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Article

Hierarchic Branch Morphology, Needle Chlorophyll Content, and Needle and Branch Non-Structural Carbohydrate Concentrations (NSCs) Imply Young Pinus koraiensis Trees Exhibit Diverse Responses Under Different Light Conditions

by
Bei Li
1,
Wenkai Li
2,
Sudipta Saha
1,
Xiao Ma
1,
Yang Liu
3,
Haibo Wu
1,
Peng Zhang
1,4,* and
Hailong Shen
1,5,*
1
College of Forestry, Northeast Forestry University, Harbin 150040, China
2
Jingyu County Natural Resources and Forestry Bureau, Baishan 134300, China
3
School of Geography and Tourism, Harbin University, Harbin 150086, China
4
Key Laboratory of Sustainable Forest Ecosystem Management-Ministry of Education, Northeast Forestry University, Harbin 150080, China
5
State Forestry and Grassland Administration Engineering Technology Research Center of Korean Pine, Harbin 150040, China
*
Authors to whom correspondence should be addressed.
Horticulturae 2025, 11(7), 844; https://doi.org/10.3390/horticulturae11070844
Submission received: 11 June 2025 / Revised: 12 July 2025 / Accepted: 14 July 2025 / Published: 17 July 2025
(This article belongs to the Section Floriculture, Nursery and Landscape, and Turf)
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Abstract

Research on young trees’ adaptation to shade has predominantly focused on leaf-level responses, overlooking critical structural and functional adaptations in branch systems. In this study, we address this gap by investigating hierarchical branch morphology–physiology integration in 20-year-old Pinus koraiensis specimens across four distinct light conditions classified by photosynthetic photon flux density (PPFD): three in the understory (low light, LL: 0–25 μmol/m2/s; moderate light, ML: 25–50 μmol/m2/s; and high levels of light, HL: 50–100 μmol/m2/s) and one under full light as a control (FL: 1300–1700 μmol/m2/s). We measured branch base diameter, length, and angle as well as chlorophyll and non-structural carbohydrate (NSC) content in branches and needles. Branch base diameter and length were more than 1.5-fold higher in the FL Korean pine trees compared to the understory-grown ones, while the branching angle and ratio in the LL Korean pine trees were more than two times greater than those in the FL trees. As light levels increased, Chlorophyll a and b and total chlorophyll (Chla, Chlb, and Chl) concentrations in the needles all significantly decreased. Starch, glucose, and NSC concentrations in both needles and branches were the highest in the trees under FL and lowest under ML (except for glucose in branches). Understory young P. koraiensis trees morphologically and physiologically adapt to limited light conditions, growing to be more horizontal, synthesizing more chlorophyll in needles, and attempting to increase their light-foraging ability. We recommend gradually expanding growing spaces to increase light availability for 20-year-old Korean pine trees grown under canopy level.

