Article
Growth and Morphological Patterns of Norway Spruce
(Picea abies (L.) Karst.) Juveniles in Response to
Light Intensities
Pavel Bednář 1, * , Jiří Souček 1 , Jan Krejza 2,3
Citation: Bednář, P.; Souček, J.;
Krejza, J.; Černý, J. Growth and
Morphological Patterns of Norway
Spruce (Picea abies (L.) Karst.)
Juveniles in Response to Light
Intensities. Forests 2022, 13, 1804.
https://doi.org/10.3390/f13111804
Academic Editor: Alessio
Giovannelli
Received: 26 August 2022
Accepted: 21 October 2022
Published: 29 October 2022
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Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
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Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Forests 2022, 13, 1804. https://doi.org/10.3390/f13111804
and Jakub Černý 1
1 Research Station of Silviculture in Opočno, Forestry and Game Management Research Institute, Na Olivě 550,
517 73 Opočno, Czech Republic
2 Department of Forest Ecology, Faculty of Forestry and Wood Technology, Mendel University in Brno,
Zemědělská 3, 613 00 Brno, Czech Republic
3 Global Change Research Institute of the Czech Academy of Sciences, Bělidla 4a, 603 00 Brno, Czech Republic
* Correspondence: pavelbednar13@seznam.cz; Tel.: +420-731-188159
Abstract: (1) Background: The growth and morphology of Norway spruce (Picea abies (L.) Karst.)
juveniles were observed under different light conditions due to overstory canopy openness. The aim
was to investigate the photo-morphological responses of juveniles for the development of a desirable
forest structure. The research was located in a higher altitude zone in central Europe. (2) Methods:
Light was estimated using hemispherical photographs. Eight different parameters of natural regen-
eration were measured on individuals within 1 × 1 m around each point on a 3 × 3 m grid. A total
of 1214 sapling measurements (from 10 to 431 cm in height) were taken. (3) Results: Light affected
the growth and morphological development of juveniles, resulting in variation in the lateral crown
growth and crown length. Acclimations manifested as trade-offs between height and lateral crown
growth. Similar shifts were found within relative height growth (the terminal length to the total
height) and apical dominance ratio (the terminal length to the branch length). The crown length was
proven to be highly capable to acclimation. Light influenced the density weakly, however, when
a regeneration index (density × median height) was considered, significant relations with light
conditions were discovered.
Keywords: Norway spruce; photo-morphological plasticity; light intensities; conifer juveniles; natural
regeneration; growth acclimation
1. Introduction
Forest regeneration and establishment are critical stages during which considerable
losses in species diversity can often occur [1]. Natural regeneration (or natural regeneration
in conjunction with artificial regeneration) often results in a large initial seedling bank.
This is favored by a high proportion of unoccupied growing space and a diversity of
microsite characteristics. As the plants increase in size and fill in the available space,
competition becomes more intense and the growing conditions become more homogeneous.
As the initial seedling bank progressively declines, species diversity may dramatically
decrease as less competitive species are eliminated [2]. These competitive relationships
must be considered in silvicultural techniques [3–5]. When the transformation of pure
Norway spruce (Picea abies (L.) Karst.) stands is adopted, natural regeneration of Norway
spruce (combined with planting of reintroduced tree species) must be controlled to prevent
reintroduced species from being overtopped and out-competed [6]. One of the main
silvicultural tools for this purpose is adjusting light intensities through overstory canopy
openness. Thus, forest structure and heterogeneity are encouraged [7,8].
Uneven-aged forest management is characterized by silvicultural practices that create
or maintain stands made up of trees of different age classes [9,10]. Different terms and
https://www.mdpi.com/journal/forests
Forests 2022, 13, 1804 2 of 20

definitions are used for the numerous forms of uneven-aged silviculture [10,11]; however,
all these silvicultural systems are based on the creation of canopy openings or gaps to
allow light to reach the floor of the stand [5,7,8]. The degree of growth heterogeneity, and
structure of the understory induced by the variability of light levels, can be as variable
and advanced as the shade tolerance of particular tree species is high [12,13]. In similar
studies of forest ecology where light is an independent variable, it is important to consider
reference levels of other site conditions, such as nutrients and water supply. This is because
shade tolerance is shifted by the availability or scarcity of other site factors [14–16]. It is
therefore advantageous to perform such an investigation on nutrient-poor sites as a basic
level of nutrient resources in combination with average water supply, which can then
be used as a reference compared to more optimal conditions (such as nutrient-enriched
conditions) where shade tolerance is generally increased [14,15].
Tree species with different levels of shade tolerance show different levels of both
morphological and physiological plasticity under various light conditions [4,8,17]. The
response of crown morphology and height growth depending on shade tolerance or intol-
erance has been described in different studies [3,18,19]. In general, the conclusion of those
studies is that shade-tolerant tree species can adapt their crown morphology more acutely
and in more morphological traits (such as crown length, width, and projection or terminal
length) along the light gradient compared to shade-intolerant tree species. Shade-tolerant
tree species can effectively reduce their growth and survive under low light intensities due
to their diminished growth rate and photo-morphological responses resulting in lower
mortality rate in shaded conditions; after their release, they react immediately to increased
light intensities with an increased growth rate [15,20,21]. However, they do not reach the
same growth rate under canopy-open conditions as shade-intolerant tree species [22].
The three following objects of interest related to photo-morphological patterns of
Norway spruce juveniles were analyzed in this paper:
1. The existence and intensity of morphological plasticity (i.e., the ability to respond to
the different availability of resources by acclimation of the morphological traits) as a
response to different light intensities;
2. The definition of microsites where the morphological plasticity was realized most distinctly;
3. The influence of light intensities on the general development of Norway spruce
juveniles (including, for instance, their density) and a consideration of all observed
patterns for further use in silvicultural guidelines.

2. Materials and Methods

2.1. Research Sites
The research was conducted on acidic sites, i.e., Fageta piceoso-abietina or Fageta abietina-
piceosa, due to the fact that a large proportion of these sites were occupied by secondary
Norway spruce monocultures [23]. Selected research plots were represented by forest
stands with a pure even-aged Norway spruce overstory that was tended by thinnings from
below in young and mid-aged growing stages. Then in the growing stage of forest maturity,
shelter-wood cuts of different intensities or gaps were created 15–25 years ago with no
further silvicultural interventions since that time. The observations took place in the central
part of the Czech Republic in the Bohemian–Moravian Highlands within altitudes of ca.
670–770 m above sea level (Figure 1). The soils of the research plots were determined as
cambisols (mainly Eutric Cambisol; less often Dystric Cambisol or Areno-dystric Cambisol).
The whole area lies in the geological entity of Moldanubikum with the typically prevailing
acidic bedrocks of mainly amphibolite with veins of granodiorite or granite [24]. The mean
annual air temperature is 4.9 ◦ C; the mean annual precipitation is 850 mm with 55%–60%
falling in the growing season [24,25], and the number of days when the snow cover is
present usually ranges between 100 and 120 days [26].
Forests 2022, 13, x FOR PEER REVIEW 3 of 21

Forests 2022, 13, 1804 3 of 20

Figure 1. Location of research plots for applied measurements showing the positions both within
FigureEurope
1. Location of research
and the plots for applied measurements showing the positions both within
Czech Republic.
Europe and the Czech Republic.
2.2. Experiment Design
2.2. Experiment
Three Design
research plots were established for this study (Table 1), taking into account
experimental
Three researchconditions
plots wereasestablished
follows: for this study (Table 1), taking into account
experimental
• conditions
Overstory, as follows:
consisting of pure mature even-aged Norway spruce stands (approximately
• 100 years
Overstory, old); of pure mature even-aged Norway spruce stands (approxi-
consisting
•
mately A100
gradient of overstory canopy openness characterized by a significant gradient of
years old);
• light intensity
A gradient of overstory within the plots
canopy ranging
openness from complete
characterized by acanopy closure
significant up to open
gradient of sky
conditions of a forest gap; the maximum size of a gap in the
light intensity within the plots ranging from complete canopy closure up to open sky study was considered
as ca.of16
conditions m in diameter
a forest (i.e., ca. 2/3
gap; the maximum sizeof
ofoverstory
a gap in themature
study tree
was height) and as
considered thus ca.
200–220 m 2 (ca. 2 ares) of the total size; thus, the light conditions ranged from 0.04 to
ca. 16 m in diameter (i.e., ca. 2/3 of overstory mature tree height) and thus ca. 200–
220 m20.46(ca.of the total
2 ares) sitetotal
of the factor (TSF);
size; thus, the light conditions ranged from 0.04 to 0.46
of• the No
totalfurther silvicultural
site factor (TSF); interventions were carried out within natural regeneration
• (once silvicultural
No further it started) orinterventions
within the mature overstory
were carried once the
out within initialregeneration
natural regeneration cut
(shelter-wood cut or gaps) occurred.
(once it started) or within the mature overstory once the initial regeneration cut (shel-
ter-wood cut or gaps) occurred.
Table 1. Fundamental characteristics of the research plots.

