3.1. Sapwood Metrics
- (a)
Description of the metrics
Table 2 shows the descriptive statistics of the sapwood metrics and reveals distinct differences between species. Pine exhibited the highest mean sapwood width at 64 ± 24 mm, followed by silver fir at 58 ± 20 mm, and spruce, which had the narrowest sapwood at 41 ± 13 mm. This trend was reflected in the proportion of sapwood area, where pine also had the highest mean value (71 ± 15%), compared to silver fir (68 ± 15%) and spruce (57 ± 12%). Notably, pine also displayed the greatest variability in absolute sapwood width (SD = 24 mm), while spruce was the least variable in both its width (SD = 13 mm) and proportion (SD = 12%). These overall statistics confirm clear, species-specific patterns in sapwood characteristics.
- (b)
Sapwood proportion
For both spruce and silver fir, a clear decreasing trend in sapwood proportion was observed with increasing sawlog diameter (
Figure 6). At any given diameter class, spruce consistently exhibited a lower sapwood proportion than fir. For example, in the 150 mm diameter class, the sapwood proportion was approximately 65% for spruce and 75% for fir. This decreased to around 52% and 60%, respectively, in the 500 mm diameter class. This trend suggests that in these two species, the relative allocation of radial growth shifts with age. In the initial growth phase of trees, the cross-section of the trunks consists mainly of sapwood. As the trees age, heartwood gradually forms, particularly to provide mechanical support. The distinct difference between spruce and fir may reflect species-specific physiological strategies related to growth and water conduction.
In contrast, the sapwood proportion for pine remained relatively high and more stable, starting around 80% and decreasing only slightly to approximately 70% in the 400 mm diameter class before appearing to increase again. This high sapwood proportion may reflect an adaptive strategy of this pioneer species, centered on efficient water conduction and delayed duraminization, likely enhancing its tolerance to various soil types. Pine maintains a high proportion of conductive xylem area, which can be accurately measured using techniques based on wood moisture content rather than staining methods [
39]. This has direct implications for processing: large-diameter pine sawlogs will contain a proportionally larger volume of sapwood, which is less durable and non-colored, and has a higher moisture content. This directly affects the yield of colored heartwood and dictates different requirements for the drying process.
The knowledge of sapwood properties allows for more advanced sorting; larger sawlogs can be preferentially allocated to products where heartwood properties are desired by the customers. The authors of [
7,
40] also documented sapwood thickness variations in relation to tree characteristics, confirming that these patterns are inherent and could be predictable. The ability to model these species-specific trends provides a powerful tool for optimizing raw material allocation before the first cut is made.
- (c)
Sapwood width
For all three species, the mean sapwood width showed a clear increasing trend with sawlog diameter. However, the rate of increase differed. Pine consistently presented the greatest sapwood width, starting at approximately 40 mm in the 100 mm diameter class and increasing to more than 140 mm in the 550 mm class. Silver fir displayed an intermediate width, while spruce consistently had the narrowest sapwood, growing from roughly 30 mm to 80 mm across the same diameter range.
It is important to note that for diameter classes exceeding 450 mm, the sample size for all species decreased substantially. Overall, 500 and 550 mm classes had 1164 and 77 samples respectively, across all species. The apparent upward trend in sapwood proportion for pine in the largest diameter classes, for instance, may not be representative of the general population and should be interpreted with caution due to the low number of observations.
- (d)
Statistical analysis
The test for the homogeneity of regression slopes, via the diameter and essence interaction term, was highly statistically significant (F(2, 28,560) = 125.02,
p < 0.001). This finding indicates that the assumption of parallel slopes was violated. Therefore, the relationship between sawlog diameter and sapwood proportion was fundamentally different among the species. Because the interaction was significant, we analyzed the slopes from the interaction model as shown in
Appendix A,
Figure A1. The estimated slopes (the rate of change in sapwood proportion per mm of diameter) were pine: −0.038; silver fir: −0.073; spruce: −0.073.
A post hoc pairwise comparison of these slopes (
Table A1) revealed that the slope for pine was significantly different from that of both silver fir (
p < 0.001) and spruce (
p < 0.001). However, the difference between slopes in silver fir and spruce was not statistically significant (
p = 0.9909). This indicates that the sapwood proportion of pine decreased significantly more slowly with increasing sawlog diameter, whereas silver fir and spruce exhibited a statistically similar and steeper decline. This finding is critical for accurately modeling wood properties, as it shows that pine follows a distinct developmental pattern regarding sapwood proportion compared to spruce and fir.
3.2. Knots Metrics
- (a)
Description of the metrics
As detailed in
Table 3, knot characteristics varied by species and type. Particularly, sound knots were larger in diameter than black knots for all three species, a trend that was most pronounced in pine. Pine sawlogs consistently presented the largest and most frequent sound knots. In contrast, spruce was characterized by having the smallest knots of both types and the lowest frequency of black knots. However, the knot diameters measured in our study are comparable in magnitude to those modeled for Scots pine and Norway spruce in other previous European studies [
41,
42].
