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Article

Substrate and Fertilization Used in the Nursery Influence Biomass and Nutrient Allocation in Fagus sylvatica and Quercus robur Seedlings After the First Year of Growth in a Newly Established Forest

by
Odunayo James Rotowa
1,2,*,
Stanisław Małek
1,
Michał Jasik
1 and
Karolina Staszel-Szlachta
1
1
Department of Ecology and Silviculture, Faculty of Forestry, University of Agriculture in Kraków, Al. 29 Listopada 46, 31-425 Kraków, Poland
2
Department of Forestry and Wildlife Management, Faculty of Agriculture, Nasarawa State University, Keffi, Nigeria
*
Author to whom correspondence should be addressed.
Forests 2025, 16(3), 511; https://doi.org/10.3390/f16030511
Submission received: 7 February 2025 / Revised: 9 March 2025 / Accepted: 12 March 2025 / Published: 14 March 2025
(This article belongs to the Section Forest Ecophysiology and Biology)
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Versions Notes

Abstract

:
This study evaluates the efficacy of innovative peat-free organic substrates and liquid fertilizers as alternatives to traditional peat substrates in the cultivation of Fagus sylvatica L. and Quercus robur L. seedlings in a newly established forest in Southern Poland. The experiment was conducted in a 2 × 2 × 4 experimental layout using a randomized complete block design, comprising eight treatments that combined four substrate types (three novel organic substrates and one peat-based control) with two types of fertilizers (solid and liquid). After one year of growth, biomass and nutrient allocation in the roots, shoots, and leaves of the seedlings were analyzed. The results showed that while solid fertilization enhances biomass accumulation, liquid fertilization supports more uniform growth across different substrates, particularly in oak seedlings. Also, peat substrates recorded the highest nutrient allocation. However, one novel substrate (R22) performed comparably, indicating its potential as a viable peat alternative. Significant interspecies differences were observed, with beech seedlings allocating more biomass to aboveground organs, while oak seedlings favored belowground nutrient allocation. These findings suggest that while peat substrates and solid fertilizers currently provide better outcomes, the innovative R22 substrate shows promise for sustainable forestry practices. Further refinement of the liquid fertilizer was recommended to enhance effectiveness.

1. Introduction

Environmental factors exert a profound influence on physiological traits, thereby demonstrating key aspects of developmental plasticity in plants [1,2,3]. The success of forest plantations is contingent upon the traits of their seedlings. Biomass from forests has the potential to provide a continuous, largely carbon-neutral supply of materials for the forestry industry [4]. The cultivation practices employed in nurseries can have a significant impact on the functional characteristics and field performance of seedlings. These practices can influence post-transplant rooting and early growth, as evidenced by the studies conducted by Villar-Salvador et al. [5] and Grossnickle and MacDonald [6]. Conversely, some research indicates that seedlings with limited nutrient availability may exhibit greater resilience to transplant shock and summer drought [7,8]. Fertilization can enhance plant survival through a variety of mechanisms. For example, root growth potential and hydraulic conductance are enhanced by phosphorus (P) and nitrogen (N) availability [9,10] which increases the capacity of fertilized seedlings to absorb soil water [11]. N- and P-deficient plants frequently exhibit alterations in their biomass accumulation and allocation patterns [2,12].
In Poland, the extensive restructuring of coniferous monocultures has been necessitated by the declining health and quality of the trees. European beech (Fagus sylvatica L.), a temperate species that is widely distributed across central and Western Europe [13], along with oaks (Quercus spp., including Quercus robur L.), are the dominant species in Polish forests and constitute a significant portion of European temperate vegetation. Beech and oak forests are of great importance to the biosphere, contributing to biomass production, oxygen generation, atmospheric regulation, and more [14]. Due to their superior wood quality, beech and oak have become more commercially desirable than several conifer species and are favored in climate change adaptation strategies for both ecological and economic reasons in Europe [15]. Global changes have demonstrated that shifts in tree species and their symbiotic associations have an impact on biogeochemical processes [16,17,18]. Consequently, it is imperative to intensify efforts to enhance the health and sustainability of forest stands dominated by these highly valued species, particularly as Polish forests transition from pine to species such as beech and oak.
Peat is widely recognized as a foundational component in nursery substrates due to its exceptional physical, chemical, and biological properties. Its remarkable water-retention capabilities and consistent quality make it a preferred medium for the nurturing of plants [19,20]. Nevertheless, the release of carbon (C) from peat soils over time has the potential to give rise to environmental concerns. In contrast to forests, which act as C sinks, peatlands have the potential to release stored C into the atmosphere, which can have a significant impact on climate change. Given that peatlands store an estimated one-third of the world’s soil C, which exceeds the combined capacity of global forests, the rapid conversion of peat to CO2 in plantations has the potential to elevate greenhouse gas levels, thereby threatening our treasured ecological systems [21]. Gruda [22] notes that Europe excavates approximately 20,000 cubic meters of peat annually, which has the effect of exacerbating environmental degradation [23]. In light of the urgent global environmental challenges associated with peat use in nursery substrates, it is of the utmost importance to prioritize the preservation of peatlands over their destruction. These, therefore, underscore the urgent need for sustainable alternatives.
Over the years, there has been an increased adoption of compost utilization in place of or as a mixture with peat [24,25]. The process of production is time-consuming and labor-intensive. If not carried out correctly, compost can harbor pathogens, weed seeds, and plant diseases [26]. As a result of the contemporary emphasis on sustainability and environmental awareness, there is a pressing need to develop a novel peat-free organic substrate using sustainable, cost-effective, and eco-friendly materials as viable alternatives to peat. The novel substrate used in this study was designed to overcome these challenges, with properties, including water capacity, bulk density, and solid density, meeting those of standard peat substrates [27].
In previous studies, the novel substrates and liquid fertilizer used in this research demonstrated promising physicochemical properties, such as water retention capacity, bulk density, and nutrient content, comparable to those of traditional peat substrates [27]. Specifically, the R22 substrate exhibited enhanced structural stability and nutrient availability after the nursery production cycle [28]. Building on these findings, this study extends the evaluation to assess the performance of seedlings after one year of growth in a forest environment. This approach allows us to determine whether the promising outcomes observed in the nursery translate into sustainable growth and nutrient allocation under field conditions as recommended by Rotowa et al. [28]. Integrating these advancements, our study aims to explore the practical potential of these innovations for sustainable nursery and forestry practices.
The objective of this study is to evaluate the impact of novel peat-free organic substrate and liquid fertilizer, developed by the University of Agriculture in Kraków, Poland, on the nutrient allocation and biomass production in Fagus sylvatica and Quercus robur seedlings after one year of growth in a forest environment. This study hypothesizes that novel peat-free organic substrates and liquid fertilizers will again result in comparable or superior seedling growth, biomass allocation, and nutrient distribution compared to traditional peat substrates and solid fertilizers. This hypothesis is grounded in the demonstrated physicochemical benefits of the innovative substrates and fertilizers after the nursery production cycle. By evaluating these plant materials under both nursery and field conditions, the study seeks to bridge the gap between experimental advancements and practical forestry applications.