Table 1. Fundamental characteristics

Elevation of the research plots. Area
Plot Name (ID) Geographical Coordinates Dimension (m) Number of Sampled Sub-Plots
(m a.s.l.) (m2 )
Geographical Elevation Area Number of Sampled Sub-
Plot Name (ID)
Drtič (A) 49◦ 36 0 5000 N; 15◦ 530 0500 E 672a.s.l.) Dimension
27 × 42 (m) 1134
Coordinates (m (m2) Plots126
Drtič (A)
Tornádová 49◦ 40
(B) 49°36′ 0 1200 N; 15◦ 550 5600 E
50″ N; 15°53′ 05″ E 672
737 12 ×
2721
× 42 1134
252 126 28
Tornádová (B)
Odvozní (C) 49°49 40 1700N;N;15°55′
◦
40′ 0
12″ 15◦ 56056″
1300 E 737
766 1212× 21
9× 252
108 28 12
Odvozní (C) 49° 40′ 17″ N; 15°56′ 13″ E 766 9 × 12 108 12
Field-Map technology (IFER, Prague, Czech Republic) was utilized to identify the position
Field-Map technology (IFER, Prague, Czech Republic) was utilized to identify the
of the mature overstory for both tree positions and their crown projections and to locate borders
position of the mature overstory for both tree positions and their crown projections and
of research plots. All present mature trees within the plot areas and all neighbors whose crown
to locate borders of research plots. All present mature trees within the plot areas and all
projections intervened in the area of the plot were included. Afterward, the same equipment
neighbors whose crown projections intervened in the area of the plot were included. Af-
was used to cover the research plots in a grid of 3 m × 3 m squares. Every segment (3 m × 3 m)
terward, thegrid
of the samewasequipment wasmarked
permanently used toatcover the research
its central plots in(Figure
point (centroid) a grid S1).
of 3 Centroids
m × 3 m were
squares. Every segment (3 m × 3 m) of the grid was permanently marked
then targeted for subsequent measurements when hemispherical photos were taken at its central
here and
point (centroid) (Figure S1). Centroids were then targeted for subsequent measurements
the number of Norway spruce juveniles was measured within 1 m × 1 m sub-plots around
when the
hemispherical
centroids. photos were taken here and the number of Norway spruce juveniles
was measured within 1 m × 1 m sub-plots around the centroids.
2.3. Natural Regeneration Measurements
2.3. NaturalAround
Regeneration Measurements
each centroid (within the 1 × 1 m sub-plot), the number of all present Norway
spruce individuals was counted (individuals both below and over 10 cm in height were
Forests 2022, 13, 1804 4 of 20

included), and the morphological parameters were measured for all seedlings and saplings
over 10 cm in height (according to [3]).
Seven different morphological parameters of Norway spruce juveniles were measured,
as listed in Table 2. The chosen morphological parameters were employed in studies of the
photo-morphological plasticity of different tree species in the past [27–29].

Table 2. Parameters of Norway spruce natural regeneration that were measured within the study.

Parameter Acronym Juveniles Involved

Total density DEN all
Total height H (cm) above 10 cm
Live crown length Cl (cm) above 10 cm
Terminal leader length Tl (cm) above 10 cm
Length of the longest branch Bl (cm) above 10 cm
Width of crown in the widest part of crown CW1 (cm) above 10 cm
Width of crown perpendicular to the widest part of crown CW2 (cm) above 10 cm
Diameter at height of 5 cm above ground D (mm) above 10 cm

Stem diameter was taken as an average from two mutually perpendicular measure-
ments, and it was measured using a digital caliper with an accuracy of 0.1 cm. It was
measured 5 cm above the ground, due to the possible occurrence of morphological anoma-
lies within the root collar [30,31]. Other morphological parameters were measured using a
telescopic tape measure or a Messfix telescopic measuring stick (NEDO, Switzerland) with
an attached level and a precision of 1 cm. In total, 1214 juveniles of Norway spruce were
measured with heights ranging from 10 to 431 cm within 166 sub-plots on three different
research plots.

2.4. Indirect Estimation of Light Intensities

We used color hemispherical photography evaluation as an indirect method to estimate
the potential light intensity [5,32]. This method is also fully compatible with instantaneous
direct methods, such as photon flux density measurement [5,33]. A self-leveling platform
(Régent Instruments Inc., Quebec City, QC, Canada) with a remote-triggered Nikon Coolpix
995 digital camera (Nikon, Tokyo, Japan) with a fine-quality, calibrated fish-eye lens was
employed. The whole self-leveling platform was fixed atop a Manfrotto 269 HDBU tripod
(Manfrotto, Cassola, Italy) with a maximum height of 8 m; this enabled photos to be taken
from 1.7 m (the essential height of the tripod) to 4.5 m for photographing the tallest juveniles.
Photographs were taken during windless weather and standard overcast sky conditions—
i.e., when the solar disk was completely obscured (according to Čater et al. 2013 [5]). The
camera was leveled and the fish-eye lens oriented toward magnetic north using a compass.
The standardized post-processing and evaluation of the photos were performed with
WinSCANOPY 2012 (Régent Instruments Inc., Quebec City, QC, Canada). The quality of
photos was controlled visually, and batches (of maximally 20) of similar photos (mainly
based on features of exposure and contrast) were created. Then the process of distinguishing
sky vs. canopy was performed by selecting different colors of a picture and sorting them
into one of these categories. Finally, black and white thresholding was tested for several
random pictures of the batch to check the accuracy of the thresholding visually. The value
of 60◦ as a zenith angle detecting an evaluated sector of the picture view was used as
the most accurate and effective [5]. The growing season was considered from 1 May to
30 September, according to Löf et al. (2007) [21] and Čater et al. (2013) [5]. Other required
values, such as the geographical coordinates, the elevation, and the magnetic declination,
were considered. Five different characteristics (two classifying the openness of the canopy
Forests 2022, 13, 1804 5 of 20

and three classifying light intensities (Table 3)) were taken into account for subsequent
analysis of this study according to the manual of WinSCANOPY software [34].

Table 3. Characteristics of light intensities and canopy openness based on hemispherical photography
evaluation used in this study.

Entity Acronym Description Open Area Conditions

Gap Fraction GFr pixels of “Sky” in two dimensions 100%
Openness Op pixels of “Sky” in three dimensions 100%
Direct Site Factor DSF sum of direct light of a growing season 1
Indirect Site Factor ISF sum of indirect light of a growing season 1
Total Site Factor TSF sum of total light of a growing season 1

2.5. Data Evaluation

2.5.1. The Effect of Light Intensities on the Density of Natural Regeneration
First, all individuals of all sub-plots in the research plots were considered for the
density evaluation. Then, to determine if there was a relationship between light intensity
and density along the dimension of natural regression (i.e., height), height classes deter-
mined by the tallest juvenile within a sub-plot were created and only those sub-plots were
subsequently tested. The height classes were as follows: up to 15 cm, 20 cm, 30 cm, 40 cm,
50 cm, 60 cm, 70 cm, 80 cm, 90 cm, 100 cm, 110 cm, 120 cm, 130 cm, and 140 cm.

2.5.2. Morphological Patterns of Juveniles under Size-Independent Evaluation

The influence of light intensities on two measured parameters (H and D (Table 2))
and six calculated indexes (Table 4) was analyzed. The size (i.e., height classes) was not
considered here; the exact value of a parameter or index for a sub-plot was calculated as
the median of all present juveniles occurring within a particular sub-plot, according to
Dai (1996) [35]. However, to make this size-independent evaluation useful for individuals
of varying heights, parameters and indexes involving the parameters of total height (H) or
terminal length (Tl) were used. For calculation of the regeneration index (RI), the number of
juveniles higher than 10 cm was considered as the total number of juveniles in the equation
for which the height was being measured.

2.5.3. Morphological Patterns of Juveniles under Size-Dependent Evaluation

Natural regeneration was divided into height classes according to Lundquist and
Fridman (1996) [36], Duchesneau et al. (2001) [37], and Chrimes and Nilson (2005) [19]. The
regular height classes concerning a range of heights in this study were as follows: 10–50 cm,
51–100 cm, 101–150 cm, and >151 cm.
The exact value of a parameter or an index of a height class was calculated within a
sub-plot as a median of values of all juveniles falling into a particular height class. Thus,
when more height classes were present within one certain sub-plot, more values of the
same parameter (or index) were obtained, but for different height classes (Table 4). In
total, for size-dependent evaluation, the influence of light intensities on all seven measured
morphological parameters (Table 2) and eight calculated indexes was tested (Table 4).