1. Introduction

Mixed forests exhibit higher productivity and greater resilience to global climate change than pure forests [1]. Replanting trees in selectively harvested sites with abundant pre-regenerated young trees is a common and effective way to establish mixed forests [2]. However, trees that have regenerated naturally or been replanted often find themselves under shaded conditions beneath the canopy.
Plants growing under the canopy always show plastic phenotypic responses to different light conditions [3]. The plasticity exhibited by plants during shoot development is remarkable, as it allows them to adapt to various harmful external and internal conditions in order to survive and thrive. Moreover, shoot branching has been shown to be significantly affected by phytohormones, nitrogen, light, and sugars [4]. The pattern exhibited during shoot branching is controlled by axillary bud development, which is also significantly reliant on environmental conditions, especially light [5]. For instance, far-red light inhibits lateral bud growth in tomato [6], and this form of light is much more abundant under the canopy [7]. Branching frequency, branch base diameter, the physiological and biochemical properties of plant branches, and leaf morphology are all affected by light intensity [8]. The live crown ratio and maximum and average branch diameters of spruce (Picea glauca) mixed with aspen (Populus tremuloides) significantly decreases with an increase in aspen density (implying a decrease in light intensity) [9]. Plants modulate branch growth in response to light availability. However, quantitative studies on light-mediated branch development remain limited, despite the critical role of non-vertical growth angles in roots and shoots for efficient above- and below-ground resource capture [10]. Furthermore, it remains unclear how young conifers growing in the field change their branching patterns to adapt to a constantly changing light environment.
Alterations in plant branching patterns subsequently affect the leaves growing on these branches. Shifts in branch angle alter light interception by leaves, thereby driving plasticity in chlorophyll content. Chlorophyll is an important pigment in leaves that is involved in photosynthesis and plays a crucial role in absorbing, transmitting, and converting solar energy into electrochemical energy. Furthermore, optimal photosynthesis and proper leaf development rely on the spatiotemporal regulation of chlorophyll metabolism [11]. In many species, it has been found that shaded leaves have higher chlorophyll concentrations than those grown under sunlight [12]; however, the relationship between chlorophyll composition (e.g., the ratio of chlorophyll a to b) and light gradients in relation to young conifers is not well-defined.
NSCs, mainly composed of starch and soluble sugar, are important energy supply materials during plant growth and metabolism. Shade significantly impacts NSC concentrations in woody species’ leaves, so NSC storage may help trees survive long periods in low-light environments [13,14,15]. Additionally, regarding Cunninghamia lanceolata and Schima superba, it has been found that both species show a decrease in NSC, soluble sugar, and starch content with an increase in shade [16]. These research results indicate that plants in shaded environments are subject to various light conditions generated by surrounding plants, leading to diverse morphological and physiological responses [10,17,18]. In addition, research has revealed that a novel rice tiller angle mutant could regulate starch biosynthesis in gravity-sensing cells [19], indicating that starch is related to branch architecture. Understanding these response patterns and their biological mechanisms is crucial for promoting the growth of understory trees and precisely regulating the relationship between the target trees and their surrounding plants and habitat factors.
The Korean pine (Pinus koraiensis Sieb. et Zucc.) is a valuable evergreen arbor species prized for its high-quality timber, nutritious edible nuts, and ecological significance as the climax species in zonal mixed forests containing broad-leaved species in the humid areas of the middle temperate zone in China [2,20,21,22]. However, the primary Korean pine and broad-leaved mixed forests have been seriously damaged and almost entirely transformed into secondary forests lacking Korean pine seed sources or overharvested forests, with fewer—and younger—Korean pine trees compared to the original forests [22,23,24]. These secondary forests can be transformed into near-climax mixed forests again by introducing Korean pine trees through the “planting conifers and preserving broad-leaved trees” approach, especially in areas lacking Korean pine seed sources [23,25,26].
Commonly, the Korean pine trees in primary mixed forests undergo a lengthy growth period of 80–120 years under the canopy before reaching the upper layer. The planted Korean pine trees in secondary forests and the naturally regenerated Korean pine trees in overharvested forests can also undergo an under-canopy growth period before ascending to the upper canopy [21]. Throughout this process, under-canopy light conditions can significantly affect the growth of the planted or naturally regenerated Korean pine trees until they become the upper-layer trees. A light environment created by surrounding trees has been proven to be the critical ecological factor influencing the growth of Korean pine trees before they ascend to the upper layer [27,28].
Korean pine seedlings display considerable plasticity in needle morphology, tree height, and crown growth in response to varying light intensities, with needle chlorophyll content increasing with canopy openness and needle NSC levels decreasing as light intensity decreases [29].
While previous research has yielded valuable insights, three main limitations exist: (1) the focus has been on using seedlings as testing materials; (2) predominantly stand-level research scales have been employed; and (3) there is a scarcity of data obtained from trees that have undergone long-term field acclimation to realistic light regimes. Consequently, linkages between branch architecture, light-harvesting efficiency, and downstream physiological processes (chlorophyll metabolism, NSC allocation, etc.) remain unresolved at the individual-tree level, hindering precision silviculture.
In this study, we aimed to address these limitations by selecting young Korean pine specimens (about 20 years old) that had grown for a long period under the canopy of a secondary forest composed of naturally regenerating broadleaved trees as the target trees; selecting scarcely researched morphological and physiological indices that have not yet been proven to be important; and elucidating the response mechanisms of the developmental and physiological characteristics of P. koraiensis trees in adapting to different light environments. Our main aim was to clarify the following issue: How does an individual 20-year-old Korean pine’s branching architecture adapt to understory low light? Will this architecture, in turn, influence the light-foraging strategy for needles through chlorophyll synthesis, NSC content, and storage? We hypothesized that young conifers growing in the understory would exhibit a different branching architecture and coordinated set of chlorophyll and NSC production adjustments in response to variations in light availability. This work will provide further theoretical references and a scientific basis for determining the optimal light environment for the growth of P. koraiensis trees under the canopy and addressing issues related to the sluggish growth and slow regeneration of P. koraiensis trees under deciduous broad-leaved canopies.

