Abstract
Ancient trees represent a significant ecological niche, constituting a vital habitat for a variety of species. Moreover, they serve as a cultural heritage and are worthy of preservation. The systematic analysis of volatile organic compounds (VOCs) in the wood of 11 species of ancient trees in Sichuan Province was performed using headspace gas chromatography–mass spectrometry (GC–MS). The results revealed significant compositional variations were demonstrated across different species of ancient trees. This finding indicates species-specific chemical defense mechanisms that have evolved in response to environmental adaptation. For instance, Magnolia sargentiana exhibited a preference for terpene-based preservation, whereas Acer grosseri demonstrated a preference for lipid-derived aldehydes. Terpenoids, which are the dominant constituents (e.g., α-cedrene and β-cedrene in Magnolia sargentiana and hexanal in Acer grosseri), demonstrate antimicrobial, insect-repellent and ecological signaling functions. These findings contribute to the advancement of knowledge regarding the role of VOCs in woods in ecological interactions and lay the foundation for the development of natural antimicrobial, flavorant, and medicinal products.
Download PDF
Full Article
Composition Characteristics of Volatile Organic Compounds in Different Tree Species and their Chemical Ecological Functions
Ye Hu ,a,b,c,d,*,† Yuzhi Pang
,a,e,† Shiji Jiaba
,a,e,† Chunxia Peng
,a,e,† Xiang Nong
,a,b,c,d Xuan Liu
,g Jian Li
,f Hao Yang
,f Wei Liu
,a,b,c,d Xinyu Wei
, a,b,c,d and Jie Xiao
a,b,c,d
Ancient trees represent a significant ecological niche, constituting a vital habitat for a variety of species. Moreover, they serve as a cultural heritage and are worthy of preservation. The systematic analysis of volatile organic compounds (VOCs) in the wood of 11 species of ancient trees in Sichuan Province was performed using headspace solid-phase microextraction coupled with gas chromatography–mass spectrometry (HS-SPME-GC–MS). The results revealed significant compositional variations across different ancient trees species. This finding indicates species-specific chemical defense mechanisms that have evolved in response to environmental adaptation. For instance, Magnolia sargentiana exhibited a preference for terpene-based preservation, whereas Acer grosseri demonstrated a preference for lipid-derived aldehydes. Terpenoids, which are the dominant constituents (e.g., α-cedrene and β-cedrene in Magnolia sargentiana and hexanal in Acer grosseri), demonstrate antimicrobial, insect-repellent and ecological signaling functions. These findings contribute to the advancement of knowledge regarding the role of VOCs in woods in ecological interactions and lay the foundation for the development of natural antimicrobial, flavorant, and medicinal products.
DOI: 10.15376/biores.21.2.4780-4791
Keywords: Volatile organic compound; Heartwood; GC‒MS analysis
Contact information: a: Bamboo & Forest Institute of Science, Technology and Industrial Innovation, Leshan Normal University, Leshan 614000, China; b: Mabian Dafengding Bamboo Forest Ecosystem Provincial Observation and Research Station, Leshan Normal University, Leshan 614000, China; c: Provincial Engineering and Technology Research Center for Innovative, Development of Bamboo Fiber Nutrition, Leshan Normal University, Leshan 614000, China; d: Bamboo Resource Conservation and Utilization Key Laboratory of Sichuan Province, Leshan 614000, China; e: School of Life Science, Leshan Normal University, Leshan 614000, China; f: Laojunshan National Nature Reserve Conservation Center, Yibin 644000, China; g: School of Economics and Management, Leshan Normal University, Leshan 614000, China; *Corresponding author: fuscahuye@163.com; †These authors contributed equally to this work and share first authorship
INTRODUCTION
Volatile organic compounds (VOCs) are important components of secondary metabolites in plants and play a pivotal role in the survival and competitive dynamics of plant species by modulating the ‘plant–environment’ interaction network. (Hu and Zhang 2022). Research has demonstrated that plant VOCs can mediate a variety of ecological functions. These include antibacterial defence (e.g., α-pinene inhibits fungal spore germination), pest repellence (e.g., β-caryophyllene deters coleopteran pests), and allelopathic suppression (e.g., methyl jasmonate regulates the growth of neighboring plants) (Rashaduz et al. 2024; Tays et al. 2025). The ecological functions of these organisms are facilitated by the secretion of bioactive molecules, including terpenes and aldehydes. In recent years, innovations in gas chromatography–mass spectrometry (GC–MS), particularly the synergistic application of static headspace sampling and comprehensive two-dimensional gas chromatography (GC × GC), have significantly enhanced the detection sensitivity and compositional resolution of trace volatile organic compounds (VOCs) (Yu et al. 2020). This advancement has furnished substantial technical assistance for research endeavors concerning the mechanisms of chemical interaction in plants (Sotaro and Akiko 2024).