2.6. Statistical Analysis

All statistical analyses were performed with p < 0.05. William’s test was employed
using QC Expert version 3.3 (Trilobyte Statistica Software Ltd., Pardubice, Czech Republic)
to detect outlying values. The software STATISTICA version 10.0 (StatSoft Inc., Tulsa, OK,
USA) was used to test the significance of models of light intensities and morphological
features; quadratic polynomials were used to intersperse the values. When a model was
found to be significant, the Spearman correlation coefficient (rs ) was tested (to test the
Forests 2022, 13, 1804 6 of 20

robustness of correlation against the effect of extreme values, nonlinear correlation, and non-
normality). For the visualization of certain indexes within the largest research plot, Wafer
plots were constructed in STATISTICA software, and then those outputs were overlaid on
the schema of the plot with 50% transparency in a freeware piece of graphics software.
Redundancy analysis (RDA) was performed to investigate the effect of the main light
components (direct and indirect—DSF and ISF) and the similarities within intensities
of photo-morphological plasticity of different observed parameters and indexes. In this
analysis, all measured parameters were included, as were indexes calculated for size-
independent evaluation (i.e., regardless of height classes); the density was also included.
RDA was performed with R software version 3.6.0 [38] in R Studio version 1.2.1335 [39]
using the R package vegan [40].

Table 4. Photo-morphological indexes calculated for Norway spruce juveniles considered within two
different approaches of data evaluation.

Indexes Used for Testing of Photo-Morphology of Norway Spruce Juveniles under Size-Independent Evaluation *
Title of Index Acronym Equation
Relative height growth RHG Tl/H
Apical dominance ratio ADR Tl/Bl
Relative crown length RCL Cl/H
Relative crown width RCW (CW1 + CW2)/2 × H
Height–diameter ratio HDR H/D
Regeneration index ** RI (cm/m2 ) juveniles’ number * median height
Indexes used for testing of photo-morphology of Norway spruce juveniles under size-dependent evaluation ***
Title of index Acronym Equation
Height–diameter ratio HDR H/D
Relative crown length RCL Cl/H
Relative crown width RCW (CW1 + CW2)/2 × H
Crown width–length ratio CWLR (CW1 + CW2)/2 × Cl
Crown projection CP (cm2 ) ((CW1 + CW2)/4)2 × π
Index of spatial distribution ISD (CW1 + CW2)/2 × D
Crown volume according to equilateral cone CVEC (cm3 ) π/12 × (CW1 + CW2)2 /4 × Cl
Crown volume according to quadratic paraboloid CVQP (cm3 ) π/8 × (CW1 + CW2)2 /4 × Cl
H: total height (cm); Cl: live crown length (cm); Tl: terminal leader length (cm); Bl: length of the longest branch
(cm); CW1: width of crown in the widest part of crown (cm); CW2: width of crown perpendicular to the widest part
of crown (cm); D: diameter at height of 5 cm above ground (mm); * Indexes were calculated according to Messier
et al. (1999) [27], Duchesneau et al. (2001) [37], Robakowski et al. (2004) [41], Grassi and Giannini (2005) [18], and
Szymura (2005) [29]. ** The number of individuals in this equation was considered as the total number of juveniles that
were higher than 10 cm for which the total height (H) was measured and then used to calculate a median of heights.
*** Indexes were calculated according to Šebík and Polák (1990) [42], Takahashi (1996) [43], Messier et al. (1999) [27],
Claveau et al. (2002) [44], Duchesneau et al. (2001) [37], Grassi and Giannini (2005) [18], and Szymura (2005) [29].

3. Results
3.1. The Effect of Light Intensities on the Regeneration Density
Only a weak relationship between light intensities and the density of Norway spruce
natural regeneration was found. No significant effect was demonstrated when gap fraction
(GFr), openness (Op), or indirect site factor (ISF) was considered through all tested classes
of the maximal height. However, when direct site factor (DSF) or total site factor (TSF) was
contemplated, a few significant relations were shown. For both light variables, significant
(p < 0.05) relations were found when height classes up to 30, 40, 50, and 70 cm were
investigated. Significance was also proven for DSF when height classes up to 60 and 100 cm
Forests 2022, 13, 1804 7 of 20

were tested (Table A1 in Appendix A). However, none of those significant relationships
were established by the significance of the Spearman correlation coefficient (rs ). In addition,
the significance of relations between DSF and the density of Norway spruce up to 80
and 90 cm was narrowly rejected with p-values of 0.0529 and 0.0562, respectively. All
significant models showed a gentle decrease in density with increasing light intensities (i.e.,
negative correlation). In general, the effect of light conditions on the density of Norway
spruce juveniles was found to be very weak and limited; when a relationship was found, it
occurred only for juveniles up to 1 m in height.

3.2. General Morphological Patterns of Juveniles under Size-Independent Evaluation

3.2.1. Height Growth
A significant relationship between light intensities and relative height was established
for all five light or canopy openness variables. Moreover, all of the discovered models
were highly significant (p < 0.001), and their significance was proven to be robust against
negative influences by the significance of the Spearman correlation coefficient (Table 5).
Relative height growth (RHG) increased moderately with increasing low light intensities;
the increment became progressive with approximately 20% of the light intensity of open
area conditions. This means the terminal length related to total height was higher in
high light intensities. A comparison of RHG and the apical dominance ratio (ADR) is
stated below (see Section 3.2.2) and visualized in Figure 2A. Parameter H was not found to
significantly correlate with light intensities.

Table 5. Significantly proven regression models of light intensities and photo-morphological acclima-
tions of Norway spruce natural regeneration under size-independent evaluation.

Index Regression Model Variable (x) R2 p rs

RHG = 0.1904 − 0.0151 × x + 0.0005 × x2 GFr 0.26 0.0000 *** 0.3698 *
RHG = 0.1911 − 0.015 × x + 0.0005 × x2 Op 0.26 0.0000 *** 0.3577 *
RHG RHG = 0.1104 − 0.1592 × x + 0.8724 × x2 DSF 0.10 0.0004 *** 0.1949 *
RHG = 0.1813 − 1.1899 × x + 3.5281 × x2 ISF 0.25 0.0000 *** 0.2878 *
RHG = 0.1105 − 0.1999 × x + 1.0346 × x2 TSF 0.12 0.0002 *** 0.2097 *
ADR = 0.3433 − 0.0285 × x + 0.001 × x2 GFr 0.24 0.0000 *** 0.4362 *
ADR = 0.347 − 0.0287 × x + 0.001 × x2 Op 0.25 0.0000 *** 0.4218 *
ADR ADR = 0.1865 − 0.0733 × x + 1.1619 × x2 DSF 0.08 0.0015 ** 0.2257 *
ADR = 0.3402 − 2.4533 × x + 7.4787 × x2 ISF 0.25 0.0000 *** 0.3784 *
ADR = 0.179 − 0.0943 × x + 1.374 × x2 TSF 0.09 0.0006 *** 0.2442 *
RCL = 0.5603 − 0.0024 × x + 0.0002 × x2 GFr 0.12 0.0001 *** 0.4242 *
RCL = 0.559 − 0.0023 × x + 0.0002 × x2 Op 0.12 0.0001 *** 0.4301 *
RCL RCL = 0.5272 + 0.6574 × x − 0.53 × x2 DSF 0.11 0.0001 *** 0.3337 *
RCL = 0.5723 − 0.3511 × x + 2.2008 × x2 ISF 0.14 0.0000 *** 0.4395 *
RCL = 0.5113 + 0.7212 × x − 0.5476 × x2 TSF 0.12 0.0000 *** 0.3560 *
RI = 116.5526 − 10.4147 × x + 0.798 × x2 GFr 0.09 0.0006 *** 0.2430 *
RI RI = 126.1057 − 12.5477 × x + 0.8617 × x2 Op 0.10 0.0003 *** 0.2568 *
RI = 215.7408 − 2436.291 × x + 10498.1164 × x2 ISF 0.16 0.0000 *** 0.3567 *
Note: RHG: relative height growth; ADR: apical dominance ratio; RCL: relative crown length; RI: regeneration
index (cm/m2 ); GFr: gap fraction; Op: canopy openness; DSF: direct site factor; ISF: indirect site factor; TSF: total
site factor; p < 0.05 (*), p < 0.01 (**), p < 0.001 (***).
 Forests 2022, 13, x FOR PEER REVIEW 8 of 21
Forests 2022, 13, 1804 8 of 20

