees (1.2 m, 1.8 m, 4.3 m,
and 6.1 m) and nine selected wood quality attributes. Only growth rate proxies (ring
width, earlywood width, latewood width and ring area) are affected by spacing between
trees. Based on ring width, this happens only when widely different spacing (1.2 vs. 6.1,
1.8 vs. 6.1 and 1.2 vs. 4.3) are considered. One can therefore advise growing bigger trees
through wider spacing to lower rotation age, after determining the optimal spacing for
growth rate and volume production. Sampling height affected more wood quality attributes
(ring width, earlywood width, latewood width, ring area and intra-ring density variation)
than did spacing intensity. Some wood quality attributes were found to greatly change from
juvenile wood to mature wood (ring width, earlywood width, ring area, tracheid length,
tracheid diameter and intra-ring density variation) while other wood quality attributes
showed a limited (ring density and latewood width) or totally absent (latewood proportion)
variation. However, no significant effects of spacing between trees, height, and their
interaction were found on the remaining (ring density, latewood proportion, tracheid
length, and tracheid diameter) wood quality attributes. Since wood quality is a subjective
judgment one makes based on the end use of the resource and preferably in economic
terms, one must also keep in mind that other wood quality attributes not considered in this
study (knots, taper, stem form, compression wood, spiral grain, juvenile wood proportion,
lignin and extractive proportion, strength and stiffness) may be negatively impacted by
wide spacing.

Author Contributions: Conceptualization, C.S.M.; methodology, C.S.M. and A.K.; formal analysis,
V.G.; resources, C.S.M., A.K. and J.D.S.; data curation, C.S.M.; writing—original draft preparation,
C.S.M. and V.G.; writing—review and editing, C.S.M., V.G., A.K. and J.D.S.; supervision, C.S.M.;
funding acquisition, C.S.M., A.K., and J.D.S. All authors have read and agreed to the published
version of the manuscript.
Funding: Natural Resources Canada funded this research through the Forest Innovation
Program (FIP).
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Contact corresponding author for data sharing.
Acknowledgments: C.S.M. is grateful to M. Christopher Helmeste for constructive discussions in an
earlier stage of this project.
Conflicts of Interest: The authors declare no conflict of interest.

References
1. Natural Resources Canada. 8 Facts about Canada’s Boreal Forest. Available online: https://www.nrcan.gc.ca/our-natural-
resources/forests/sustainable-forest-management/boreal-forest/8-facts-about-canadas-boreal-forest/17394 (accessed on
26 October 2022).
2. Herman, M.; Dutilleul, P.; Avella-Shaw, T. Growth rate effects on temporal trajectories of ring width, wood density, and mean
tracheid length in Norway spruce (Picea abies (L.) Karst.). Wood Fiber Sci. 1998, 30, 6–17.
3. Larson, P.R.; Kretschmann, D.E.; Clark, A., III; Isebrands, J.G. Formation and Properties of Juvenile Wood in Southern Pines: A Synopsis;
General Technical Report, FPL-GTR-129; Department of Agriculture, Forest Service, Forest Products Laboratory: Madison, WI,
USA, 2001; p. 42.
4. Alteyrac, J.; Zhang, S.Y.; Cloutier, A.; Ruel, J.C. Influence of stand density on ring width and wood density at different sampling
heights in black spruce (Picea mariana (Mill.) B.S.P.). Wood Fiber Sci. 2005, 37, 83–94.
5. Lasserre, J.-P.; Mason, E.G.; Watt, M.S.; Moore, J.R. Influence of initial planting spacing and genotype on microfibril angle, wood
density, fibre properties and modulus of elasticity in Pinus radiata D. Don corewood. For. Ecol. Manag. 2009, 258, 1924–1931.
[CrossRef]
6. Zhang, S.Y.; Chauret, G. Impact of Initial Spacing on Tree and Wood Characteristics, Product Quality and Value Recovery in Black Spruce
(Picea Mariana); Forintek Canada Corp: Fredericton, NB, Canada, 2001.
Forests 2022, 13, 1807 15 of 16

7. Zobel, B.J.; Sprague, J.R. Juvenile Wood in Forest Trees; Springer Series in Wood Science: Berlin/Heidelberg, Germany, 1998; p. 300.