Research has focused predominantly on the functions of VOCs in above-ground plant organs, such as leaves and flowers—for instance, their role in attracting pollinators—while the compositional characteristics and ecological adaptability of wood-derived VOCs have received insufficient attention (Loreto and Schnitzler 2010). Because of prolonged environmental adaptation, the VOCs present in the wood of plants exhibit characteristics such as prolonged release cycles and high compositional stability. They might form unique chemical defense systems through terpenoid metabolites (e.g., cedrene) and phenolic derivatives (e.g., lignin degradation products) (Pichersky and Gershenzon 2002). Notably, while research on VOCs in coniferous species (e.g., Pinus massoniana and Picea asperata) has been relatively extensive, the VOCs diversity, functional differentiation, and resource potential of broadleaved tree species—particularly rare and endangered ones—remain to be systematically analyzed (Yan et al. 2020).
Sichuan Province’s abundant rainfall, diverse soil and rock types, and rugged terrain create an ideal environment for a wide variety of plant species (Li et al. 2021). Sichuan Province is located within the biodiversity hotspot of Southwest China and is home to national-level protected plants, with Davidia involucrata serving as a notable example. Old forests containing ancient trees are essential ecosystems for life on earth (Gilhen-Baker et al. 2022). The chemical diversity of VOCs in ancient tree wood might be closely related to its specific ecological adaptability (Meritxell et al. 2021). However, owing to limitations in technical approaches and the complexity of research subjects, studies on the VOCs in the wood of broadleaved tree species in this region remain scarce. Specifically, in addition to sporadic reports on Davidia involucrata and Populus koreana, VOCs of most species (e.g., Fagus lucida and Acer flabellatum) remain unclear (Shen et al. 2024). The extant research in this field has focused primarily on the identification of components, with a paucity of quantitative correlation analyses between terpenoid markers (e.g., α-cedrene) and ecological functions (e.g., plant insect resistance) (Turlings and Erb 2018). Conventional steam distillation techniques frequently resulted in the degradation of heat-sensitive components, such as furans. This complicated the precise reflection of wood VOCs (Ninkuu et al. 2021).
Based on these findings, the present study investigated the compositional differences of VOCs in the wood of 11 broadleaved tree species. These species, including a national first-class protected plant, were selected from Sichuan Province. This study employed headspace solid-phase microextraction coupled with gas chromatography–mass spectrometry (HS–SPME–GC–MS) to systematically analyze VOCs. The objective of this study was to address the following scientific inquiries: First, the objective was to identify the distinctive components of VOCs in the wood of diverse broadleaved tree species and to elucidate their chemotaxonomic significance. Second, the investigation addressed the ecological functions (e.g., antibacterial and insect-repellent properties) and concentration thresholds of key terpenoid compounds (e.g., cedrene and cedrol). Third, the resource utilization potential of rare VOCs in wood was assessed (e.g., prioritized for developing natural preservatives) (Schreiner et al. 2018; Zeng et al. 2021).
EXPERIMENTAL
Materials
The experimental samples were collected from 11 types of wood in Sichuan Province. Tree cores with a diameter of 1 cm and a length of 20 cm were drilled at breast height using an increment borer.