Figure 2. Example of visualized similarities between relative height growth (RHG) and apical
dominance ratio (ADR)
Figure 2. Example (A) andsimilarities
of visualized example of relativerelative
between crown height
lengthgrowth
(RCL) (RHG)
(B) of Norway
and apical spruce
dom-
juveniles under a mature overstory canopy on the largest research plot Drtič (A). For all
inance ratio (ADR) (A) and example of relative crown length (RCL) (B) of Norway spruce juveniles three indexes,
aunder
deep agreen
mature overstory
color canopy
indicates highlyon the largestgrowth
suppressed researchregimes
plot Drtič (A). For all three
characterized by (i) indexes, a deep
short terminal
green (as
length color indicatesinhighly
numerator RHG suppressed
and ADR); (ii) growth
lateralregimes characterized
crown growth by (i)by
encouraged short
longterminal length
branch length
(as numerator in RHG and ADR); (ii) lateral crown growth encouraged
(as denominator in ADR); (iii) shortened crown length (as numerator in RCL). by long branch length (as
denominator in ADR); (iii) shortened crown length (as numerator in RCL).
3.2.2. Lateral Crown Growth
3.2.2.The
Lateral
apicalCrown Growthratio was significantly influenced by light intensities for all
dominance
five light
The or canopy
apical openness
dominance variables
ratio (p < 0.01). Moreover,
was significantly influencedall byfive regression
light intensitiesmodels
for all
were proven
five light or to be robust
canopy against
openness negative(pinfluences
variables by the significance
< 0.01). Moreover, of the Spearman
all five regression models
correlation
were proven coefficient (Table
to be robust 5). For
against all five influences
negative regressionby models, a positive of
the significance correlation
the Spearman was
observed when ADR was increasing with light intensity. This means
correlation coefficient (Table 5). For all five regression models, a positive correlation was the terminal length
increased
observed under
when ADR higherwaslight intensities
increasing while
with lighttheintensity.
lateral growth was slower.
This means In lowlength
the terminal light
intensities,
increased underlateralhigher
growth prevailed,
light intensitiesand the annual
while height
the lateral increment
growth was suppressed.
was slower. In low light
ADR, comprising
intensities, lateralboth branch
growth length (Bl)
prevailed, andthe
and terminal
annuallength
height(Tl) and thus was
increment expressing shifts
suppressed.
within the level of height increment and lateral growth, was
ADR, comprising both branch length (Bl) and terminal length (Tl) and thus expressingshown to be a suitable index
for observation
shifts within theoflevel
this phenomenon.
of height increment Whenand considering only lateral
lateral growth, growthtoexpressed
was shown be a suitable by
relative crown width (RCW) within such a large gradient of
index for observation of this phenomenon. When considering only lateral growth ex-juvenile dimensions (although
H was theby
pressed denominator
relative crownfor RCW),
width no significant
(RCW) withincorrelation
such a largewithgradient
light intensities
of juvenilewasdimen-
found.
sionsIn(although
addition, changes
H was the within both RHGfor
denominator andRCW),
ADR along the light gradient
no significant correlation were reason-
with light
ably similar. When
intensities was found. the ranges of median values of sub-plots were compared, RHG was
foundIntoaddition,
range within 15.2 times (the ratio of the highest median
changes within both RHG and ADR along the light gradient were rea- value of the sub-plot to
the smallest median value of the sub-plot), while the same approach
sonably similar. When the ranges of median values of sub-plots were compared, RHG was showed a range of
23.3
found times for ADR.
to range withinHowever,
15.2 times when
(theall setsofofthe
ratio juveniles
highest were
median compared,
value of thethe range
sub-plot was to
much wider and represented approximately 68 times and 131
the smallest median value of the sub-plot), while the same approach showed a range of times for RHG and ADR,
respectively. In addition, the visualization of both indexes was performed (Figure 2A) on
23.3 times for ADR. However, when all sets of juveniles were compared, the range was
the largest research plot, Drtič (A). This demonstrated a similar rate of photo-morphological
much wider and represented approximately 68 times and 131 times for RHG and ADR,
acclimations for both indexes. The same positive correlation of both ADR and RHG was
respectively. In addition, the visualization of both indexes was performed (Figure 2A) on
also confirmed by RDA, which showed a similar level of effect of both DSF and ISF (i.e.,
the largest research plot, Drtič (A). This demonstrated a similar rate of photo-morpholog-
direct and indirect components of light (Figure 3)).
ical acclimations for both indexes. The same positive correlation of both ADR and RHG
was also confirmed by RDA, which showed a similar level of effect of both DSF and ISF
(i.e., direct and indirect components of light (Figure 3)).
ForestsForests
2022, 13, x FOR
2022, PEER REVIEW
13, 1804 9 of 209 of 20

FigureFigure
3. The3.results of redundancy
The results analysis
of redundancy (RDA)(RDA)
analysis wherewhere
two factors were involved—indirect
two factors site site
were involved—indirect
factorfactor
(ISF) and direct site factor (DSF).
(ISF) and direct site factor (DSF).

3.2.3.3.2.3. Relative
Relative Crown Crown
Length Length
A significant
A significant positive
positive correlation
correlation between
between light light intensities
intensities and relative
and relative crown crown
lengthlength
(RCL)(RCL) was established
was established for allfor all light
five five light or canopy
or canopy openness
openness variables.
variables. All ofAlltheofmodels
the models
werewere
highlyhighly significant
significant (p < 0.001),
(p < 0.001), and their
and their significance
significance was proven
was proven to be to be robust
robust against
against
negative influences by the significance of the Spearman correlation coefficient (Table 5). 5).
negative influences by the significance of the Spearman correlation coefficient (Table
RCL increased
RCL increased with with
light light intensities,
intensities, whichwhich
proved proved the ability
the ability of Norway
of Norway spruce spruce juveniles
juveniles
to reduce crown length under low light intensity. The range
to reduce crown length under low light intensity. The range of photo-plasticity of this of photo-plasticity of this
index was lower compared to both RHG and ADR when within
index was lower compared to both RHG and ADR when within the comparison of median the comparison of median
valuesvalues of sub-plots
of sub-plots (the highest
(the highest median median
valuevalue of sub-plot/the
of sub-plot/the smallest
smallest median median
valuevalue
of of
sub-plot); the range was a multiple of 4.3 (when the whole
sub-plot); the range was a multiple of 4.3 (when the whole set of juveniles was set of juveniles was investigated,
the range
investigated, was
the a multiple
range of approximately
was a multiple 6.3). The
of approximately variation
6.3). of RCLofwithin
The variation the largest
RCL within
the largest research plot (A) is shown in Figure 2B. Within the RDA analysis, huge were
research plot (A) is shown in Figure 2B. Within the RDA analysis, huge similarities
found between the effects of DSF and ISF on RCL, RHG, and ADR (Figure 3).
similarities were found between the effects of DSF and ISF on RCL, RHG, and ADR
(Figure 3). Regeneration Index—RI
3.2.4.
A significant
3.2.4. Regeneration positive correlation between the regeneration index (RI; RI = the number
Index—RI
of individuals × median of the height) and three variables (GFr, Op, ISF) was established.
A significant
For those threepositive correlation
regression models, between
a highthelevelregeneration index
of significance (RI;found
was RI = the(p number
< 0.001), and
of individuals × median of the height) and three variables (GFr, Op, ISF) was
the significance of the Spearman correlation coefficient was shown for all three (Table 5). established.
For those three when
However, regression models,
both DSF andaTSF high level
were of significance
tested, was influence
no significant found (p <was 0.001), and
revealed.
the significance of the Spearman correlation coefficient was shown for all
This means that if dimension was not considered, the density of natural regeneration three (Table 5).
However,
weakly when both DSF
correlated andlight
with TSFconditions
were tested, no significant
within a limitedinfluence
range of was revealed.
its height (maximally
This means that if dimension was not considered, the density of natural
up to 1 m (see Section 3.1)). However, when RI was tested, significance was proven. In regeneration
weakly correlated
addition, the with
commonlight conditions within a2limited
density (pieces/m ) negativelyrangecorrelated
of its height with(maximally up
light conditions
to 1 (albeit
m (seepoorly),
Sectionwhile3.1)). the
However, when RI was tested,
2 significance was
regeneration index (cm/m ) correlated positively. This shows proven. In the
addition, the common density (pieces/m 2) negatively correlated with light conditions
fundamental importance of regeneration height within the regeneration index. The total
(albeit poorly),
height (andwhile
otherthe regeneration
parameters index (cm/m
or indexes 2) correlated positively. This shows the
expressing the height growth) correlated positively
fundamental
with lightimportance
in this study of as
regeneration height within
well. Furthermore, based on thethe
regeneration
RDA, a positiveindex.correlation
The total of RI
height (and
with lightother parameters
intensity can also or indexes expressing
be concluded (Figure 3). the height growth) correlated
positively with light in this study as well. Furthermore, based on the RDA, a positive
3.2.5. Stem
correlation of RI Diameter
with light intensity can also be concluded (Figure 3).
Statistical analyses did not prove the effect of light intensities on the thickness of the
3.2.5.stem
Stemwhen
Diameter
diameter (D) did not correlate with light intensities. Moreover, no significant
influence of light intensities
Statistical analyses did not provewas the
found onof
effect the height–diameter
light intensities on the ratio (HDR) where
thickness of the D is
included as a denominator (when H alone was also not found
stem when diameter (D) did not correlate with light intensities. Moreover, no significant to significantly correlate
with light intensities in this general approach of size-independent
influence of light intensities was found on the height–diameter ratio (HDR) where D is evaluation).
Forests 2022, 13, 1804 10 of 20

3.3. Morphological Patterns of Juveniles under Size-Dependent Evaluation

3.3.1. Height Growth
Terminal length (Tl) and total height (H) were tested, and significant regression models
for both were established with a positive correlation between height growth and light
intensities. The Tl was more dependent on light intensities; it was significantly influenced
by light in three height classes (10–50 cm, 51–100 cm, >151 cm) (Table A2 in Appendix A).
By contrast, total height was significantly correlated only with light intensities in the tallest
height class (>151 cm) (Table A2 in Appendix A). A distinctive effect of light on Tl was
found within the smallest height class (10–50 cm), where it significantly correlated with
four variables of light (GFr, Op, ISF, TSF) with a high level of significance (p < 0.01).