- (b)
Dimensions of sound knots and variability
Figure 7 illustrates the variations in the average diameter of sound knots and their standard deviation as a function of sawlog diameter classes. For all three species, the average diameter of sound knots increased with sawlog diameter. Comparatively, as mentioned previously, pine tended to have the largest sound knots, followed by silver fir, with spruce having the smallest.
This marked increase directly reflects branch growth dynamics; branches that persist longer on larger trees, which are often older or have benefited from more growing space, reach larger dimensions. These findings are consistent with models for other conifer species that link larger branch and knot diameters to increasing tree size and stem diameter.
The standard deviation of sound knot diameter increased with the sawlog diameter class, indicating a greater dispersion of knot sizes in larger sawlogs. Pine was distinguished by higher variability compared to spruce and fir, which both exhibited similar levels of variation on this criterion.
This suggests greater heterogeneity in the development of living branches in pine. Such variability may stem from a broader range of branch vigor within the crown (e.g., a mix of dominant, vigorous branches and smaller, suppressed ones), a more plastic response to environmental conditions, or different genetic controls on growth. Indeed, several studies have highlighted significant genetic influences on growth traits in pine, which may contribute to this variability. This heterogeneity complicates the prediction of sawn timber quality from pine sawlogs and underscores the importance of detailed, individual sawlog assessment. It should be noted that the lower number of pine samples in this study could amplify the apparent heterogeneity.
- (c)
Dimensions of black knots and variability
Figure 8 shows the relationship between the mean diameter of black knots and the sawlog diameter class for the three species. A positive correlation was observed for all species, where the average diameter of black knots increased with sawlog diameter. This trend may reflect, in part, biological processes such as crown dynamics, where dominant trees develop larger branches that persist longer before dying due to crown recession and natural pruning. It may also result from the fact that large-diameter sawlogs are often taken from the lower part of the stem (i.e., butt logs), where older and larger branches had formed early in the tree’s development, before being naturally pruned. The resulting black knot diameter therefore corresponds to the maximum size the branch achieved while it was alive. In inter-species comparison, pine consistently presented the largest black knots, followed by silver fir, with spruce having the smallest.
The variability in black knot diameter, indicated by the standard deviation, also increased with sawlog diameter class. This suggests a greater range of dead branch sizes being occluded within larger sawlogs. Pine was distinguished by higher variability compared to silver fir and spruce, which showed more constrained and similar levels of variation. This pronounced heterogeneity in pine likely reflects a greater plasticity in its response to stand density and competition for this pioneer species, leading to more diverse branch growth and mortality scenarios. This high variability in a key wood defect complicates quality assessment based on external sawlog features alone and stresses the challenge in predicting the final grade of sawn timber from pine.
- (d)
Number of sound and black knots per linear meter
Figure 9 shows that the number of sound knots per linear meter decreased as sawlog diameter increased for all three species. The reason is that large-diameter sawlogs in the wood supply are typically butt sawlogs, from which lower branches have naturally shed over time, resulting in fewer sound knots from the original living crown. In contrast, small-diameter sawlogs are often top sawlogs that originate entirely within the live crown, where nearly all knots are sound. This distinction between the living crown and the dead branches is critical for accurately modeling knots characteristics. Among the species, silver fir exhibited the highest density of sound knots, particularly in the intermediate diameter classes.
- (e)
Statistical analysis
The ANCOVA test was found significant for the linear models of both black and sound knots, showing that there was no homogeneity of the regression slopes. Therefore, there was an interaction of the species on knot variability.
The analysis investigated how knot diameter variability is influenced by tree species, sawlog diameter, and mean knot size. All four linear models were highly significant (
p < 0.001), explaining 66%–79% of the variance (adjusted R
2) in knot standard deviation (
Figure 10). The summary of the models is available in
Appendix B,
Table A2. The key findings relate to the moderating effect of species on these relationships.
A significant interaction was found between sawlog diameter and species for both black and sound knot variability, indicating that the relationship between sawlog size and knot variability differs among species. For black knots, the variability in spruce increased significantly less with sawlog diameter compared to pine (p = 0.005). The trend for silver fir was not significantly different from pine. For sound knots, the increase in variability with sawlog diameter was significantly steeper for pine than for either silver fir (p < 0.001) or spruce (p = 0.004). The slopes for silver fir and spruce did not differ significantly from each other.
The relationship between the mean diameter of knots and their standard deviation (a measure of heteroscedasticity) was also significantly moderated by tree species. The variability of black knots in spruce showed a significantly different relationship with mean knot size compared to that of pine (p = 0.003). Specifically, the slope of this relationship was less pronounced for spruce. A similar interaction was observed for sound knots, where the relationship between mean size and variability for spruce was significantly different from that of pine (p = 0.010). In this case, the variability in sound knots increased more sharply with their mean size in spruce than in pine.
Pine consistently showed the strongest positive relationship between sawlog diameter and sound knot variability. Furthermore, the relationship between mean knot size and its own variability was distinct for spruce compared to pine and silver fir, particularly for sound knots.
3.3. Sapwood and Heartwood Density and Moisture Content
The seasonal data, presented in
Figure 11, reveals distinct and physiologically trends in wood moisture content (MC) and density for all three species, particularly within the sapwood.