2. Materials and Methods

2.1. Study Site

The study site was situated in Barbarka, within the Miechów Forest District (Figure 1). The research area is situated at an altitude of approximately 370 m above sea level in the Olkuska Upland, southern Poland (50°15′54.2″ N 19°53′36.5″ E). It is located within a forest complex managed by the National Forest Holding. The area in question encompasses several gaps created by the clear-cutting of a Populus spp. plantation. The Miechów Forest District is distinguished by its diverse upland landscape. The Olkuska Upland is a compact karst plateau composed of limestone and marl. The climate is continental, exhibiting a notable temperature range of 21 °C and a considerable amount of precipitation during the growing season. The mean annual air temperature in the Forest District is 8.2 °C, with July being the warmest month (19.6 °C) and January the coldest (–3.0 °C).

2.2. Substrate Composition and Seedling Production

At the nursery stage of the experiment, the control variant (C) peat substrate was produced at the Nursery Farm in Nędza (50.167964 N, 18.3138334 E). The substrate was composed of 93% peat and 7% perlite, with the addition of dolomite (3 kg per m3 of substrate) to achieve a pH of 5.5. The novel peat-free substrates (R20, R21, and R22) were manufactured from diverse components from coniferous woody materials (mainly Pinus sylvestris L.), with specific proportions designed to optimize water retention and nutrient availability (Table 1). Physicochemical properties, such as water capacity, bulk density, and porosity, were measured following standardized protocols to ensure comparability (Table 2). The peat-free substrates and liquid fertilizer used in this study were developed under the project POIR.04.01.04-00-0016/20, which was funded by the National Centre for Research and Development (NCBiR) from national resources and the European Regional Development Fund. This project was spearheaded by the Department of Ecology and Silviculture, University of Agricultural in Kraków. The comprehensive procedure for preparing the novel substrate and fertilizer was previously described by Rotowa et al. [27].
In each experimental variant, seedlings of both species were cultivated in 75 Marbet V300 polystyrene containers, each containing 53 cells with a volume of 275 cm3. Subsequently, the various substrates were filled into containers, and beech and oak seeds were manually sown at the Suków-Papierna Nursery Farm (Daleszyce Forest District) on 19–20 April 2022. To enhance germination, seeds of both species were scarified before sowing. After sowing, the containers were transferred into a vegetation hall for four weeks, after which they were relocated to an external production field. Manual weeding was conducted during the seedling growth phase. The seedlings were cultivated for five months following the procedures employed in the container nursery, as outlined by Szabla and Pabian [29].
Osmocote fertilizer was incorporated into the substrate during its preparation prior to sowing, with a total application rate of 3 kg m3 for each medium. The mixture comprised Osmocote 3–4 M (2 kg) and Osmocote 5–6 M (1 kg). The Osmocote 3–4 M formulation contained 16% N (7.1% N-NO3− and 8.9% N-NH4+), 9% P2O5, 12% K2O, 2.0% MgO, and microelements (B, Fe, Cu, Mn, Zn, and Mo). The Osmocote 5–6 M formulation included 15% N (6.6% N-NO3− and 8.4% N-NH4+), 9.0% P2O5, 12% K2O, 2.0% MgO, and the same microelements. A novel liquid fertilizer regime was introduced, utilizing two distinct compositions. These application rates were determined based on the nutrient demand of the seedlings at different growth stages and the need to ensure adequate nutrient availability without causing leaching losses. The first liquid fertilizer contained 4.78% N, 1% P2O5, 2.64% K2O, 2.65% CaO, 1.4% MgO, 0.71% SO3, and 0.14% Na2O. It was initially applied at a total volume of 3.14 dm3 (0.048 dm3 m−2) to provide an immediate supply of essential macronutrients required for early root and shoot development. The second liquid fertilizer contained 0.798% N, 0.166% P2O5, 0.440% K2O, 0.441% CaO, 0.234% MgO, 0.118% SO3, and 0.023% Na2O, applied at a total volume of 15.09 dm3 (0.229 dm3 m−2). This staggered approach was designed to sustain nutrient availability throughout the critical growth phases, promoting steady biomass accumulation and nutrient allocation. During seedling production, the first liquid fertilizer was applied eight times at 10-day intervals, while the second was applied 15 times at 5-day intervals. These fertilization schedules were consistently maintained for both European beech and pedunculate oak seedlings to optimize nutrient use efficiency while minimizing environmental impact. This approach aligns with previous research findings on staged nutrient applications to enhance seedling performance in forest nurseries [27,29].

2.3. Experimental Layout

After the nursery production cycle, the seedlings were transported and planted into the forest on 5 September 2022. The experimental design employed a 2 × 2 × 4 factorial layout within a randomized complete block design (RCBD), comprising four substrate types (three novel and one peat-based control) and two fertilizer types (solid and liquid), resulting in eight treatment combinations. Each treatment was replicated three times within blocks to minimize spatial variability. The blocks were stratified based on topographic and soil characteristics to ensure uniform environmental conditions. Subplots, each containing 49 seedlings were established with consistent spacing (1 × 1.7 m) to standardize growing conditions across treatments. A total of 24 subplots were established for each species. Four substrates (R20, R21, R22, and peat) and two fertilizer treatments were employed to cultivate the seedlings. The first fertilizer treatment was a solid fertilizer utilized in the Suków container nursery (SR20, SR21, and SR22 variants), while the second was a novel liquid fertilizer developed by the University of Agriculture in Kraków, Poland (UR20, UR21, and UR22 variants). In both fertilization cases, the peat substrate served as the control variant (SC and UC).