3.3.2. Lateral Crown Growth

To observe morphological development at the level of crown lateral growth, three
parameters (branch length (Bl), crown width 1 (CW1), crown width 2 (CW2)) and three
indexes (relative crown width (RCW), crown projection (CP), index of spatial distribution
(ISD)) were tested. All of them were significantly correlated with light conditions within
observed height classes (Table A3 in Appendix A). In general, all observed parameters and
indexes (where crown width was situated as a numerator within indexes) demonstrated a
strong negative correlation with light intensities that expressed an initiation of the lateral
growth under low light and its suppression under high light. The parameters Bl and CW1
and the index CP were correlated with light intensities only in two cases each (throughout
the whole gradient of height classes), and thus they were rarely correlated with light com-
pared to other observed parameters and indexes of lateral crown growth. This significance
was established for all three in height classes 51–100 cm and 101–150 cm. For a similar
range of natural regeneration (height ranging from 51 to 150 cm), the correlation of CW2
was also proven; however, four regression models were proven for this parameter. The ISD
significantly correlated with light conditions only when the highest class of juveniles was
considered (>151 cm); however, for this height class, the correlation was proven by three
significant regression models. The highest level of correlation was demonstrated by RCW,
as eight significant regression models were proven for this index, which was correlated
with light conditions within a wide range of dimensions—for juveniles taller than 51 cm, in
particular (i.e., the height classes 51–100, 101–150, and >151 cm). This index was especially
appropriate for the tallest juveniles (height class >151 cm), where four significant regression
models were found.
These results clearly demonstrate that photo-morphological plasticity at the level of
the lateral growth of the crown has not yet developed within the smallest height class
(10–50 cm) of Norway spruce natural regeneration. In this height class, no significant
influence of light conditions on observed parameters and indexes of the lateral crown
growth was established.

3.3.3. Relative Crown Length and General Shape of the Crown

The number of established regression models that covered all observed height classes
showed the highest capability to acclimation at the level of crown length and general crown
shape. Cl and RCL were shown to be correlated with light intensities within three height
classes (10–50 cm, 51–100 cm, >151 cm) through 8 and 10 significant regression models,
respectively. The correlation of CWLR was demonstrated within all four height classes
through 16 significant regression models. Since Cl was a parameter and RCL an index
where the crown length was situated as a numerator of the index, both showed the same
trend within the light gradient. Thus, the crown length was positively correlated with
light intensities, which means it was suppressed under low light and enhanced under
high light. Compared to the photo-morphological plasticity of lateral growth that was
not found within the smallest dimension of juveniles of 10–50 cm (see Section 3.3.2), the
photo-plasticity of the crown length was shown even in this initial stage of the natural
regeneration. In particular, RCL was frequently found to be significantly affected by light
Forests 2022, 13, 1804 11 of 20

intensities (five regression models) in the smallest height class. Furthermore, three models
both for Cl and CWLR were also proven for the smallest juveniles (Table A4 in Appendix A).
CWLR, which aggregated both the lateral growth of the crown and its length, showed
particular importance—it expressed the general shape of the crown. CWLR highly cor-
related with light intensities as 16 significant regression models through all dimensions
of juveniles were proven (Table A4 in Appendix A). The lateral growth was situated as a
numerator and the crown length as a denominator within the index; CWLR demonstrated
a significantly negative correlation with light intensities (when the lateral growth alone
demonstrated the same negative correlation while the crown length exhibited a positive
correlation). This means that under low light intensities, juveniles were creating umbrella-
shaped crowns (i.e., a high value of CWLR) through an acceleration of the lateral growth
and shortening of the crown length. An exception was seen in this pattern only within the
smallest juveniles (10–50 cm) where the CWLR changes were created through crown length
acclimations. This was due to a lack of significant photo-plasticity in the lateral growth
found for the smallest juveniles (see Section 3.3.2), while the acclimations of crown length
within the smallest height class were broadly demonstrated by the significant plasticity of
both Cl and RCL.

3.3.4. Stem Diameter and Height–Diameter Ratio

A significant relation between light intensities and D within all observed height classes
was not established, fully proving previous findings of no relation of this natural regenera-
tion parameter on light (see Section 3.2). However, while no effect of light intensities on
HDR was found within general evaluation (under size-independent evaluation), these anal-
yses showed the effect of the light on HDR in three height classes (Table A5 in Appendix A).
No relationship between light and HDR was found in the smallest height class (10–50 cm),
while the most distinctive relationship was found for the tallest height class (>151 cm). This
finding clearly confirms the correlation of total height (H) and light intensities which was
shown for the same height class (>151 cm), while within other height classes the correla-
tion of the H and the light was missing and was represented (within the height growth)
through a distinctive correlation of Tl (see Section 3.3.1). In general, regardless of particular
parameters or indexes, height growth was clearly correlated with light conditions. There
was no correlation between stem diameter in this or other analyses. Thus, these proven
relations between HDR and light conditions took place through acclimations of the H
to light intensities while D remained unchanged throughout the changing light gradient.
This was confirmed by a positive correlation between HDR and light when all significant
regression models exhibited an increment in HDR with the increment in light. This shows
the same trend of correlation as with height growth.

3.3.5. Crown Volume

No effect of light intensities on the crown volume was found regardless of which
algorithm for the crown volume calculation was used for the testing (crown volume
according to equilateral cone or crown volume according to quadratic paraboloid).

4. Discussion
Other authors have established the same positive correlation between light conditions
and height growth of Norway spruce natural regeneration [19,45]. The lower correlation of
total height (H) compared to the terminal length (Tl) can be explained in that the response of
height growth to particular light conditions is not immediate. On the contrary, it takes time
from when the height growth is again in full compliance with light conditions, as has been
established by Nilson and Lundqvist (2001) [46], Robakowski et al. (2004) [41], and Chrimes
and Nilson (2005) [19]. This is in compliance with the results of Diaci and Firm (2011) [3],
who demonstrated a significant correlation between Tl and light intensity both for Norway
spruce and silver fir (Abies alba Mill.) seedlings, but did not find a significant correlation
with H. Tl has a higher probability of being adaptive to current light conditions, while H
Forests 2022, 13, 1804 12 of 20