The wood samples were sourced from ancient trees. The method of morphological identification was utilized by Professor Hu Chao for species identification. The methods of core sampling, historical documents, and archival records were utilized by Professor Xie Jiulong for tree age determination. These include samples from Davidia involucrata, a nationally protected first-class plant (from a 170-year-old tree); Fagus lucida, a precious species of Fagaceae (100-year-old tree); Carpinus fangiana, a species of Betulaceae (from a 120-year-old tree); Magnolia sargentiana, a rare species of Magnoliaceae (160-year-old tree); Cinnamomum jensenianum, a common economic plant of Lauraceae (120-year-old tree); Camellia oleifera, an oil plant of Theaceae (150-year-old tree); Enkianthus ruber, an ornamental plant (110-year-old tree); and Acer flabellatum, a common landscaping tree species with a 130-year-old tree. Sorbus folgneri (an ornamental Rosaceae plant with a 190-year-old tree), Lindera limrichtii (an ornamental Lauraceae plant with a 100-year-old tree), and Prunus conradinae (an ornamental Rosaceae plant with a 130-year-old tree) were also analyzed.
Methods
The wood samples were promptly ensconced in hermetically sealed bags, meticulously labeled with the time and location of collection, altitude, and pertinent information, and expeditiously conveyed to the laboratory at a temperature of 4 °C. The samples were placed in a constant-temperature drying oven (DHG-9070A) and dried at 60 °C until they reached a constant weight. This process required approximately 72 h. A plant grinder (RT-34) was subsequently used to grind the dried samples into wood powder, with particle sizes ranging from 80- to 100-mesh. The ambient temperature was maintained at 25 °C, and the relative humidity was maintained at 40% throughout the process. The wood powder samples were subsequently meticulously portioned into brown, wide-mouth bottles, hermetically sealed, and stored in a desiccator for subsequent use.
This study employed HS–SPME–GC–MS to systematically analyze VOCs. Two grams of wood powder were taken from a brown wide-mouth bottle into a headspace vial. The vials were hermetically sealed, subsequently placed in a headspace sampler, and subjected to a heating and equilibration process for a designated period. The headspace gas was subsequently sampled for one min and analyzed by gas chromatography‒mass spectrometry (GC‒MS).
The chromatographic column utilized was an MI-5STUS (30 m × 0.25 mm × 0.25 μm). The inlet temperature was set to 250 °C. The temperature program was as follows: the initial temperature was held at 40 °C for two min, then increased to 100 °C at a rate of 5 °C per min, and finally increased to 165 °C at a rate of 4 °C per min and held for two min. Helium was utilized as the carrier gas. The column flow rate was set to 0.80 mL/min. The injection volume was 1 microliters. The septum purge flow rate was 6 milliliters per min (mL/min), and the mass spectrometer (MS) employed an electron ionization (EI) source. The detector voltage was set to 0.90 kV. The ion source temperature was set to 230 °C. The interface temperature was set to 200 °C. The solvent delay time was set to 5.00 min. The detector gain was set to 1.50 kV + 0.40 kV.
The NIST17-1.lib database was utilized to identify volatile components with a match degree of 90% or higher in conjunction with the literature for qualitative analysis. The relative content of each volatile component was calculated using the peak area normalization method. The generation of GC‒MS spectra was carried out using Origin Pro 8.1 software.
RESULTS AND DISCUSSION
Compositional Characteristics of the Main Volatile Components in the Wood of Ancient Trees
An analysis of the compositional characteristics of volatile components in the wood of ancient trees revealed significant differences. The MS data of the plant samples are presented in Table 1.
Among the ancient tree samples, those with terpenoid compounds as the dominant volatile components included Lindera limprichtii and Cinnamomum jensenianum. The analysis of volatile components in the ancient tree samples revealed the predominance of terpene compounds in Magnolia sargentiana, D. involucrata, and Acer flabellatum. The analysis of the ancient tree samples revealed that those containing aldehyde compounds as the primary volatile organic compounds include Carpinus fangiana, Fagus lucida, Camellia oleifera, Prunus conradinae, and Enkianthus ruber. The volatile components of the ancient tree sample from Sorbus folgneri were primarily composed of patchoulane.