8. Drew, D.M.; Downes, G.M.; Seifert, T.; Eckes-Shepard, A.; Achim, A. A Review of Progress and Applications in Wood Quality
Modelling. Curr. For. Rep. 2022, 1–16. [CrossRef]
9. Cown, D.J. Corewood (juvenile wood) in Pinus radiata—Should we be concerned? N. Z. J. For. Sci. 1992, 22, 87–95.
10. Koubaa, A.; Zhang, S.Y.; Makni, S. Defining the transition from earlywood to latewood in black spruce based on intra-ring wood
density profiles from X-ray densitometry. Ann. For. Sci. 2002, 59, 511–518. [CrossRef]
11. Jozsa, L.A.; Middleton, G.R. A Discussion of Wood Quality Attributes and Their Practical Implications; FPInnovations; Special
Publication SP-34: Vancouver, BC, Canada, 1994; p. 42.
12. Molteberg, D.; Hoibo, O. Modelling of wood density and fibre dimensions in mature Norway spruce. Can. J. For. Res. 2007, 37,
1373–1389. [CrossRef]
13. Bowyer, J.L.; Shmulsky, R.; Haygreen, J.G. Le Bois Et Ses Usages; Centre collégial de développement de matériel didactique:
Montréal, QC, Canada, 2005; p. 528.
14. Obernberger, I.; Thek, G. Physical characterisation and chemical composition of densified biomass fuels with regard to their
combustion behaviour. Biomass Bioenergy 2004, 27, 653–669. [CrossRef]
15. Law, K.N.; Lapointe, M. Chemimechanical pulping of boles and branches of white spruce, white birch, and trembling aspen. Can.
J. For. Res. 1983, 13, 412–418. [CrossRef]
16. Beaulieu, J. Genetic variation in tracheid length and relationships with growth and wood traits in eastern white spruce
(Picea glauca). Wood Fiber Sci. 2003, 35, 609–616.
17. Mvolo, C.S.; Koubaa, A.; Beaulieu, J.; Cloutier, A.; Mazerolle, M.J. Variation in Wood Quality in White Spruce (Picea Glauca
(Moench) Voss). Part I. Defining the Juvenile—Mature Wood Transition Based on Tracheid Length. Forests 2015, 6, 183–202.
[CrossRef]
18. Mvolo, C.S.; Koubaa, A.; Defo, M.; Beaulieu, J.; Yemele, M.-C.; Cloutier, A. Prediction of tracheid length and diameter in white
spruce (Picea glauca (Moench) Voss). IAWA J. 2015, 36, 186–207. [CrossRef]
19. Olson, M.E.; Anfodillo, T.; Rosell, J.A.; Petit, G.; Crivellaro, A.; Isnard, S.; León-Gómez, C.; Alvarado-Cárdenas, L.O.; Castorena,
M. Universal hydraulics of the flowering plants: Vessel diameter scales with stem length across angiosperm lineages, habits and
climates. Ecol. Lett. 2014, 17, 988–997. [CrossRef] [PubMed]
20. Anfodillo, T.; Petit, G.; Crivellaro, A. Axial conduit widening in woody species: A still neglected anatomical pattern. IAWA J.
2013, 34, 352–364. [CrossRef]
21. Fries, A.; Ericsson, T.; Morling, T. Measuring relative fibre length in Scots pine by non-destructive wood sampling. Holzforschung
2003, 57, 400–406. [CrossRef]
22. Schimleck, L.R.; Dahlen, J.; Auty, D. Radial patterns of specific gravity variation in North American conifers. Can. J. For. Res. 2022,
52, 889. [CrossRef]
23. Koubaa, A.; Zhang, S.Y.; Isabel, N.; Beaulieu, J.; Bousquet, J. Phenotypic correlations between juvenile-mature wood density and
growth in black spruce. Wood Fiber Sci. 2000, 32, 61–71.
24. Koga, S.; Zhang, S.Y. Relationships between wood density and annual growth rate components in balsam fir (Abies balsamea).
Wood Fiber Sci. 2002, 34, 146–157.
25. Lindström, H. Fiber length, tracheid diameter, and latewood percentage in Norway spruce: Development from pith outwards.
Wood Fiber Sci. 1997, 29, 21–34.