2. Materials and Methods

2.1. Study Site

The Maoershan Experimental Forestry Farm of Northeast Forestry University (45°23′~45°26′ N, 127°34′~127°39′ E) is situated in Maoershan Town, Shangzhi City, Heilongjiang Province. This region has a temperate continental monsoon climate with a growing season spanning from May to September. The annual mean air temperature is 3 °C, ranging from −19.7 °C in January to 22 °C in July. There are around 120–140 days with no frost on an annual basis, and the average amount of precipitation is between 600 and 800 mm, primarily (80%) falling from May to September, with 2471.3 h of annual sunshine [2,21,23].

2.2. Research Design and Sampling

This study focused on understory Korean pine trees obtained from permanent plots established in 2012. The Korean pine trees were planted in 1989 and replenished in 2003 at micro-sites where Korean pine trees were scarce [23]. The trees used in this experiment were the ones that had been replenished and were 20 years old when they were sampled with three replicates. Plot information is given in Table 1.
Based on the photosynthetic photon flux density (PPFD) measured using a dual radiometer (Spectrum Technologies, Inc., Aurora, IL, USA), we categorized under-canopy light environments into three levels, namely, low light (<25 μmol/m2/s, LL), moderate light (25–50 μmol/m2/s, ML), and high light levels (50–100 μmol/m2/s, HL), employing full light (1300–1700 μmol/m2/s, FL) as a control. Under each light condition, three Korean pine trees exhibiting vigorous growth that were mutually similar in height and diameter at breast height (DBH) were selected as sample trees. We investigated branch morphology, chlorophyll content, and NSC concentrations using these sample trees. In mid-June, during the peak growth period, healthy branches were selected from the upper half of the mid-third of the crown of trees to standardize developmental stage and light exposure history. These branches were cut and placed in sterile polythene bags before being transported to the laboratory in a box with ice bags for a laboratory experiment.

2.3. Measurement of Branch Morphology and Needle Chlorophyll Concentrations

The DBH, height, and health statuses were similar among the sampled trees (for more information, please refer to Table 2). We measured the morphological traits of the P. koraiensis branches and needles of the topmost whorl (with current-year and 2-year-old branches and needles), including the number of branches, branch base diameter, branch length in two years, and branching angle. The branching ratio (OBR) can be used to quantify topological hierarchy, as it correlates strongly with carbon allocation efficiency in conifers; it was calculated using Formula (1).
OBR = (Nt − Ns)/(Nt − N1)
where Nt is the total number of branches in all branch-class levels; Ns is the number of branches at the last branch-class level; and N1 is the number of branches at the first branch-class level. The first-class branches are attached to the main stem, while the secondary class branches are attached to the first branch, and the rest of the branches are attached in the same manner; i.e., branches directly attached to the main stem were considered class 1 branches, and lateral branches attached to class 1 branches were considered level 2 branches (this scheme is clearer in Figure 1).
The quantities of level 1, level 2, and total branches of the sampled P. koraiensis trees under FL were 111, 479, and 590, and those of the P. koraiensis trees under LL, ML, and HL were 98, 213, and 311; 88, 215, and 303; and 108, 251, and 359, respectively. Branch base diameter, branch length, and branching angle were measured for the main stem tip branch and top whorl branches.
Needles were sampled from the upper one-third of the crown of the young P. koraiensis trees, mainly in the morning from 8:00 to 11:00 on 18 June 2021. Fresh annual needles were cleaned and weighed to 0.2 g. Then, they were cut up, placed in a 50 mL centrifuge tube, extracted with 40 mL of a 1:1 mixture of 99.5% acetone and 95% ethanol by volume at 10–25 °C, and protected from light until the material decolorized and turned white. The absorbance value was measured using a spectrophotometer, and the chlorophyll content was calculated according to the method described by He J et al. [30].