2.4. Soil Sample Collection and Plantation Establishment

The research plot, comprising 0.7 ha, was established on a Populus spp. harvest site with similar parent material and soil type to that of an old-growth forest. Based on the primary active root zones of young seedlings; soil samples were collected from five different locations within each subplot at two layers. The top mineral layer at depths of 0–10 cm and the second layer at 10–20 cm were collected using polyvinyl chloride (PVC) tubes. The layer of the top 0–10 cm is not only conducive to initial root growth and early seedling establishment, but rich organic matter nutrients also make it essential. This layer is also the primary domain of microbial activity and nutrient cycles, and the 10–20 cm layer was chosen to test beyond the immediate root zone the availability of nutrients, as well as the potential effects of leaching from fertilization. The inclusion of these two depths provides a comprehensive analysis of nutrient dynamics within the soil, ensuring that the study captures both immediate and longer-term soil fertility impacts on seedling growth [30]. A total of 480 soil samples were collected. Each sample was then air-dried, sieved through a 2 mm mesh, ground, and prepared for the analysis of its soil properties following standard soil preparation protocols [31]. To ensure consistency in analysis, the soil samples were air-dried, sieved, and ground. Air-drying stabilized the samples and prevented microbial activity that could alter nutrient content. Sieving through a 2 mm mesh removed debris and homogenized the samples, ensuring uniformity across all the analyses. Grinding the samples into a fine powder increased surface area, enhancing the accuracy of nutrient extractions and spectrometric measurements. This process maintained comparability between soil samples, ensuring that the results accurately reflected the nutrient composition of the study site.
The physicochemical properties of the samples were determined following established methods described by Ostrowska et al. [31] and Staszel et al. [30]. The pH values of the soil were measured potentiometrically in both water and 1 M KCl. Hydrolytic acidity was evaluated using the Kappen method, while exchangeable acidity and content were estimated using the Sokołow method. The total N and C contents were determined using a LECO CNS True Mac Analyzer (Leco, St. Joseph, MI, USA). To quantify alkaline cations (Ca2+, Mg2+, K+, and Na+), 1 M ammonium acetate was employed (iCAP 6500 DUO, Thermo Fisher Scientific, Cambridge, UK), utilizing inductively coupled plasma optical emission spectrometry (ICP-OES). The analyses were conducted at the Laboratory of Forest Environment Geochemistry and Land Intended for Reclamation, Department of Ecology and Silviculture, and the Faculty of Forestry at the University of Agriculture in Krakow, Poland.
The field experiment was conducted using a randomized complete block design. In each subplot, 49 seedlings were planted following a standardized spacing of 1 × 1.7 m both between and within rows, ensuring consistent plant density across all the treatments. Before transplantation, the seedlings were selected based on specific morphological criteria to ensure uniformity and quality. The selection criteria included a minimum height of 25 cm, a root system length of at least 15 cm, a well-developed collar diameter, and healthy stem and leaf formation. These parameters were used to enhance transplant success and minimize variability in initial seedling vigor. The planting depth was standardized to ensure that the root collar was level with the soil surface, a technique aimed at optimizing root establishment and reducing transplant shock. For both species combined, a total of 2352 seedlings were established (147 seedlings per treatment). At the end of the 2023 growing season, three seedlings were selected from each of the 49 seedlings in every subplot (nine seedlings per treatment). The seedlings were assessed for root, shoot, and leaf biomass and nutrient allocation. These seedlings were carefully uprooted to preserve an intact root segment. The mean height of the seedlings in each subplot was recorded and this mean value formed the choice of the selected seedling. The seedlings were selected from each of the eight treatment groups for root, shoot, and leaf biomass and the subsequent laboratory analysis for nutrient allocation, leading to the assessment of 144 seedlings for both species in the laboratory experiment. To protect the new forest from animal interference, the area was fenced after the plantation was established.

2.5. Parameter Assessment and Nutrient Analysis

Following the conclusion of the 2023 growing season, the biomass of different plant organs, including leaves, shoots, and roots, was quantified. To validate the hypothesis, biomass allocation (roots, shoots, and leaves) and nutrient concentrations (C, N, P, K, S, Ca, and Mg) were measured after one year of growth in a newly established forest. These parameters were chosen based on their relevance to seedling survival and productivity under field conditions. The samples were dried at 65 °C for 48 h to achieve a constant weight, then ground and homogenized. For each organ, 0.5 g of the ground sample was placed into a flask and subjected to acid digestion using a 3:1 mixture of nitric and hydrochloric acids. After digestion, the samples were filtered into a 50 mL flask, and the concentration of elements was determined using inductively coupled plasma optical emission spectrometry (ICP-OES). Quality control measures were implemented throughout the nutrient analysis process to ensure accuracy and reliability. Calibration standards were prepared and run alongside the samples to verify the accuracy of the ICP-OES readings. Additionally, blank samples and duplicates were included in the analysis to monitor and correct for any potential contamination or analytical drift. The samples were analyzed for N, sulfur, and C contents using a LECO CNS TruMac analyzer (Leco, St. Joseph, MI, USA). The phosphorus, potassium, calcium, and magnesium (P, K, Ca, and Mg) contents were analyzed using a Thermo iCAP 6500 DUO ICP-OES spectrometer (Thermo Fisher Scientific, Waltham, MA, USA) following mineralization in a 3:1 mixture of nitric and hydrochloric acids. These analyses were conducted at the Laboratory of Forest Environment, Geochemistry, and Land Intended for Reclamation in the Department of Ecology and Silviculture, Faculty of Forestry, University of Agriculture in Kraków, Poland.