26. Mvolo, C.S.; Koubaa, A.; Beaulieu, J.; Cloutier, A.; Defo, M.; Yemele, M.-C. Phenotypic correlations among growth and selected
wood properties in white spruce (Picea glauca (Moench) Voss). Forests 2019, 10, 589. [CrossRef]
27. Auty, D.; Achim, A.; Macdonald, E.; Cameron, A.D.; Gardiner, B.A. Models for predicting clearwood mechanical properties of
scots pine. Forest science 2016, 62, 403–413. [CrossRef]
28. Lenz, P.; Cloutier, A.; MacKay, J.; Beaulieu, J. Genetic control of wood properties in Picea glauca—An analysis of trends with
cambial age. Can. J. For. Res. 2010, 40, 703–715. [CrossRef]
29. Rossi, S.; Deslauriers, A.; Griçar, J.; Seo, J.W.; Rathgeber, C.B.K.; Anfodillo, T.; Morin, H.; Levanic, T.; Oven, P.; Jalkanen, R. Critical
temperatures for xylogenesis in conifers of cold climates. Glob. Ecol. Biogeogr. 2008, 17, 696–707. [CrossRef]
30. Yang, K.C. Impact of spacing on juvenile wood and mature wood properties of white spruce (Picea glauca). Taiwan J. For. Sci. 2002,
17, 13–29.
31. Yang, K.C.; Hazenberg, G. Impact of spacing on tracheid length, relative density, and growth rate of juvenile wood and mature
wood in Picea mariana. Can. J. For. Res. 1994, 24, 996–1007. [CrossRef]
32. Herman, M.; Dutilleul, P.; Avella-Shaw, T. Intra-ring and inter-ring variations of tracheid length in fast-grown versus slow-grown
Norway spruces (Picea abies). IAWA J. 1998, 19, 3–23. [CrossRef]
33. Jaakkola, T.; Makinen, H.; Saren, M.P.; Saranpaa, P. Does thinning intensity affect the tracheid dimensions of Norway spruce?
Can. J. For. Res. 2005, 35, 2685–2697. [CrossRef]
34. Kang, K.Y.; Zhang, S.Y.; Mansfield, S.D. The effects of initial spacing on wood density, fibre and pulp properties in jack pine
(Pinus banksiana Lamb.). Holzforschung 2004, 58, 455–463. [CrossRef]
35. Corriveau, A.; Beaulieu, J.; Mothe, F. Wood density of natural white spruce populations in Quebec. Can. J. For. Res. 1987, 17,
675–682. [CrossRef]
Forests 2022, 13, 1807 16 of 16

36. Dutilleul, P.; Herman, M.; Avella-Shaw, T. Growth rate effects on correlations among ring width, wood density, and mean tracheid
length in Norway spruce (Picea abies). Can. J. For. Res. 1998, 28, 56–68. [CrossRef]
37. Fu, S.; Chen, H.Y.H.; Bell, F.W.; Sharma, M.; Delaney, J.R.; Peterson, G. Effects of timing of glyphosate application on jack pine,
black spruce, and white spruce plantations in northern Manitoba. For. Chron. 2008, 84, 37–45. [CrossRef]
38. Brändström, J. Micro and ultrastructural aspects of Norway spruce tracheids: A review. IAWA J. 2001, 22, 333–353. [CrossRef]
39. Dinwoodie, J.M. Variation in Tracheid Length in Picea Sitchensis CARR; Special Report No 16; Department of Scientific and Industrial
Research, Forest products research Laboratory: Princes Risborough, United Kingdom, 1963; p. 60.
40. Briggs, D.G.; Smith, W.R. Effect of silvicultural practices on wood properties of conifers: A review. In Douglas Fir: Stand
Management for the Future; Oliver, C., Hanley, D., Johnson, J., Eds.; University of Washington Press: Seattle, WA, USA, 1986;
pp. 108–117.
41. Jaakkola, T.; Makinen, H.; Saranpaa, P. Wood density in Norway spruce: Changes with thinning intensity and tree age. Can. J. For.
Res. 2005, 35, 1767–1778. [CrossRef]
42. Pape, R. Influence of thinning and tree diameter class on the development of basic density and annual ring width in Picea abies.