2.4. Measurement of Non-Structural Carbohydrate Concentrations

The samples were oven-dried to a constant weight at 65 °C for at least 72 h, ground several times, sieved through a 100-mesh (0.15 mm aperture) sieve, pulverized using a SCIENTZ-48L pulverizer (Ningbo Scientz Biotechnology Co., Ltd., Ningbo, China), and stored in plastic bags for further measurement. The non-structural carbohydrate (NSC) content was determined as the sum of soluble sugars (SSs) and starch (ST) content, which, in turn, was determined using the anthrone colorimetric method according to Hansen and Møller [31].

2.5. Statistical Analysis

All the measurements were conducted using three biological replicates. All the data were subjected to ANOVA to search for statistically significant differences between different light conditions. We used the Least-Significant Difference test as a post hoc test (p < 0.05). Pearson’s correlation matrix was used to find the correlation coefficients (r) of all pairwise combinations. Data analysis and graph generation were conducted using R 4.2.2 software.

3. Results

3.1. Branch Morphology and Needle Chlorophyll Content of Pinus koraiensis Under Different Light Conditions

3.1.1. Branch Characteristics of Pinus koraiensis

There was no significant difference in the branch base diameters of the Korean pine trees at different light levels in the understory (p > 0.05), but the branch base diameters under HL were relatively higher than those under ML and LL (18% and 19% higher, respectively), whereas they were almost the same under LL and ML. Notably, under FL, the branch base diameter was significantly higher than that under the canopy (p < 0.05—61%, 90%, and 92% higher than that under HL, ML, and LL) (Figure 2A).
The branches of the Korean pine trees under FL were significantly longer than those under the canopy (p < 0.05).
However, compared to what was observed for branch size, the pattern in the response of the branching angles of the Korean pine trees was different (Figure 2C): the branches of the Korean pine trees under LL exhibited a near-flat angle, reaching 74°, which is significantly higher than the angles of the trees under ML, HL, and FL (11°, 15°, and 42° higher, respectively). Additionally, different branching patterns were observed between under-canopy light conditions and full-light conditions. The top whorl branching angles of the Korean pine trees that had adapted to under-canopy light environments over the long term were greater than 45° and tended towards a flat pattern, whereas the angles under full-light conditions were less than 45° and tended towards an erect pattern.
The branching ratios of the low-light Korean pine trees were significantly higher than those of the full-light trees (p < 0.05, Figure 2D).

3.1.2. Needle Chlorophyll Concentrations of Pinus koraiensis

The Chla, Chlb, and Chl concentrations in the Korean pine needles all significantly decreased as light conditions improved (p < 0.05). Chla/Chlb (Chlorophyll a/Chlorophyll b) levels in needles were the highest under FL and significantly higher than that under ML (p < 0.05). As light conditions improved, Chla/Chlb initially declined and then increased (Figure 3D).

3.2. Needle and Branch NSCs of Pinus koraiensis Under Different Light Conditions and Their Relationships

The highest starch, glucose, and NSC concentrations were all found in the Korean pine trees under FL, both in branches and needles (Figure 4). The glucose, starch, NSC, and starch/NSC levels in the needles all decreased and then increased as light conditions improved, with the lowest readings obtained from the needles under ML. An “S” shaped curve was observed for the starch/NSC ratio in branches (Figure 4D).
There were no significant relationships between light and NSC concentrations in needles and branches (Figure 5, p > 0.05). However, light conditions, branch diameter, and branch length were significantly and negatively related to Chla, Chlb, Chl, and Chla/Chlb concentrations (Figure 5, p < 0.05). Branch glucose and NSC concentrations were negatively related to branching ratio but positively related to branch diameter and branch length. Overall, the Chla, Chlb, and Chl concentrations in the Korean pine needles were all negatively and significantly related to light conditions (Figure 5, p < 0.05). Generally, tree morphology and physiology are closely related to light conditions.