2.6. Statistical Analysis

Data analysis was conducted using a multifaceted approach. A descriptive analysis was conducted to compare the performance across treatments. The Shapiro–Wilk [32] and Levene [33] tests were employed to verify compliance with normality, homogeneity of variance, and linearity assumptions. To reveal significant differences between treatments for each measurement, post hoc analyses were performed with DMRT. The significance of these relationships and the differences between species combinations were revealed using ANOVA. Before performing each model, the homogeneity of variances and the normality of residuals were assessed, and data were log-transformed where necessary. A clustered heatmap was used to identify the hierarchy clustering using colors to represent values. The row (nutrient) and the column (substrate and fertilizer treatment) of the data matrix were ordered according to the output of clustering. The values were standardized based on mean and standard deviation using color to show the relative expression of individual nutrient allocation between individual treatments. A dendrogram was used to separate the homogeneous relation within the row and column. This dendrogram represents the hierarchical clustering of treatments based on nutrient allocation patterns in the studied species. Clusters were generated using Ward’s method, with Euclidean distance as the similarity metric. Branches indicate treatments with similar nutrient allocation profiles, and the length of the branches reflects the degree of similarity. Heapmap Pearson correlation coefficients (r) were further calculated to examine linear relationships between biomass and nutrient variables. The relationships were deemed significant at the 0.05 level [34]. Correlation coefficients were interpreted as follows: r ≤ 0.35 indicated low or weak correlations (deep blue color) r from 0.36 to 0.67 indicated modest or moderate correlations (light blue to light red color) r from 0.68 to 1.0 indicated strong or high correlations (red color) Regression analysis focused on nitrogen (N) and phosphorus (P) due to their primary role in plant growth, nutrient allocation, and physiological processes. Data visualizations were performed using Python (version 3.10, Python Software Foundation, Wilmington, DE, USA).

3. Results

3.1. Soil Properties

Both the beech and oak sites exhibit slightly acidic soils, which are characteristic of forest environments. The pH values are marginally lower at a depth of 20 cm compared to 10 cm. The oak site demonstrates a consistent N level across both depths. The C content was observed to be higher at the 10 cm depth for both sites, with a subsequent decrease at 20 cm. Furthermore, P levels also decrease with depth at both sites. The carbon/nitrogen (C/N) ratio remains stable across depths, indicating balanced C and N cycling within the soil. Exchangeable cations, including Ca, K, Mg, and Na, were more concentrated at the 10 cm soil depth than at 20 cm across both sites. This suggests that nutrient availability in the upper soil layer may play a crucial role in seedling nutrient uptake, particularly during the early stages of growth. A statistical analysis of soil properties revealed no significant differences between the two sites despite minimal numerical variations across the soil depths (Table 3). This suggests that soil composition was relatively uniform across the study area, allowing for a controlled comparison of the different treatments’ effects on seedling growth.

3.2. Biomass Allocation in Studied Organs of F. sylvatica and Q. robur Seedlings

A biomass allocation analysis revealed significant interactions between substrate type and fertilization method. F. sylvatica demonstrated a stronger dependence on the peat-based control substrate, whereas Q. robur exhibited greater adaptability to novel substrates. Solid fertilization was generally more effective in enhancing both species’ shoot and root biomass accumulation. In contrast, the seedlings treated with liquid fertilization showed greater adaptability to alternative growing media after one year of growth in a new forest (Figure 2 and Figure 3). For the beech seedlings, those grown in the peat-based control (SC) exhibited superior shoot and root biomass accumulation compared to those grown in novel substrates under solid fertilization. However, biomass allocation in the seedlings grown in novel substrates remained consistent across all the organs under liquid fertilization (Figure 2). The response of oak seedlings was somewhat different. While the peat-based control still supported better overall growth and showed variation in root, shoot, and leaf allocation under solid fertilization. Biomass allocation remained relatively stable in more stable organs (root and shoot), with no significant differences among the seedlings raised with liquid fertilization (Figure 3). In both species, a positive correlation was observed between root biomass and both shoot and leaf biomass, suggesting that root development is closely linked to aboveground growth. Similarly, shoot and leaf biomasses were positively correlated (Figure 4a–c). Notably, the regression lines for beech in root–shoot and shoot–leaf relationships were steeper than those for oak, indicating a stronger dependence of shoot and leaf biomass on root biomass in beech compared to oak.

3.3. Allocation of Nutrients in F. sylvatica and Q. robur Seedlings One Year After Planting in the Forest

The heatmap analysis (Figure 5 and Figure 6) illustrates the impact of different treatments on nutrient allocation in the roots, shoots, and leaves of both F. sylvatica and Q. robur after one year of growth in a new forest. A consistent trend emerged across all the treatments, with roots generally exhibiting the highest nutrient allocation compared to other organs. Notably, the seedlings treated with traditional solid fertilizers demonstrated significantly higher nutrient concentrations across all organs, whereas those treated with novel liquid fertilizers had consistently lower nutrient levels. These findings emphasize the superior efficacy of solid fertilizers in nutrient uptake.
Beech seedlings treated with solid fertilizers, particularly in the SC treatment, exhibited the highest concentrations of all the elements. Interestingly, the novel substrate SR22 showed nutrient levels comparable to those of the peat control (SC), indicating its potential as an effective alternative (Figure 5a). In contrast, the leaves treated with liquid fertilizers showed an opposite trend. It is noteworthy that the three novel substrates perform better than the peat substrate (Figure 5d). The nutrient content in the shoots followed a similar pattern. The shoots in the control treatment (SC) exhibited superior performance, demonstrating the most favorable outcomes among all the treatments. The SR22 treatments under solid fertilization exhibited neutral performance, with close clusters within the treatment groups and among the nutrient levels with the peat (SC) substrate (Figure 5b). The situation with shoots treated with liquid fertilizers was extremely contrary. Peat shoots treated with liquid fertilizers showed reduced nutrient content, with UR21 having a higher concentration of virtually all the elements compared to the other liquid treatments except Mg and P (Figure 5e). Again, the roots of beech seedlings showed the highest nutrient content in the SC treatment, with significant levels of C, N, and Mg (Figure 5c). The roots treated with liquid fertilizers exhibited a more competitive performance. The UR22 treatment performs better in the concentration of crucial nutrients like N, P, and K (Figure 5f). For oak leaves and shoots, the SC and SR22 treatments are tightly grouped in the dendrogram, this suggests that the two treatments have similar effects on nutrient concentration in these organs. In particular, Figure 6a,b indicated that the treatments have a consistent impact on the uptake of N, P, and K. The roots of the oak seedlings showed the highest nutrient content in all the treatments except in SR20 (Figure 6c). The situation was closely similar in the seedlings raised with liquid fertilizer (Figure 6f) The dendrogram groupings for nutrient content in the oak roots demonstrate clear differences among the fertilization methods with respective substrates (Figure 6c). The result of correlation analysis indicates a strong interdependence between biomass and nutrient concentration, including N, P, and K. (Figure 7a,b). Beech exhibits a robust positive correlation between biomass production and the availability of all the nutrients (Figure 7b). In contrast, oak exhibits weaker and more complex correlations between biomass and these nutrients (Figure 7b).