reflects the history of height growth through changing light conditions. Although light
conditions were reasonably stable for a long time (at least for 15 years or more) in this study,
some changes did occur (due to overstory growth and overstory canopy development).
A similar situation can be concluded for the index RHG, as RHG plasticity was more
carried out through Tl changes than H changes. The parameter of Tl was also proven by
Jaloviar et al. (2013) [16] as being associated with light intensities both for Norway spruce
natural regeneration and silver fir underplantings. By contrast, they found no significant
effect of light on the RHG of either tree species [16].
A high level of photo-morphological plasticity in this study was found for lateral
crown growth. However, within the evaluation involving height classes, none of those
adaptations (at the level of parameters Bl, CW1, and CW2 or indexes RCW, CP, and ISD)
were employed in the smallest height class (10–50 cm). Similar results were established by
Duchesneau et al. (2001) [37] for balsam fir (Abies balsamea (L.) Mill.) when they found a
generally higher level of correlations of morphological plasticity and light for the height
class of 100–200 cm compared to the height class of 50–100 cm. Similar findings were
confirmed for other conifer shade-tolerant tree species [28,44]. However, a continual photo-
morphological acclimation through a gradient of the dimension of natural regeneration can
be limited, because as Wright et al. (1998) [15] found, a height of 6 m can be a threshold for
many different tree species when the described trend starts to weaken or even disappears.
It was widely established that the plasticity of crown morphology plays a key role in
the adaptation ability to survive under a wide range of light intensities, and thus this fea-
ture is uniquely distinctive for shade-tolerant tree species [12,13,47,48]. Crown morphology
has a crucial effect on light use efficiency and thus on the competitive strength of an individ-
ual [44,48–50]. This competitiveness within natural regeneration can increase if light availability
becomes limiting due to overstory shelter and can vary according to the ratio of diffuse and
direct light in understory microsites [8]. Consequently, shade-tolerant tree species have the
ability to adapt their growth just through acclimations via photo-morphological plasticity of the
crown, shoots, and needles or leaves [51,52]. Thus, shade-tolerant conifer tree species crown
morphology varies from conical under high light conditions to umbrella-shaped under low light
conditions [18,53,54]. This morphological development consists of acclimations via a reduction
in height growth and crown length [29,55] and initiation of crown lateral growth to maximize
carbon gain under shade [41,54,56]. In addition, the described pattern is aimed at reducing
self-shading within the crown [18,55,57]. As light intensities increase, height increment is initi-
ated, making the light more available for the lowest whorls, which initiates the establishment of
a continual conical-shaped crown [58]. Acclimation ability (expressed as different increment
allocation patterns resulting in different architecture) varies between tree species and plays a
key role in photosynthetic efficiency (essential under low light conditions) and thus offers a
competitive advantage to particular tree species [22,28,49,52].
Within the 10–50 cm height class, juveniles invest in vertical growth (proven significant
correlation of Tl and light) to reach the best social position through height increment
with no lateral crown growth having yet taken place. However, to optimize increment
allocation (i.e., carbon gain) and to avoid self-shading, the ability to reduce crown length is
already fully developed even in this early stage of natural regeneration (broadly proven
by significant correlations of Cl, RCL, and CWLR). As Takahashi (1996) [43] stated, the
whole process is a balance between investment into height growth to reach future profit and
investment into surviving when the current total height is preserved, as is broadly shown
in other studies [18,22,29,41]. Although the index CWLR integrates both lateral crown
growth and crown length, it significantly correlates with light. This takes place through
morphological acclimation of crown length and can be concluded from the established
plasticity of both Cl and RCL in the smallest height class when no proof of morphological
plasticity for lateral crown growth was found. The two described patterns of photo-
morphological acclimations (height growth and crown length) were also present in juveniles
taller than 51 cm when the ability to employ photo-morphological acclimations to crown
lateral growth was supplemented.
Forests 2022, 13, 1804 13 of 20

As a consequence, other studies considered both ADR and RCL as indexes for express-
ing the level of shade tolerance, and deducing the light intensities under which juvenile
shade-tolerant trees have been growing [29,41].
The crown length is considered the essential photo-morphological characteristic for
distinguishing between shade tolerance and semi-tolerance, since an absence of this accli-
mation for shade semi-tolerant tree species, such as eastern white pine (Pinus strobus L.),
has been shown [27,44,55].
Our finding of the correlation between HDR and light verified the classification of
Norway spruce as a shade-tolerant tree species because this level of plasticity is also
considered a typical ability of shade-tolerant tree species [59] and is absent in shade semi-
tolerant tree species [27,44]. Although we did not prove a significant correlation between
HDR and light conditions in the general evaluation, it was established for juveniles taller
than 51 cm when evaluated according to height classes.
Thus, only the initial stage of natural regeneration did not exhibit acclimation of HDR,
whereas a correlation of HDR with all observed light variables for the highest stratum of
natural regeneration (height class >151 cm) was established. The observed plasticity of
HDR was created through acclimations of height growth when stem thickness was not
related to light intensities. No correlation between D and light can be explained by intense
intraspecific competition within natural regeneration. To some extent, the increment is
invested into height growth to reach a favorable position in relation to neighboring trees. To
a greater extent, it is invested into adaptations of the crown to make photosynthesis under
shade as effective as possible to reach a high probability of survival [18,43,58] while invest-
ment into stem radial growth is reduced and affected by the density of juveniles. In such
conditions of dense natural regeneration, the intraspecific competition was a critical factor
when compared to the results of Coates and Burton (1999) [60], who established the signifi-
cant influence of light on D for five different conifer tree species five years after planting. It
is clear that in young planted seedlings, competition was substantially lower compared
to natural regeneration in our study. Furthermore, the crucial influence of density on D
was also described by Chrimes and Nilson (2005) [19], as well as Petritan et al. (2012) [61],
who established the significance of this finding for conifers rather than broadleaf species.
Thus, the correlation of stem radial increment and light intensities is fully revoked when
intraspecific (or interspecific) competition appears; only the pattern of height growth and its
dependency on light remains. Consequently, HDR plasticity, fully arranged through height
growth plasticity, occurs. This also confirmed the finding of the independence of D on light
intensities in the natural regeneration of other conifers, as Kučeravá et al. (2012) [62] found
the same for silver fir and Claveau et al. (2002) [44] observed the same for four conifer
species from a group of five. On the other hand, in this study, only D was tested when there
were no available data about radial increment. Radial increment can be more sensitive to
certain light conditions, as was established by Duchesneau et al. (2001) [37] for balsam fir
or Greis and Kellomäki (1981) [63] for Norway spruce.
This study established only a weak relationship between regeneration density and
light. Lunquist and Fridman (1996) [36] and Hasenauer and Kindermann (2002) [64] also
did not find a correlation between Norway spruce regeneration density and the basal area
of the overstory. Similarly, Szewczyk and Szwagrzyk (2010) [65] did not find a correlation
for Norway spruce either, while for silver fir, they found a significant correlation. Jadud’
et al. (2014) [66], who evaluated the influence of light intensity on the density of seedlings
after directseeding (using the same amount of seeds within various treatments), found
no correlation between light intensities and density for Norway spruce or silver fir. We
established that there was a fragile relationship for natural regeneration up to a maximum
of 1 m in height, which corresponds well with the findings of Roženbergar et al. (2007) [67].
They did not find a significant correlation between light and the density of silver fir and
observed a continual decrease in its density with its increasing height until a height of
110 cm when density developed asymptotically to axis x. Thus, the height of approximately
1 m can work as a general threshold when the (intraspecific) competition (resulting to
Forests 2022, 13, 1804 14 of 20

autoreduction) starts to dominate, and the influence of light (which slightly affects the
density of juveniles up to 1 m and is reflected in height growth as well) is retracted. By
contrast, the regeneration index (RI) where dimension was considered was significantly
correlated with light in this study, which was confirmed by Grassi et al. (2004) [68] both for
Norway spruce and silver fir. The importance of dimension (height) on the development
of density has been clearly demonstrated. Furthermore, the importance of microclimates
in microsites is an important factor, as shown by Kupferschmid and Bugmann (2005) [69],
Hunziker and Brang (2005) [70], and Kathke and Bruelheide (2010) [71]. More studies
focused on the complex mechanisms of various factors affecting Norway spruce natural
regeneration density should be conducted in the future.

5. Conclusions
The results of our research indicate that Norway spruce juveniles have immense
photo-morphological plasticity in lateral crown growth, crown length, and height growth.
The ranges of plasticity (expressed by the maximal and minimal values of RHG and ADR)
for height growth and crown lateral growth acclimated to light conditions to a similar
extent. However, the plasticity of lateral crown growth was absent in the smallest height
class (10–50 cm). The capability for acclimations in crown length was established robustly
throughout the whole range of natural regeneration dimensions. HDR showed a significant
positive correlation with light intensities for juveniles from 51 cm in height. This shows
the positive impact of overstory shelter on the mechanical stability of juveniles (making
the HDR value smaller due to the shade of the overstory shelter) and is a representative
example of nature’s automation (also known as biological automation) principles in tending
in young forest stands. However, this means that when the overstory canopy is intensively
opened (or even removed) by silvicultural intervention or by disturbance, the risk of
mechanical lability of Norway spruce understory will occur if no further tending is applied
in this natural regeneration.
The existence and intensity of photo-morphological acclimations firmly established
Norway spruce as a shade-tolerant species. Thus, light conditions in the forest stand
understory (which can be easily influenced by silvicultural techniques aimed at the forest
stand overstory) can effectively encourage forest stand structural heterogeneity as Norway
spruce juveniles follow light conditions with their growth. Moreover, through the use of
appropriate silviculture techniques, it is possible to take full control of the growth vigor of
Norway spruce juveniles. This can be effectively used for the transformation of Norway
spruce monocultures to uneven-aged mixed stands, once it is deemed appropriate for
Norway spruce to be one of the species in the future species composition of the forest stand.
For such silviculture practice, two main growth regimes can be generally distinguished:
suppressed and enhanced. These regimes are not distinguished by distinct thresholds but
can be found at the level of ca. 0.2 TSF. From this intensity, both RHG and ADR progressively
increase, whereas until ca. 0.15 TSF they very slightly increase with light (RCL increases
constantly with TSF within the whole range). When it is required to suppress the vigor of
Norway spruce juveniles (e.g., the target being to encourage advanced artificial regeneration
of reintroduced tree species during Norway spruce monoculture transformation), light
intensity should not exceed this threshold by too much (to support reintroduced species).
When Norway spruce is to be encouraged to reach advanced growing stages earlier, the light
intensity should be over 0.2 TSF. However, regardless of these two main growth regimes, the
growing response to different light conditions is continual after the threshold is exceeded,
and therefore the encouraged heterogeneity of the overstory canopy openness leads to
high growth diversity and also a high structure of Norway spruce regeneration. Therefore,
variability of patches of overstory canopy created by a combination of sections with a closed
canopy, mid-open canopy, and small gaps can be favorable for this purpose. Regeneration
density did not correlate well with light intensities (and if some weak trends were found,
they demonstrated a negative correlation with light). These results demonstrated that
the regeneration establishment is more an issue of appropriate forest microclimate, where
Forests 2022, 13, 1804 15 of 20

perhaps both temperature and humidity regimes play key roles and light is the further
additional variable, which indicates that a shelter-wood cut can be more favorable for
regeneration establishment than the creation of gaps. And after it reaches certain growing
stage of about 1 m in height, intraspecific competition and autoreduction start to be driving
forces of its density.