4. Discussion

4.1. There Is Some Evidence Showing That Branch Morphological Traits Are Linked with Light-Foraging Strategies

The P. koraiensis trees grown under FL tended to grow better than those in the understory. Our findings also revealed that the branch production rate of first- and second-order branches was greater for the full-light trees than those grown in the understory, a finding consistent with the observations made by Kwon et al. [32] regarding Platycodon grandiflora. The number of branches as well as leaf length and width decreased significantly under low light-intensity conditions. According to Casal and Fankhauser [18], one of a plant’s tactics for avoiding shadow consists of inhibiting branching at the base of the plant and improving elongation of the stem.
In this study, we investigated the morphological indicators of branches of young P. koraiensis trees in different light environments. It was found that the branch angles of the young P. koraiensis branches in shaded environments were significantly higher than those in the full-light environment, suggesting that in environments with insufficient light, young Pinus koraiensis trees can expand the total area of light acquisition by increasing the branch angle and thus obtaining more light. This adaptive response highlights the ability of young P. koraiensis trees to adjust to varying light conditions. Branch growth is phototropic, resulting in different branch patterns in different light environments [33]. In a study on spruce (Picea asperata), it was found that the number of spruce branches decreased significantly with an increase in light intensity after thinning mixed forests [34]. Our results also align with this finding. As light conditions improved, the branches of young P. koraiensis trees gradually changed from being scarce and elongated to numerous and shorter [2,35]. P. koraiensis’s strategy for adapting to changes in external light environments consists of harvesting light by expanding its light interception ability to efficiently use light energy. However, increased horizontal branching may not fully compensate for receiving more light, resulting in shorter and thinner branches. While the sample size used (n = 3 per light treatment) limits the generalizability of some of our statistical inferences, the observed trends align with the findings of previous studies. In future studies, we will prioritize expanded replication.

4.2. Weak Light Use by Needles Could Compensate for Less Light Foraging via Variation in Branch Traits

Chlorophyll, indispensable in the process of photosynthesis, can be used to measure the strength of plant photosynthesis, and chlorophyll synthesis is mainly affected by light [36]. In this study, we observed a gradual decrease in chlorophyll content with an increase in light intensity, a result consistent with the findings reported by Minotta and Pinzauti [37]. Chlorophyll concentrations play a crucial role in the light-capturing ability of leaves, indicating that leaves of understory trees need more chlorophyll to capture light in order to support their growth owing to the limited amount of light available to them [29]. Leaves of shade-tolerant plants, such as Alocasia macrorrhiza, undergo changes in photosynthetic structures, including increases in chlorophyll concentrations, to optimize efficiency in low light [38]. It was also found that the chlorophyll content in A. thaliana plants grown under low light decreased, whereas this decrease was attenuated in C. hirsuta plants [39,40]. Chla/Chlb levels in many species have been found to be higher in leaves exposed to higher levels of light. However, in Arabidopsis thaliana (L.) Heynh. cv. Landsberg erecta, Chla/Chlb levels increase in low and high irradiance ranges and plateau at intermediate irradiance, reflecting the complexity of photosynthetic acclimation to different light levels, a result that aligns with our findings [41]. Furthermore, Chla/Chlb levels in the all-light environment were higher than those in the other light environments, indicating that Chlb predominantly drives chlorophyll changes under varying light conditions, a common adaptation of plants in low-light habitats [42]. Under insufficient light conditions, the proportion of blue-violet light increases, leading to a higher content of chlorophyll b, which absorbs blue-violet light and facilitates the absorption and utilization of weak light by leaves [10]. Research indicates that P. koraiensis seedlings exhibit lower photosynthetic rates in the understory [29], resulting in reduced net assimilation and growth rates under the canopy [43,44].
NSCs fulfil distinct functional roles, including transport, energy metabolism, and osmoregulation, and provide substrates for the synthesis of defense compounds or exchange with symbionts involved in nutrient acquisition or defense. Branches and roots usually contain storage pools that may be mobilized only in small proportions during normal functioning but can make up most of the C source during catastrophic events that occur on decadal/centennial timescales, like severe and repeated herbivory, wind and ice-storm damage (leading to branch breakage), or fire [45]. Other than being one of the photosynthetic products of plants, NSC also reflects their survival strategies [46]. NSC levels in branches and needles varied under different light conditions, with the levels of starch, glucose, and NSC in the trees under FL being higher than those in the trees in the understory. In contrast, among the Korean pine trees grown in the understory, the trees under ML had lower NSC concentrations than those under LL and HL, possibly because the active radial growth and construction of photosynthetic organs in the Korean pine trees grown under ML result in difficulties in accumulating NSC. In contrast, because of environmental constraints in LL conditions, the trees likely reduced their energy consumption to conserve resources, which might be one reason why their NSC levels were higher than those under ML. Carbon storage is important for species that regenerate in persistently shady habitats [47], and carbohydrate storage in stems and roots enhances long-term survival in the shade by enabling seedlings to cope with periods of biotic and abiotic stress [48]. Without root NSC measurements, we cannot draw more comprehensive conclusions regarding whole-tree NSC dynamics in P. koraiensis. Furthermore, light duration and tree age would also affect the experimental outcomes. In future work, we will focus more on conducting controlled experiments on whole-tree NSC dynamics. Moreover, NSC also functions as a source of stored energy and carbon for biosynthesis [49]. Furze et al. [50] showed that branches were the largest reservoir of total NSC. In our study, branches had higher NSC content than needles since NSCs are used to provide C skeletons for structural growth [45].
Furthermore, numerous studies have provided experimental evidence that C storage is an active process that allows the buffering of environmental fluctuations and supports long-term plant growth [49]. Under FL, P. koraiensis displayed greater vitality and put more resources into active development rather than storage, as evident from the higher glucose and NSC content in the branches and the lower starch/NSC ratio. Starch is inactive and accumulates as a storage compound when photosynthetic supply exceeds growth demand [13,51]. The Korean pines under FL had higher starch/NSC levels in their needles, indicating their higher photosynthetic rates served to address more than growth demand, allowing for starch accumulation for future active growth. In contrast, the Korean pines under ML and HL had lower starch/NSC ratios than those under LL, indicating more active growth since higher light intensity usually brings vigorous growth [52]. Plants rely on both newly assimilated carbon and stored reserves of NSC for growth and other physiological functions, such as respiration, osmotic regulation, and defense [53]. Previous research has shown that NSCs are transferred from mature bamboo to young shoots, facilitating their fast growth [54]. Light is one indispensable factor for NSC content, composition, and distribution. As research has shown, as canopy openness increases, NSC levels in Korean pine needles also increases [29]. Based on the NSC results, we recommend changing the growth environment (to increase PPFD over 50 μmol/m2/s) to ensure Korean pine trees receive more light under moderate light conditions.