In
Table 4, the analysis shows a pronounced difference in moisture content (MC) between the two wood types across all three species. Sapwood consistently exhibited substantially higher MC, with mean values ranging from 93% for pine to 105% for silver fir. In contrast, heartwood was drier, with mean MC values between 23% (spruce) and 43% (silver fir). This clear distinction aligns with the primary physiological roles of sapwood in water conduction and heartwood in structural support.
The species ranking in our data (silver fir > spruce > pine) are similar in the XyloDensMap reference data [
37]. Silver fir is the species with the highest moisture content in both datasets. Furthermore, the maximum MC values recorded in our study (136%–172%) show correspondence with the maximums reported in XyloDensMap (150%–180%), suggesting a similar upper limit for moisture saturation across the species.
- (a)
Seasonal dynamics of moisture content in sapwood
The most noticeable trend observed across spruce, silver fir, and pine was a consistent and marked decrease in sapwood moisture content from early spring (March) to late summer (August). For all species, sapwood MC began at its peak, typically between 110%–120% and progressively declined to 70%–85% by August. This pattern directly corresponds to the seasonal cycle of tree physiology. The high initial MC in spring reflects the mobilization of water for budbreak and the onset of transpiration. As the summer progresses, higher temperatures and increased evapotranspiration rates lead to a gradual desiccation of the sawlogs post-harvest, a well-documented phenomenon as initial moisture content is typically high, especially in sapwood, and decreases over time due to drying [
43,
44]. This seasonal variation in wood MC has practical implications for the timber industry, as it affects both the sawlog weight during transport and the amount of water evaporated during the drying of sawn products. Managing a log yard effectively requires adapting to these seasonal supply variations [
45].
- (b)
Contrasting seasonal behavior of sapwood and heartwood MC dynamics
In contrast to the dynamic nature of sapwood, the heartwood in all three species demonstrated remarkable stability in both moisture content and density throughout the observation period. Heartwood MC remained low and relatively constant, typically ranging between 25% and 45%, with minimal variation. This stability is expected, as heartwood is composed of non-living cells and does not participate in active water transport [
46]. Studies on various softwood species show that sapwood typically has higher initial moisture content and drying rates than heartwood [
47]. This fundamental physiological difference highlights the contrast between the water-conducting sapwood and the inert, structurally focused heartwood. It is worth noting that the low MC observed in the heartwood, sometimes falling below the fiber saturation point (FSP), may partially reflect drying that occurred between tree felling and the CT scanning.
- (c)
Inter-species variations
While the overall trends are similar, subtle differences between species are apparent.
Silver fir and spruce showed very similar patterns: a steep, consistent decline in sapwood MC and highly stable heartwood properties. This suggests a comparable physiological response to seasonal changes and post-harvest drying.
Pine displayed a slightly more different pattern, particularly in its heartwood. A minor but noticeable increase in heartwood MC was observed from April to July. While the variability was large, this could be an artifact of the smaller sample size for this species in certain months as indicated by the sawlog count bars. Furthermore, the initial sapwood MC in pine appeared slightly lower, likely due to its reduced porosity compared to the other two species.
3.4. Current Limitations
While this study is based on an exceptionally large dataset, it has certain limitations. The sample sizes were imbalanced across the three species. These factors may affect the robustness of the observed trends at the upper end of the sawlog diameter range. Future work should aim to increase the number of sawlogs for the higher sawlog diameter classes. Moreover, further work ongoing will focus, from a statistical point of view, on the variations of quality metrics (means and standard deviations) in relation to the number of measured sawlogs.
It is also important to acknowledge the absence of information on the health status of the study sample as a potential limitation of this method. In fact, confounding effect of fungal decay, such as butt rot caused by
Heterobasidion annosum [
48], on density readings, and consequently on MC estimation, warrants further investigation. Similarly, colonization by bark beetles like
Ips typographus also warrants consideration as the fungi they introduce are known to alter wood’s physical properties [
49]. Over time, these fungal infestations can cause weight loss and increase wood permeability, which would also introduce variability into density-based moisture content estimations.
Additionally, another limitation of this study is the method used to estimate MC. The calculation relied on a single, species-specific average basic density (ID) sourced from the XyloDensMap database [
37]. This approach, while based on a large and reliable dataset, does not account for the natural, inter-tree variability in basic density that exists within any given species. This simplification may contribute to the notably low mean MC values observed, particularly in the heartwood of spruce, which were lower than typically expected. Due to the industrial and retrospective nature of the data collection, direct verification of these values on the study sample was not possible.
Nevertheless, the methodology is grounded in established physical principles and standard formulas [
35]. While the absolute MC values should be interpreted with caution, the primary value of these results lies in their relative comparisons and observed trends. The contrast between sapwood and heartwood dynamics, the clear seasonal decline in sapwood MC, and the comparative differences between the three species are robust and physiologically consistent findings. Therefore, the results serve as a valid, large-scale indicator of seasonal and structural moisture dynamics, even if the absolute precision for any single log is limited. Future work will aim to incorporate more direct MC measurement techniques to refine and validate these estimations.