Supplementary Materials: The following supporting information can be downloaded at: https:
//www.mdpi.com/article/10.3390/f13111804/s1. Figure S1.
Author Contributions: Conceptualization, P.B.; idea, P.B. and J.S.; methodology, P.B.; investigation,
P.B.; measurements, P.B. and J.Č.; data acquisition, P.B.; data curation, P.B. and J.Č.; data process-
ing, P.B.; data analysis, P.B.; formal analysis, P.B.; interpretation of results, P.B.; validation, P.B.;
visualization P.B. and J.K.; supervision, P.B.; funding acquisition, P.B.; funding administration, P.B.;
writing—original draft, P.B.; writing—reviewing & editing, J.K., J.Č. and J.S. All authors have read
and agreed to the published version of the manuscript.
Funding: This research was funded by the Ministry of Agriculture of the Czech Republic, institutional
support MZE-RO0118, and by National Agency for Agricultural Research (NAZV)—project number
NAZV QK1810443 “Methods for minimization of damage caused by wind and snow in forests stands
in connection with climate change”.
Data Availability Statement: The data presented in this study are available on request from the
corresponding author.
Acknowledgments: We thank our practical forestry colleagues, namely František Slavíček, Jiří Bína,
and Petr Bednář, who were involved in this research activity and research plot establishment. We
thank Estelle Noyer for valuable help and Padraig O’Tuama for his language corrections and for text
comments. We thank Lena Macrie Hunt for her very valuable help with language correction and
with text improvements and comments. We thank technicians Luboš Janáček; Jan Škareda and Petr
Smutný for helping us with fieldwork. We thank the two anonymous reviewers for their valuable
reviews, comments, and recommendations.
Conflicts of Interest: The authors declare no conflict of interest. The funders had no role in the design
of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or
in the decision to publish the results.

Appendix A

Table A1. Significant regression models of Norway spruce natural regeneration density and light
intensities.

Height up
Variable (x) Equation; DEN (No. of Juveniles/m2 ) R2 p rs
to (cm)
30 DEN = 16.5597 − 71.5915 × x + 91.067 × x2 0.09 0.0334 * −0.0786 NS
40 DEN = 14.2696 − 42.8019 × x + 39.1018 × x2 0.06 0.0366 * −0.0826 NS
50 DEN = 15.5823 − 46.297 × x + 38.3294 × x2 0.07 0.0145 * −0.1196 NS
DSF
60 DEN = 12.6875 − 12.4916 × x − 25.0481 × x2 0.04 0.0464 * −0.0804 NS
70 DEN = 13.5962 − 20.4831 × x − 11.5114 × x2 0.05 0.0257 * −0.0858 NS
100 DEN = 12.4144 + 8.674 × x − 71.9446 × x2 0.04 0.0419 * −0.0830 NS
30 DEN = 16.9492 − 72.669 × x + 91.9606 × x2 0.09 0.0334 * −0.0546 NS
40 DEN = 14.3836 − 42.0798 × x + 36.6371 × x2 0.06 0.0423 * −0.0878 NS
TSF
50 DEN = 15.2769 − 41.107 × x + 26.4127 × x2 0.06 0.0193 * −0.0937 NS
70 DEN = 13.33 − 16.5955 × x − 20.0969x2 0.04 0.0367 * −0.0679 NS
Note: DEN: total density of all juveniles regardless of their height (pieces/m2 ); DSF: direct site factor; ISF: indirect
site factor; TSF: total site factor; p < 0.05 (*), p > 0.05 (NS ).
Forests 2022, 13, 1804 16 of 20

Table A2. Significantly proved regression models of light intensities and morphological parameters
expressing height growth of Norway spruce natural regeneration under size-dependent evaluation.

Height Class Regression Model Variable (x) R2 p rs

10–50 cm Tl = 3.5544 − 0.2377 × x + 0.009 × x2 GFr 0.18 0.0001 *** 0.4052 *
10–50 cm Tl = 3.4941 − 0.2304 × x + 0.0088 × x2 Op 0.18 0.0001 *** 0.4020 *
10–50 cm Tl = 3.458 − 20.8439 × x + 72.033 × x2 ISF 0.22 0.0000 *** 0.4187 *
10–50 cm Tl = 1.9048 + 5.5023 × x + 0.4773 × x2 TSF 0.06 0.0059 ** 0.1944 *
51–100 cm Tl = 9.7984 − 48.9895 × x + 151.7929 × x2 ISF 0.09 0.0275 * 0.3180 *
51–100 cm Tl = 8.1192 − 17.2793 × x + 73.168 × x2 TSF 0.08 0.0469 * 0.2650 *
>151 cm Tl = 17.7294 − 90.283 × x + 356.3486 × x2 DSF 0.48 0.0001 *** 0.5342 *
>151 cm Tl = −0.6218 + 46.0722 × x + 41.9031 × x2 ISF 0.14 0.0363 * 0.4228 *
>151 cm Tl = 21.1318 − 126.4246 × x + 439.5598 × x2 TSF 0.48 0.0001 *** 0.5471 *
>151 cm H = 20.712 + 1827.4301 × x − 3017.8096 × x2 DSF 0.21 0.0139 * 0.4374 *
>151 cm H = −68.8182 + 2527.9261 × x − 4391.4594 × x2 TSF 0.22 0.0134 * 0.4194 *
Note: Tl: terminal length (cm); H: total height (cm); GFr: gap fraction; Op: canopy openness; DSF: direct site
factor; ISF: indirect site factor; TSF: total site factor; p < 0.05 (*), p < 0.01 (**), p < 0.001 (***).

Table A3. Significantly proved regression models of light intensities and morphological param-
eters and indexes expressing lateral growth of Norway spruce natural regeneration under size-
dependent evaluation.

Height Variable
Regression Model R2 p rs
Class (x)
51–100 cm Bl = 19.2155 + 228.5272 × x − 540.4043 × x2 ISF 0.11 0.0256 * −0.3692 *
51–100 cm CW1 = 34.4871 + 408.7886 × x − 978.2429 × x2 ISF 0.13 0.0124 * −0.4541 *
51–100 cm CW2 = 76.9358 − 0.3667 × x − 0.0132 × x2 GFr 0.09 0.0266 * −0.3917 *
51–100 cm CW2 = 79.166 − 0.5299 × x − 0.01 × x2 Op 0.09 0.0235 * −0.3987 *
51–100 cm CW2 = 59.6086 + 102.3799 × x − 358.9949 × x2 ISF 0.11 0.0124 * −0.4040 *
51–100 cm RCW = 1.8688 − 0.051 × x + 0.0006 × x2 GFr 0.26 0.0001 *** −0.5712 *
51–100 cm RCW = 1.9707 − 0.0579 × x + 0.0007 × x2 Op 0.28 0.0000 *** −0.6031 *
51–100 cm RCW = 1.8489 − 4.0591 × x + 3.2983 × x2 ISF 0.37 0.0000 *** −0.6643 *
51–100 cm CP = 672.2795 + 33124.752 × x − 80280.9727 × x2 ISF 0.12 0.0182 * −0.4297 *
101–150 cm Bl = 561.0136 − 3200.0795 × x + 5035.5044 × x2 ISF 0.59 0.0193 * −0.5000 NS
101–150 cm CW1 = 640.0183 − 3323.1949 × x + 5071.6685 × x2 ISF 0.48 0.0331 * −0.5091 NS
101–150 cm CW2 = 645.9867 − 3606.0909 × x + 5750.5151 × x2 ISF 0.59 0.0251 * −0.4455 NS
101–150 cm RCW = 6.4675 − 36.6654 × x + 58.6871 × x2 ISF 0.71 0.0120 * −0.4727 NS
101–150 cm CP = 1.0569 × 105 − 6.2668 × 105 × x + 9.8709 × 105 × x2 ISF 0.61 0.0180 * −0.5000 NS
>151 cm RCW = 3.6175 − 0.2223 × x + 0.0042 × x2 Op 0.18 0.0409 * −0.5015 *
>151 cm RCW = 1.0578 − 1.698 × x + 0.336 × x2 DSF 0.55 0.0000 *** −0.7449 *
Forests 2022, 13, 1804 17 of 20

Table A3. Cont.