5. Conclusions

In general, young P. koraiensis trees growing under full-light conditions have thicker, longer branches and a more horizontal branching pattern than those growing under a canopy. Besides having a flatter branching pattern, Korean pine trees that grow in the understory also have higher Chla, Chlb, and Chl content; lower Chla/Chlb levels; and lower NSC content than those grown under full-light conditions. Relying on these morphological and physiological changes, understory Korean pine trees can attempt to grow better under a light-limited environment. Due to insufficient light, the Korean pines under both ML and LL conditions exhibited growth limitation. Based on our findings, we recommend gradually improving light conditions for 20-year-old understory Korean pines through forest management. However, given the limitations of the current experimental results, the development of more specific management protocols still requires further supporting data, including physiological indicators across different age groups under varying forest canopies and in distinct functional organs of Korean pines.

Author Contributions

W.L., X.M., B.L., and H.S. conceived the entire study; B.L., X.M., Y.L., and W.L. performed the experiment and performed the statistical analyses, created figures, and wrote the entire manuscript; S.S. revised the manuscript; S.S., H.W., P.Z., and H.S. conducted the final revision. All authors have read and agreed to the published version of the manuscript.

Funding

This work was funded by the National Key Research and Development Program of China (Grant No. 2022YFD2201002), the Fundamental Research Funds for the Central Universities (Grant No. 2572023CT02), and the National Natural Science Foundation of China (Grant No. 31972950 and No. 32271857).

Data Availability Statement

The datasets generated during and/or analyzed in this study are available from the corresponding author on reasonable request.

Acknowledgments

The authors would like to thank Maoershan Experimental Forest Farm of Northeast Forestry University for providing access to the study area for this research.

Conflicts of Interest

The authors declare there are no conflicts of interest.