3.4. Principal Component Analysis (PCA) Results for F. sylvatica and Q. robur

The PCA results revealed the distribution patterns of nutrient allocation in the root system of F. sylvatica and Q. robur under different fertilization treatments. In both species, the substrate medium had a minimal effect on nutrient composition as, nutrient profiles remained relatively consistent across different substrates, with data points clustering closely together (Figure 8 and Figure 9). This, therefore, implies that variations in the growing medium did not significantly influence nutrient composition in the root system. In contrast, the effect of fertilization treatments (UR and SR) was more pronounced, as illustrated in Figure 10, where data points were more widely dispersed. This separation indicates that fertilization had a significant impact on nutrient content in both the oak and beech seedlings, causing noticeable variation in nutrient allocation at the end of the production cycle. This indicated a significant effect of fertilization on nutrient content for both species, with the points being spatially separated. The variation observed in these graphs suggests that the primary factor influencing growth is the type of fertilizer treatment rather than the growing medium treatment.

4. Effects of Treatments on N and P Allocation in Beech and Oak Seedlings

Among the analyzed nutrients, N and P showed significant responses to different treatments, influencing nutrient allocation in seedling organs. These two nutrients were, therefore, selected for detailed regression analysis to quantify treatment effects more precisely. The regression analysis revealed significant effects of the treatments and plant parts on the N and P concentrations in the F. sylvatica and Q. robur seedlings. For the beech seedlings, the N concentration was significantly reduced across all the treatments compared to the peat substrate. Decreases were observed in the liquid fertilizer treatments (UR22, UR20, and UR21), while the solid fertilizer treatment (SR20, SR21, and SR22) resulted in smaller reductions. Similarly, the phosphorus concentration declined under all the treatments, with the most notable reductions in the liquid fertilizer. Across plant organs, the N and P concentrations were significantly lower in the leaves and shoots, indicating preferential nutrient allocation to the root organ. In the oak seedlings, the nitrogen concentration also showed significant reductions, particularly in the liquid fertilizer, with UC (−37.10 mg g−1) also showing substantial declines. Unlike beech, oak leaves exhibited a positive N allocation (7.25 mg g−1), suggesting preferential retention in the foliage, while shoots showed a reduction. For phosphorus, the largest decreases were again observed in UR21, UR20, and UC. Similarly to beech, the oak seedlings exhibited lower P concentrations in comparison to the reference part (Table 4).