Height Variable
Regression Model R2 p rs
Class (x)
>151 cm RCW = 1.6269 − 4.1525 × x + 3.1664 × x2 ISF 0.29 0.0014 ** −0.5806 *
>151 cm RCW = 1.129 − 2.0758 × x + 0.8032 × x2 TSF 0.56 0.0000 *** −0.7423 *
>151 cm ISD = 5.5782 − 5.3475 × x − 2.0348 × x2 DSF 0.32 0.0008 *** −0.5711 *
>151 cm ISD = 10.0399 − 32.7416 × x + 44.103 × x2 ISF 0.16 0.0267 * −0.4280 *
>151 cm ISD = 5.9229 − 7.584 × x + 1.7462 × x2 TSF 0.32 0.0008 *** −0.5583 *
Note: Bl: branch length (cm); CW1: crown width 1; CW2: crown width 2; RCW: relative crown length; CP: crown
projection (cm2 ); ISD: index of spatial distribution; GFr: gap fraction; Op: canopy openness; DSF: direct site factor;
ISF: indirect site factor; TSF: total site factor; p < 0.05 (*), p < 0.01 (**), p < 0.001 (***); p > 0.05 (NS ).

Table A4. Significantly proved regression models of light intensities and morphological parame-
ters and indexes expressing the crown length of Norway spruce natural regeneration under size-
dependent evaluation.

Height
Regression Model Variable (x) R2 p rs
Class
10–50 cm RCL = 0.5913 − 0.0065 × x + 0.0003 × x2 GFr 0.08 0.0051 ** 0.3511 *
10–50 cm RCL = 0.5899 − 0.0063 × x + 0.0003 × x2 Op 0.08 0.0046 ** 0.3528 *
10–50 cm RCL = 0.5333 + 0.4538 × x − 0.304 × x2 DSF 0.05 0.0122 * 0.2499 *
10–50 cm RCL = 0.5936 − 0.549 × x + 2.2822 × x2 ISF 0.08 0.0051 ** 0.3372 *
10–50 cm RCL = 0.5244 + 0.4743 × x − 0.2612 × x2 TSF 0.06 0.0084 ** 0.2685 *
10–50 cm CWLR = 1.1184 + 0.0573 × x − 0.0019 × x2 GFr 0.06 0.0455 * −0.2854 *
10–50 cm CWLR = 1.1229 + 0.0563 × x − 0.0018 × x2 Op 0.06 0.0436 * −0.2899 *
10–50 cm CWLR = 1.1199 + 4.9663 × x − 14.5085 × x2 ISF 0.07 0.0447 * −0.2868 *
10–50 cm Cl = 12.868 − 0.1274 × x + 0.0082 × x2 GFr 0.04 0.0409 * 0.2121 *
10–50 cm Cl = 12.7175 − 0.1159 × x + 0.008 × x2 Op 0.04 0.0354 * 0.2149 *
10–50 cm Cl = 12.7194 − 11.4916 × x + 68.5436 × x2 ISF 0.05 0.0195 * 0.2264 *
51–100 cm CWLR = 2.6967 − 0.0429 × x − 0.0001 × x2 GFr 0.14 0.0041 ** −0.5547 *
51–100 cm CWLR = 2.8315 − 0.0496 × x − 6.3876 × 10−5 × x2 Op 0.16 0.0018 ** −0.5969 *
51–100 cm CWLR = 1.9634 − 2.0859 × x + 0.6197 × x2 DSF 0.08 0.0341 * −0.3290 *
51–100 cm CWLR = 2.2819 + 0.6117 × x − 11.3368 × x2 ISF 0.26 0.0001 *** −0.6968 *
51–100 cm CWLR = 2.2373 − 3.9869 × x + 3.5742 × x2 TSF 0.10 0.0180 * −0.3747 *
51–100 cm RCL = 0.5197 + 0.9095 × x − 0.897 × x2 DSF 0.08 0.0331 * 0.2753 *
51–100 cm RCL = 0.8403 − 2.2948 × x + 5.8313 × x2 ISF 0.11 0.0181 * 0.3168 *
51–100 cm RCL = 0.4589 + 1.3188 × x − 1.5389 × x2 TSF 0.10 0.0247 * 0.2902 *
51–100 cm Cl = 41.4323 − 29.6034 × x + 180.7043 × x2 DSF 0.14 0.0065 ** 0.2624 *
51–100 cm Cl = 12.3012 + 155.6449 × x − 143.2971 × x2 ISF 0.07 0.0449 * 0.2665 *
51–100 cm Cl = 38.9685 − 14.9096 × x + 162.9465 × x2 TSF 0.14 0.0053 ** 0.2773 *
101–150 cm CWLR = 13.8824 − 0.8997 × x + 0.0157 × x2 GFr 0.58 0.0322 * −0.4182 NS
101–150 cm CWLR = 15.3481 − 1.0121 × x + 0.0178 × x2 Op 0.60 0.0292 * −0.4364 NS
101–150 cm CWLR = 10.3003 − 56.1753 × x + 84.8293 × x2 ISF 0.62 0.0087 ** −0.5546 NS
Forests 2022, 13, 1804 18 of 20

Table A4. Cont.

Height
Regression Model Variable (x) R2 p rs
Class
>151 cm CWLR = 6.399 − 0.397 × x + 0.0072 × x2 GFr 0.34 0.0013 ** −0.6320 *
>151 cm CWLR = 6.1167 − 0.3702 × x + 0.0066 × x2 Op 0.33 0.0014 ** −0.6173 *
>151 cm CWLR = 1.8672 − 5.1136 × x + 5.0869 × x2 DSF 0.62 0.0000 *** −0.7537 *
>151 cm CWLR = 2.3706 − 4.9823 × x + 0.7249 × x2 ISF 0.30 0.0012 ** −0.5762 *
>151 cm CWLR = 2.0488 − 6.1746 × x + 6.5934 × x2 TSF 0.63 0.0000 *** −0.7665 *
>151 cm RCL = 0.2823 + 0.0178 × x + 1.239 × 10−5 × x2 GFr 0.19 0.0119 * 0.4051 *
151 cm + RCL = 0.189 + 0.0264 × x − 0.0002 × x2 Op 0.17 0.0206 * 0.3589 *
>151 cm Cl = −14.4542 + 1462.7692 × x − 2300.0565 × x2 DSF 0.30 0.0022 ** 0.5230 *
>151 cm Cl = −83.1842 + 1988.695 × x − 3314.3855 × x2 TSF 0.30 0.0022 ** 0.5171 *
Note: Cl: crown length (cm); RCL: relative crown length; CWLR: crown width–length ratio; GFr: gap fraction;
Op: canopy openness; DSF: direct site factor; ISF: indirect site factor; TSF: total site factor; p < 0.05 (*), p < 0.01 (**),
p < 0.001 (***); p > 0.05 (NS ).

Table A5. Significantly proved regression models of light intensities and the height–diameter ratio of
Norway spruce natural regeneration under size-dependent evaluation.

Height
Regression Model Variable (x) R2 p rs
Class
51–100 cm HDR = 4.7679 + 0.0172 × x + 0.0018 × x2 GFr 0.13 0.0054 ** 0.4334 *
51–100 cm HDR = 3.9528 + 0.0839 × x + 0.0004 × x2 Op 0.14 0.0045 ** 0.4442 *
51–100 cm HDR = 3.6928 + 9.0674 × x + 1.4289 × x2 ISF 0.17 0.0015 ** 0.4481 *
101–150 cm HDR = −31.7919 + 253.2491 × x − 414.6227 × x2 ISF 0.65 0.0400 * 0.3091 NS
>151 cm HDR = −2.9996 + 0.6657 × x − 0.0117 × x2 GFr 0.20 0.0151 * 0.4995 *
>151 cm HDR = −1.6448 + 0.5431 × x − 0.009 × x2 Op 0.20 0.0121 * 0.4875 *
>151 cm HDR = 5.4822 + 1.3029 × x + 7.8746 × x2 DSF 0.23 0.0065 ** 0.4172 *
>151 cm HDR = 7.439 − 18.9085 × x + 50.8065 × x2 ISF 0.16 0.0254 * 0.3861 *
>151 cm HDR = 5.5556 − 0.2287 × x + 12.4244 × x2 TSF 0.23 0.0057 ** 0.4322 *
Note: HDR: height–diameter ratio; GFr: gap fraction; Op: canopy openness; DSF: direct site factor; ISF: indirect
site factor; TSF: total site factor; p < 0.05 (*), p < 0.01 (**); p > 0.05 (NS ).

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