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Horticulturae 11 00844 g001
Figure 1. Simple schematic figure depicting the branch system of Korean pines, considering branch level and branch angle. This system has more than 2 levels, but for illustrative purposes, we have depicted only two levels of branches.
Figure 1. Simple schematic figure depicting the branch system of Korean pines, considering branch level and branch angle. This system has more than 2 levels, but for illustrative purposes, we have depicted only two levels of branches.
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Figure 2. Differences in the branch traits of level 1 branches of Pinus koraiensis under different light intensities: (A) differences in branch diameter, (B) branch length, (C) branching angle, and (D) branching ratio of level 1 branches of Pinus koraiensis under different light intensities. LL, low light; ML, moderate light; HL, high light levels; FL, full light. Different lowercase letters indicate significant differences among different light conditions, with p < 0.05 according to one-way ANOVA and an LSD post hoc test; non-significant results were omitted. Values are means ± SE, with n = 3 biological replicates.
Figure 2. Differences in the branch traits of level 1 branches of Pinus koraiensis under different light intensities: (A) differences in branch diameter, (B) branch length, (C) branching angle, and (D) branching ratio of level 1 branches of Pinus koraiensis under different light intensities. LL, low light; ML, moderate light; HL, high light levels; FL, full light. Different lowercase letters indicate significant differences among different light conditions, with p < 0.05 according to one-way ANOVA and an LSD post hoc test; non-significant results were omitted. Values are means ± SE, with n = 3 biological replicates.
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Figure 3. Comparisons of chlorophyll concentrations in Pinus koraiensis needles under different light intensities: (A) chlorophyll a (Chla), (B) chlorophyll b (Chlb), (C) total chlorophyll (Chl), and (D) chlorophyll a/chlorophyll b (Chla/Chlb) concentrations in Pinus koraiensis needles under different light intensities. LL, low light; ML, moderate light; HL, high light levels; FL, full light. Different lowercase letters indicate significant differences among different light conditions, with p < 0.05 according to one-way ANOVA and an LSD post hoc test; non-significant results were omitted. Values are means ± SE, with n = 3 biological replicates.
Figure 3. Comparisons of chlorophyll concentrations in Pinus koraiensis needles under different light intensities: (A) chlorophyll a (Chla), (B) chlorophyll b (Chlb), (C) total chlorophyll (Chl), and (D) chlorophyll a/chlorophyll b (Chla/Chlb) concentrations in Pinus koraiensis needles under different light intensities. LL, low light; ML, moderate light; HL, high light levels; FL, full light. Different lowercase letters indicate significant differences among different light conditions, with p < 0.05 according to one-way ANOVA and an LSD post hoc test; non-significant results were omitted. Values are means ± SE, with n = 3 biological replicates.
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Figure 4. Comparison of non-structural carbohydrates in branches and needles of Pinus koraiensis under different light intensities: (A) starch, (B) glucose, (C) NSC (starch + glucose), and (D) starch/NSC concentrations in branches and needles of Pinus koraiensis under different light intensities. LL, low light; ML, moderate light; HL, high light levels; FL, full light. Different lowercase letters indicate significant differences between different light conditions, with p < 0.05 according to one-way ANOVA and an LSD post hoc test; non-significant results were omitted. Values are means ± SE, with n = 3 biological replicates.
Figure 4. Comparison of non-structural carbohydrates in branches and needles of Pinus koraiensis under different light intensities: (A) starch, (B) glucose, (C) NSC (starch + glucose), and (D) starch/NSC concentrations in branches and needles of Pinus koraiensis under different light intensities. LL, low light; ML, moderate light; HL, high light levels; FL, full light. Different lowercase letters indicate significant differences between different light conditions, with p < 0.05 according to one-way ANOVA and an LSD post hoc test; non-significant results were omitted. Values are means ± SE, with n = 3 biological replicates.
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Figure 5. Pearson correlation matrix of the correlation between different parameters (p < 0.05). Non-significant correlations were omitted (p > 0.05). Negative and positive correlations are given in red and green, respectively; the color gradient shows the strength of the correlation. N indicates needle traits, while B indicates branch traits.
Figure 5. Pearson correlation matrix of the correlation between different parameters (p < 0.05). Non-significant correlations were omitted (p > 0.05). Negative and positive correlations are given in red and green, respectively; the color gradient shows the strength of the correlation. N indicates needle traits, while B indicates branch traits.
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Table 1. Location and community composition of sampling sites.
Table 1. Location and community composition of sampling sites.
Light LevelsLongitude (E)Latitude (N)Altitude (m)Tree Species Composition
LL127.53°45.32°312Ulmus pumila L., Fraxinus mandshurica Rupr., Quercus mongolica Fisch. ex Ledeb., Acer pictum Thunb. subsp. mono (Maxim.) H. Ohashi, Tilia amurensis Rupr., Juglans mandshurica Maxim., Phellodendron amurense Rupr., A. tegmentosum Maxim., Betula platyphylla Suk., A. tataricum subsp. ginnala (Maxim.) Wesmael
ML127.53°45.32°319U. pumila L., F. mandshurica Rupr., Q. mongolica Fisch. ex Ledeb., A. pictum Thunb. subsp. mono (Maxim.), Pinus koraiensis, Tilia mandshurica Rupr. et maxim., Populus davidiana Dode, P. koreana Rehd., B. platyphylla Suk.
HL127.53°45.32°294U. pumila L., F. mandshurica Rupr., Q. mongolica Fisch. ex Ledeb., A. pictum Thunb. subsp. mono (Maxim.), T. amurensis Rupr., Juglans mandshurica Maxim., P. koraiensis, P. amurense Rupr., Syringa reticulata subsp. amurensis (Rupr.) P. S. Green & M. C. Chang, Styphnolobium japonicum (L.) Schott.
FL127.57°45.27°300U. pumila L., F. mandshurica Rupr., Q. mongolica Fisch. ex Ledeb., A. pictum Thunb. subsp. mono (Maxim.), Tilia amurensis Rupr., J. mandshurica Maxim., P. koraiensis, P. amurense Rupr., B. platyphylla Suk., Aralia elata (Miq.) Seem.
Table 2. Basic information on the sampled trees (n = 3).
Table 2. Basic information on the sampled trees (n = 3).
Light
Levels		PPFD
(μmol/m2/s)		Height
(m)		DBH
(cm)		Tree Age
(Yr)
LL<252.48 ± 0.24 b2.02 ± 0.46 b22
ML25–502.40 ± 0.05 b1.84 ± 0.34 b22
HL50–1002.38 ± 0.16 b1.79 ± 0.45 b22
FL1300–17003.21 ± 0.08 a4.47 ± 0.85 a20
Values in tables are displayed as means ± sd (n = 3); different letters indicate differences between the four light conditions (p < 0.05). LL, low light; ML, medium light; HL, high light levels; FL, full light; DBH, diameter at breast height.
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Li, B.; Li, W.; Saha, S.; Ma, X.; Liu, Y.; Wu, H.; Zhang, P.; Shen, H. Hierarchic Branch Morphology, Needle Chlorophyll Content, and Needle and Branch Non-Structural Carbohydrate Concentrations (NSCs) Imply Young Pinus koraiensis Trees Exhibit Diverse Responses Under Different Light Conditions. Horticulturae 2025, 11, 844. https://doi.org/10.3390/horticulturae11070844