5. Discussion

The results of this study provide insights into how substrate treatments affect the growth (biomass and macronutrient allocation) within and between beech and oak seedlings after one year of growth in a new forest. The results showed significant variation in the nutrient content across treatments, especially for N, P, and K, which are the most crucial nutrients required for the continuous survival of tree seedlings beyond nursery success. In essence, the novel substrate and fertilizer did not exhibit higher biomass and allocation of more nutrients in the studied organs compared to those raised on traditional peat substrate.
The analysis of soil properties revealed favorable conditions for nutrient availability. Although there were slight numerical variations, these differences were not statistically significant (p-value > 0.05). The C content is observed to be higher at the 10 cm depth for both sites, with a subsequent decrease at 20 cm. This is primarily attributed to the greater proportion of organic matter present in the humus-accumulative A horizon. However, as the pedological profile was not opened and the thickness of the A horizon was not determined, the C/N ratio remains stable across depths, indicating balanced C/N cycling within the soil, which is essential for maintaining soil health and organic matter stability [35,36]. This suggests that the variations in soil properties at different depths and sample areas may not significantly impact the growth and survival of the oak and beech seedlings. The overall similarity in soil properties across both species and depths implies that soil factors were not the primary cause of the observed differences in seedling performance. The interaction between environmental factors and nutrient availability plays a pivotal role in shaping seedling performance in the field [37,38]. Soil properties were relatively uniform across the study site, with higher nutrient concentrations in the upper 0–10 cm layer, highlighting the importance of nutrient-rich surface soils, particularly during early root development stages.
The distribution of plant biomass is a crucial factor in nutrient allocation within plant organs [39,40,41,42,43,44]. Previous studies have highlighted the significance of N and P partitioning between plant organs as a pivotal factor in regulating growth rates [45,46,47,48,49,50,51,52,53]. A previous study on the seedlings examined in this investigation, conducted in a nursery setting, indicated that the physical and chemical properties of the novel substrates were not significantly different from those of the peat substrate [15]. However, contrasting trends in nutrient allocation, particularly between N and P, were influenced from negative to positive by fertilization [28]. Consequently, the pronounced variation in seedling response observed after one year in the forest can be attributed to the fertilization methods employed during nursery cultivation.
Both species exhibited comparable responses to varying nutrient availability, yet the magnitude of these responses differed significantly. A shift in biomass allocation from shoots to roots is a well-documented response to nutrient limitation [2,54]. This may enhance the plant’s ability to access soil resources [54] and improve water supply to aboveground parts [8,55]. An increased allocation of biomass belowground and a decreased specific leaf area have been linked to a greater ability to withstand stress, dearth, or cold winter [56,57]. These adaptations can reduce water loss and enhance the seedling’s capacity to access soil moisture [58]. Furthermore, the observed differences in biomass and nutrient allocation between F. sylvatica and Q. robur suggest species-specific strategies in responding to environmental stressors [59,60]. The pronounced allocation of nutrients to belowground organs in oak seedlings may enhance resilience to nutrient limitations and water scarcity, whereas beech seedlings’ preference for aboveground growth reflects an adaptive response to favorable soil conditions.
Both fertilization methods demonstrated a positive impact on both species. This is consistent with the findings of Trubat et al. [8] who reported that fertilization significantly influences nutrient status, aboveground and belowground biomass accumulation, and biomass allocation patterns. However, seedlings raised with the solid fertilizer treatment exhibited higher nutrient concentrations than those raised with liquid fertilizer, which contradicts the initial hypothesis. The substantial variation observed in response to fertilization effects may be attributed to deficiencies of N and P in the novel fertilizer, which resulted in decreases in leaf area, likely lowering transpiration rates and reducing water demands. Trubat et al. [8] additionally observed that nutrient deprivation can enhance the field performance of woody seedlings. They found that the root–shoot ratio was higher in N- and P-deficient seedlings than in those receiving complete nutrient solutions or slow-release fertilizers. Additionally, solid fertilizers may have provided a more even distribution of nutrients within the substrate, while the timing of application with liquid fertilizers may have been less optimal in relation to the seedlings’ growth beyond nursery phases.
Nutrient allocation exhibited considerable variation not only between the two-tree species but also within different organs of each species. This finding is consistent with the findings of various studies on European beech and pedunculate oak [28,42,61,62,63]. In contrast to beech, oak exhibited significantly higher nutrient allocation in roots, shoots, and leaves. This study reaffirmed that nutrient allocation is generally higher in the belowground organs than in the aboveground ones for both species. Notably, the response efficiencies of the studied species to nutrient treatments varied significantly. The R22 treatment of the novel substrate and UAK fertilizer formulation led to greater nutrient accumulation in the roots. This finding is consistent with previous research on the nutrient content of aboveground and belowground biomass [28,39,40,41,42,43,64].
This study revealed a significant interaction between substrate types, fertilization methods, and nutrient allocation in plant organs. It is of paramount importance to fertilize seedlings in order to ensure their vitality and subsequent success following transplantation. Plant roots serve as the primary storage organs for plant nutrients, particularly during periods of dormancy or reduced metabolic activity, as was the case for the studied seedlings. After the growing season in the nursery, emphasis was placed on the transport of nutrients to the roots for storage. This mobility and reallocation of nutrients is likely to have resulted in a greater allocation to underground growth, as observed in this study. Furthermore, the allocation of nutrients to stable plant organs occurred more rapidly in oak than in beech, which is why more nutrients were directed to the underground root growth of oak compared to beech. The significant enrichment of major elements due to higher N fertilizer indicates that plants efficiently accessed and transported substantial amounts of the applied fertilizer to all the organs, especially the root. These results validate the effectiveness of the fertilization treatment during production at the nursery and explain the enhanced growth and productivity driven by these macronutrients, which are crucial for plant growth and development [50,51,65,66]
Numerous studies across various ecological zones have demonstrated that the application of fertilizers over an extended period, spanning several weeks or multiple years, has a beneficial impact on a diverse range of tree species: Larix kaempferi [66,67], Moringa oleifera [68], Khaya senegalensis [57], Eucalyptus torelliana [69], Fagus sylvatica [28,60,70,71,72], and Quercus robus [28,71,73]. Furthermore, the effect of fertilization on container-grown seedlings within the context of this recent modern forestry practice has also been exploited, [72,74,75,76,77]. However, the continuous survival of these seedlings in the forest has remained underexplored [28]. This study highlights the significant enhancement of total plant biomass through fertilization, underscoring the practical importance of this fertilization regime in forestry practice. It not only produces superior container-grown seedlings but also promotes their growth and survival in the forest, exposed to environmental factors.
F. sylvatica allocates a greater proportion of its resources to aboveground growth when the conditions of its root system are favorable in response to changes in the supply of nutrients. In contrast, Q. robur exhibits a more balanced growth strategy, allocating more resources below ground, resulting in a consistent but less steep increase in aboveground biomass. These disparate responses to fertilization methods may be attributed to interspecific differences in nutrient transport and partitioning [78], morphological and anatomical structures [79], and nutrient resorption efficiency [80,81]. Despite the consistent nutrient supply and the absence of variation in forest soil properties, observations from Quercus ilex indicate that seedling demands may be supported by nutrient reserves until the end of the first spring after germination [82]. The differing strengths and patterns of correlations observed in biomass and nutrient concentration between beech and oak are likely reflective of their specific adaptations to the new forest environments.
The results indicate that N and P allocation were significantly influenced by fertilizer treatments, consistent with their key roles in plant metabolism. The strong treatment effects observed for N and P suggest that these nutrients are the primary drivers of seedling response under peat substrate. N and P are well known to be critical for plant growth, particularly in nutrient-limited environments [83,84,85]. Their availability affects biomass accumulation, root growth, and overall survival in seedlings. In this study, the N and P concentrations declined significantly across different treatments, with reductions observed in liquid fertilizer treatments. Unlike previous studies reporting higher N levels as beneficial for seedling growth and survival [86], this study showed a reduction in N allocation across all the treatments, particularly in the seedlings exposed to liquid fertilizer treatments. This pattern suggests a possible limitation in N uptake, potentially affecting long-term seedling establishment. Similarly, while P is widely recognized as a limiting nutrient in forest ecosystems [87,88], our study revealed consistent P reductions under experimental treatments, with liquid fertilizer again exhibiting declines.
The temperate forest environment presents significant challenges for seedling survival due to summer drought, winter frost, and sometimes soil infertility, which can impede successful establishment. Nutrient loading for seedlings has proven to be an effective strategy to alleviate post-planting stresses. Recent studies into the relationship between seedling nutrient levels and out-planting performance has introduced the concept of “nutrient loading” with N. This concept involves the “supercharging” of seedlings with N to improve their survival and growth on forest sites. Nutrient loading involves fertilizing seedlings until their N content reaches levels that meet or exceed their needs. This process has been successful with black spruce (Picea mariana) on sites with heavy plant competition, as reported by Timmer [89], Thomas et al. [90], Villar-Salvador et al. [91], and Lin et al. [92]. The adoption of this concept can aid the performance of the studied species if replicated.