Scand. J. For. Res. 1999, 14, 27–37. [CrossRef]
43. Erickson, H.D.; Harrison, A.T. Douglas-fir wood quality studies part I: Effects of age and stimulated growth on wood density and
anatomy. Wood Sci. Technol. 1974, 8, 207–226. [CrossRef]
44. Vincent, M.; Krause, C.; Koubaa, A. Variation in black spruce (Picea mariana (Mill.) BSP) wood quality after thinning. Ann. For. Sci.
2011, 68, 1115–1125. [CrossRef]
45. Savidge, R. Intrinsic regulation of cambial growth. J. Plant Growth Regul. 2001, 20, 52–77. [CrossRef]
46. Larson, P.R. A biological approach to wood quality. Tappi 1962, 45, 443–448.
47. Echols, R.M. Effects of growing space on wood specific gravity in loblolly pine. In Proceedings of the Society of American Foresters;
Society of American Foresters: Washington, DC, USA, 1960; pp. 140–143.
48. Geyer, W.A.; Gilmore, A. Effect of Spacing on Wood Specific Gravity in Loblolly Pine in Southern Illinois; Department of Forestry,
Agricultural Experiment Station, University of Illinois: Champaign, IL, USA, 1965.
49. Panshin, A.J.; de Zeuuw, C. Textbook of Wood Technology: Structure, Identification, Properties, and Uses of the Commercial Woods of the
United States and Canada, 4th ed.; McGraw-Hill Book Co.,: New York, NY, USA, 1980; p. 722.
50. Nienstaedt, H.; Zasada, J.C. White spruce. In Sylvics—Volume 1: Conifers; Burns, R.M., Honkala, B.H., Eds.; USDA, forest service:
Washington, DC, USA, 1990; Agriculture Handbook 654; pp. 389–442.
51. Little, E.L., Jr. Checklist of United States Trees (Native and Naturalized); United States, Department of Agriculture, Forest Service:
Washington, DC, USA, 1979; Agriculture Handbook, 541; p. 375.
52. Franklin, G.L. Preparation of thin sections of synthetic resins and wood-resin composites, and a new macerating method for
wood. Nature 1945, 155, 51. [CrossRef]
53. Pinheiro, J.; Bates, D.; DebRoy, S.; Sarkar, D.; R Core Team. Nlme: Linear and Nonlinear Mixed Effects Models. 2016. Available
online: https://svn.r-project.org/R-packages/trunk/nlme/ (accessed on 20 April 2016).
54. RStudio Team. RStudio: Integrated Development for R; RStudio, Inc.: Boston, MA, USA, 2020.
55. R Core Team. R: A Language and Environment for Statistical Computing. Available online: http://www.R-project.org/ (accessed
on 30 August 2022).
56. Ikonen, V.P.; Peltola, H.; Wilhelmsson, L.; Kilpeläinen, A.; Väisänen, H.; Nuutinen, T.; Kellomäki, S. Modelling the distribution of
wood properties along the stems of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as affected by
silvicultural management. For. Ecol. Manag. 2008, 256, 1356–1371. [CrossRef]
57. Mäkinen, H.; Hynynen, J. Predicting wood and tracheid properties of Scots pine. For. Ecol. Manag. 2012, 279, 11–20. [CrossRef]
58. Mäkinen, H.; Jaakkola, T.; Piispanen, R.; Saranpaa, P. Predicting wood and tracheid properties of Norway spruce. For. Ecol.
Manag. 2007, 241, 175–188. [CrossRef]
59. Fabris, S.P. Influence of Cambial Ageing, Initial Spacing, Stem Taper and Growth Rate on the Wood Quality of Three Coastal
Conifers. PhD Thesis, UBC Theses and Dissertations. The University of British Columbia, Vancouver, BC, Canada, 2000.
60. Sanio, K. Über die Grösse der Holzzellen bei der gemeinen Kiefer (Pinus sylvestris), Jahrb. Wiss. Bot. 1872, 8, 401–420.
61. Zhang, S.Y. Effect of age on the variation, correlations and inheritance of selected wood characteristics in black spruce
(Picea mariana). Wood Sci. Technol. 1998, 32, 197–204.
62. Bannan, M.W. Anticlinal divisions and cell length in conifer cambium. For. Prod. J. 1967, 17, 63–69.