AMA Style

Li B, Li W, Saha S, Ma X, Liu Y, Wu H, Zhang P, Shen H. Hierarchic Branch Morphology, Needle Chlorophyll Content, and Needle and Branch Non-Structural Carbohydrate Concentrations (NSCs) Imply Young Pinus koraiensis Trees Exhibit Diverse Responses Under Different Light Conditions. Horticulturae. 2025; 11(7):844. https://doi.org/10.3390/horticulturae11070844

Chicago/Turabian Style

Li, Bei, Wenkai Li, Sudipta Saha, Xiao Ma, Yang Liu, Haibo Wu, Peng Zhang, and Hailong Shen. 2025. "Hierarchic Branch Morphology, Needle Chlorophyll Content, and Needle and Branch Non-Structural Carbohydrate Concentrations (NSCs) Imply Young Pinus koraiensis Trees Exhibit Diverse Responses Under Different Light Conditions" Horticulturae 11, no. 7: 844. https://doi.org/10.3390/horticulturae11070844

APA Style

Li, B., Li, W., Saha, S., Ma, X., Liu, Y., Wu, H., Zhang, P., & Shen, H. (2025). Hierarchic Branch Morphology, Needle Chlorophyll Content, and Needle and Branch Non-Structural Carbohydrate Concentrations (NSCs) Imply Young Pinus koraiensis Trees Exhibit Diverse Responses Under Different Light Conditions. Horticulturae, 11(7), 844. https://doi.org/10.3390/horticulturae11070844

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