6. Conclusions

This study highlights the significant impact of substrate types and fertilization methods on the biomass and nutrient allocation of Fagus sylvatica L. and Quercus robur L. seedlings after one year of growth in a newly established forest. While traditional peat substrates combined with solid fertilizers yielded the highest overall biomass and nutrient uptake, the novel R22 substrate emerged as a promising alternative, performing comparably in several key metrics. This finding is particularly relevant in the context of sustainable forestry practices, where reducing peat use is increasingly prioritized. The observed species-specific responses emphasize the importance of tailoring forestry practices to the biological characteristics of the target species. The beech seedlings allocated more resources to aboveground growth, reflecting a strategy that could be advantageous in environments with favorable soil conditions. In contrast, the oak seedlings exhibited a more balanced growth strategy, with significant nutrient allocation to belowground organs, which may enhance their resilience in less fertile soils.
Despite the promising performance of the R22 substrate, the study revealed that seedlings treated with solid fertilizers consistently outperformed those treated with novel liquid fertilizers. This suggests that the current formulation of the liquid fertilizer may require optimization, particularly through the addition of essential nutrients like N, to enhance its effectiveness in supporting seedling growth and survival. The findings from this study contribute valuable insights into the development of sustainable forestry practices, particularly in the context of peatland conservation. The R22 substrate, with further refinement, could serve as a viable replacement for peat in the cultivation of forest seedlings. Future research should focus on optimizing liquid fertilizer formulations and exploring the long-term impacts of these treatments on seedling survival and forest establishment.
Although this study primarily focuses on the first year of seedling growth, the promising results observed in biomass accumulation, nutrient allocation, and root development suggest that certain substrates and fertilizer types may have lasting effects on seedling survival and resilience. Given the critical role that early-stage growth plays in determining the long-term success of forest seedlings, these findings provide valuable insights into the potential for sustained growth and establishment over time. For instance, solid fertilizers, which exhibited better performance in terms of biomass and nutrient accumulation, may contribute to stronger root systems and greater overall seedling vigor. This could potentially translate to higher survival rates and improved resilience to environmental stressors, such as drought or nutrient deficiency.
Similarly, while R22 demonstrated comparable performance to peat after one year in the new forest, it is essential to consider its long-term implications. The potential for R22 to sustain seedling growth over multiple years may depend on factors such as its degradation rate and nutrient release. Over time, the accumulation of organic matter and microbial activity in the R22 substrate could further enhance its suitability as a peat alternative; however, it is possible that certain physical or chemical properties may change, affecting the long-term growth of seedlings. Therefore, continuous monitoring of R22’s performance across multiple growth stages will be critical to understanding its long-term viability in forest nursery settings. Additionally, the interplay between substrate and fertilizer types in the long term could have important implications for the health and resilience of seedlings. As seedlings mature, their nutrient requirements and stress tolerance may evolve. This highlights the need for future studies that explore the effects of these variables over extended periods, including the impact of environmental conditions and changes in soil quality over time.

Author Contributions

Conceptualization, O.J.R. and S.M.; Methodology, M.J. and K.S.-S.; Validation, K.S.-S.; Formal analysis, O.J.R. and M.J.; Investigation, M.J.; Data curation, O.J.R. and K.S.-S.; Writing—original draft, O.J.R.; Writing—review & editing, S.M., M.J. and K.S.-S.; Supervision, S.M.; Project administration, S.M.; Funding acquisition, S.M. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding. However, The peat-free substrates and liquid fertilizer used in this study were developed under the project POIR.04.01.04-00-0016/20, which was funded by the National Centre for Research and Development (NCBiR) from national resources and the European Regional Development Fund.

Data Availability Statement

Data is available upon request.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. Study area of the experiment in Barbarka, Miechów Forest District, Poland. (a) Geographic location of the study site. (b) Detailed topographic map of the forest area. (c) Satellite imagery showing the experimental plot layout.
Figure 1. Study area of the experiment in Barbarka, Miechów Forest District, Poland. (a) Geographic location of the study site. (b) Detailed topographic map of the forest area. (c) Satellite imagery showing the experimental plot layout.
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Figure 2. Distribution of biomass across different treatments for Fagus sylvatica L. Alphabets ‘a–c’ denote homogeneous groups under solid fertilization and ‘e’ denote homogeneous groups under liquid fertilization; S—solid fertilization; U—liquid fertilization; R—novel substrates; C—controls substrate (peat–perlite).
Figure 2. Distribution of biomass across different treatments for Fagus sylvatica L. Alphabets ‘a–c’ denote homogeneous groups under solid fertilization and ‘e’ denote homogeneous groups under liquid fertilization; S—solid fertilization; U—liquid fertilization; R—novel substrates; C—controls substrate (peat–perlite).
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Figure 3. Distribution of biomass across different treatments for Quercus robur L. Alphabets ‘a–c’ denote homogeneous groups under solid fertilization and ‘e’ and ‘f’ denote homogeneous groups under liquid fertilization; S—solid fertilization; U—liquid fertilization; R—novel substrates; C—controls substrate (peat–perlite).
Figure 3. Distribution of biomass across different treatments for Quercus robur L. Alphabets ‘a–c’ denote homogeneous groups under solid fertilization and ‘e’ and ‘f’ denote homogeneous groups under liquid fertilization; S—solid fertilization; U—liquid fertilization; R—novel substrates; C—controls substrate (peat–perlite).
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Figure 4. (a–c) correlation among assessed biomass of seedling organs.
Figure 4. (a–c) correlation among assessed biomass of seedling organs.
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Figure 5. Matrix of nutrients in different parts of F. sylvatica seedlings for each treatment (mg/kg). S—State Forest fertilization; U—University fertilization; R—novel substrates; C—control substrate (peat).
Figure 5. Matrix of nutrients in different parts of F. sylvatica seedlings for each treatment (mg/kg). S—State Forest fertilization; U—University fertilization; R—novel substrates; C—control substrate (peat).
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Figure 6. Allocation of nutrients in different parts of Q. robur seedlings for each treatment (mg/kg). S—State Forest fertilization; U—University fertilization; R—novel substrates; C—control substrate (peat).
Figure 6. Allocation of nutrients in different parts of Q. robur seedlings for each treatment (mg/kg). S—State Forest fertilization; U—University fertilization; R—novel substrates; C—control substrate (peat).
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Figure 7. Correlation matrix analysis of nutrient and biomass in F. sylvatica and Q. robur.
Figure 7. Correlation matrix analysis of nutrient and biomass in F. sylvatica and Q. robur.
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Figure 8. Nutrient allocation in beech root grown on different substrates.
Figure 8. Nutrient allocation in beech root grown on different substrates.
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Figure 9. Nutrient allocation in oak root grown on different substrates.
Figure 9. Nutrient allocation in oak root grown on different substrates.
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Figure 10. Nutrient allocation in oak and beech root grown on different fertilization methods.
Figure 10. Nutrient allocation in oak and beech root grown on different fertilization methods.
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Table 1. Properties of the organic peat-free substrate [%].
Table 1. Properties of the organic peat-free substrate [%].
SubstrateSawdustWood Bark PerliteCore Wood Mixed Silage Wood Chips Straw
R20731042110-
R21201042163-
R225033421-10
Table 2. Mean and standard deviation of physicochemical properties of substrates used in seedling growth in the nursery.
Table 2. Mean and standard deviation of physicochemical properties of substrates used in seedling growth in the nursery.
SubstrateWater Capacity (%)Water Outflow Rate (L/min)Bulk Density (g/cm3)Solid Density (g/cm3)Air Capacity (%)Porosity (%)
R2040.5 ± 2.9 b0.595 ± 0.150 b0.115 ± 0.009 a0.64 ± 0.08 a52.1 ± 3.19 c92.6 ± 0.60 d
R2133.1 ± 2.5 d0.781 ± 0.114 a0.098 ± 0.014 c1.74 ± 0.07 a60.8 ± 3.06 a93.6 ± 0.87 c
R2237.8 ± 5.1 c0.594 ± 0.150 b0.104 ± 0.020 b1.66 ± 0.11 a55.8 ± 5.58 b93.9 ± 0.98 b
Control57.7 ± 5.4 a0.417 ± 0.145 c0.085 ± 0.007 d1.69 ± 0.14 a37.0 ± 5.72 d94.7 ± 0.42 a
F387.4556.3265.811.0717295.7976.48
p0.00000.00000.00000.38700.00000.0000
Letters with different alphabet indicate statistically significant differences between means (p < 0.05).
Table 3. Soil properties of sampled plot of F. sylvatica and Q. robur at Barbarka experimental forest of Poland.
Table 3. Soil properties of sampled plot of F. sylvatica and Q. robur at Barbarka experimental forest of Poland.
Exchangeable Cations
Soil Uptake Level (cm)pH (H2O)N C P2O5C/NCaKMgNa
Fagus sylvatica L. site
0–105.21 ± 0.550.36 ± 0.102.41 ± 0.213.14 ± 0.2113.78 ± 0.325.51 ± 0.520.18 ± 0.010.57 ± 0.060.70 ± 0.08
10–205.10 ± 0.570.34 ± 0.091.94 ± 0.152.87 ± 0.1412.90 ± 0.324.81 ± 0.390.16 ± 0.010.42 ± 0.030.63 ± 0.06
Total5.15 ± 0.560.35 ± 0.102.16 ± 0.122.99 ± 0.1213.30 ± 0.234.22 ± 0.320.15 ± 0.010.49 ± 0.030.55 ± 0.07
p-value0.187 ns0.170 ns0.061 ns0.286 ns0.065 ns0.064 ns0.062 ns0.062 ns0.062 ns
Quercus robur L. site
0–105.14 ± 0.590.45 ± 0.952.14 ± 0.224.70 ± 0.4513.31 ± 0.335.11 ± 0.530.23 ± 0.020.55 ± 0.060.81 ± 0.26
10–205.03 ± 0.610.44 ± 0.091.83 ± 0.144.22 ± 0.4012.23 ± 0.314.65 ± 0.360.21 ± 0.020.46 ± 0.030.82 ± 0.27
Total5.08 ± 0.590.45 ± 0.101.55 ± 0.134.44 ± 0.3012.73 ± 0.233.32 ± 0.320.22 ± 0.010.38 ± 0.030.82 ± 0.29
p-value0.208 ns0.945 ns0.061 ns0.427 ns0.069 ns0.061 ns0.363 ns0.061 ns0.989 ns
Mean ± SD; C and N (%); P2O5 (mg/100g) Ca, K, Mg, Na (cmol (+) kg−1) ns = not significance.
Table 4. Modeled estimate of the effects of treatments and plant organs on the N and P concentrations in the beech and oak seedlings.
Table 4. Modeled estimate of the effects of treatments and plant organs on the N and P concentrations in the beech and oak seedlings.
Fagus sylvaticaQuercus robur
SpeciesNutrientTreatment/OrganEstimate (mg g−1)Std. ErrorLower CIUpper CIEstimate (mg g−1)Std. ErrorLower CIUpper CI
BeechN SR20−22.033.29−29.09−14.98−24.374.33−33.66−15.08
SR21−21.473.29−28.52−14.41−20.134.33−29.42−10.84
SR22−12.703.29−19.76−5.65−14.534.33−23.82−5.24
UC−38.173.29−45.22−31.11−37.104.33−46.39−27.81
UR20−41.803.29−48.86−34.75−39.704.33−48.99−30.41
UR21−41.203.29−48.26−34.15−42.574.33−51.86−33.28
UR22−42.873.29−49.92−35.81−39.434.33−48.72−30.14
Leaf−27.262.01−31.58−22.947.252.651.5612.94
Stem−21.332.01−25.65−17.01−10.362.65−16.05−4.67
BeechPSR20−2.900.76−4.54−1.26−3.031.46−6.170.10
SR21−2.400.76−4.04−0.76−3.001.46−6.140.14
SR22−1.330.76−2.970.30−0.971.46−4.102.17
UC−6.000.76−7.64−4.36−6.671.46−9.80−3.53
UR20−6.570.76−8.20−4.93−7.071.46−10.20−3.93
UR21−6.470.76−8.10−4.83−7.301.46−10.44−4.16
UR22−6.430.76−8.07−4.80−6.701.46−9.84−3.56
Leaf−5.400.47−6.40−4.40−1.960.90−3.88−0.04
Stem−3.460.47−4.47−2.46−3.310.90−5.23−1.39
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Rotowa, O.J.; Małek, S.; Jasik, M.; Staszel-Szlachta, K. Substrate and Fertilization Used in the Nursery Influence Biomass and Nutrient Allocation in Fagus sylvatica and Quercus robur Seedlings After the First Year of Growth in a Newly Established Forest. Forests 2025, 16, 511. https://doi.org/10.3390/f16030511

AMA Style

Rotowa OJ, Małek S, Jasik M, Staszel-Szlachta K. Substrate and Fertilization Used in the Nursery Influence Biomass and Nutrient Allocation in Fagus sylvatica and Quercus robur Seedlings After the First Year of Growth in a Newly Established Forest. Forests. 2025; 16(3):511. https://doi.org/10.3390/f16030511

Chicago/Turabian Style

Rotowa, Odunayo James, Stanisław Małek, Michał Jasik, and Karolina Staszel-Szlachta. 2025. "Substrate and Fertilization Used in the Nursery Influence Biomass and Nutrient Allocation in Fagus sylvatica and Quercus robur Seedlings After the First Year of Growth in a Newly Established Forest" Forests 16, no. 3: 511. https://doi.org/10.3390/f16030511

APA Style

Rotowa, O. J., Małek, S., Jasik, M., & Staszel-Szlachta, K. (2025). Substrate and Fertilization Used in the Nursery Influence Biomass and Nutrient Allocation in Fagus sylvatica and Quercus robur Seedlings After the First Year of Growth in a Newly Established Forest. Forests, 16(3), 511. https://doi.org/10.3390/f16030511

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