Monoterpenes and sesquiterpenes were significant components of the extracts of numerous medicinal plants and demonstrate robust biological activities and pharmacological effects (Kagawa et al. 2003; Nakai et al. 2003; Umeno et al. 2008; Tays et al. 2025). In the present study, the terpenoid substances identified in Lindera limprichtii, Cinnamomum jensenianum, M. sargentiana, D. involucrata, and A. flabellatum were β-cedrene, α-cedrene, α-pinene, (+)-β-funebrene, cis-Thujopsene, and (+)-cuparene. The bioactive substances cedrol and cis-Thujopsene endowed the wood of Sorbus folgneri with antibacterial and insect-repellent functions. M. sargentiana and L. limprichtii were notable for their abundance of terpenoid compounds, including α-cedrene and β-cedrene, which had antibacterial and anti-inflammatory properties. These compounds might also function as natural preservatives, contributing to the protection of wood from microbial infestation (Xie et al. 2022; Rashaduz et al. 2024). As demonstrated in the studies by Gao et al. (2005) and Dong et al. (2021), Carpinus fangiana, Enkianthus ruber, and Camellia oleifera contained high levels of aldehyde compounds, including valeraldehyde and hexanal. These compounds have been shown to release volatile organic compounds, which may contribute to the resistance of plants to herbivorous insects or pathogens (Gao et al. 2005; Dong et al. 2021). These results suggested that different tree species adapt to their ecological environments by synthesizing and releasing specific volatile compounds, thereby forming diverse ecological defense mechanisms (Peng et al. 2002).
Table 1. The Main Constituents of Ancient Trees
Analysis of Aroma Constituents in the Wood of Ancient Trees
The aroma properties of the volatile compounds in the wood of 11 ancient tree species were retrieved from the online database PubChem (Table 2). Specifically, a total of six aroma compounds were identified among the volatile organic compounds of both E. ruber and F. lucida. A total of five aroma compounds were identified among the volatile components of C. fangiana and P. conradinae. A total of three aroma compounds were identified among the volatile components of M. sargentiana, D. involucrata, A. flabellatum, and C. oleifera. The volatile components of both L. limprichtii and C. jensenianum were found to contain two aroma compounds. A single aroma compound was identified among the volatile components of S. folgneri.
In summary, the wood of the ancient tree species E. ruber, F. lucida, C. fangiana, and P. conradinae contained a greater number of aroma compounds. Compounds such as ethyl acetate and α-cedrene were detected in D. involucrata and C. jensenianum; these compounds exhibited fruity and woody aroma characteristics. These compounds might attract specific pollinating insects, thereby facilitating plant reproduction (Tak and Isman 2015; Chen et al. 2020). The high-content aldehyde compounds presented in E. ruber may function as defensive signaling molecules, thereby attracting natural enemies or repelling pests. Concurrently, the fruity aroma characteristics of these plants might also function as a form of attraction for pollinators (Zhu-Salzman et al. 2005; Meng et al. 2025). The broadening of research in the field has given rise to novel insights into the ecological functions of these volatile aromatic components, thereby offering a novel perspective for studying the interactions between plants and their environment.
Table 2. Odor Descriptions of Aroma Constituents in the Wood of Ancient Trees
Identification of Medicinal Active Constituents in Volatiles from Ancient Tree Wood
Based on the findings reported in the literature, a total of 25 volatile metabolites were identified across 11 ancient tree wood species (Ru et al. 2014). These results were derived from the Traditional Chinese Medicine Systems Pharmacology Database (TCMSP). Among these, 16 exhibited active medicinal components. Among the 16 metabolites examined, 15 were found to bind to 589 target proteins, corresponding to 466 diseases (Table 3). These diseases primarily included cancer, tumors, cardiovascular diseases, hypertension, inflammation, diabetes, Alzheimer’s disease, and neurological disorders.
Table 3. Identification of the Key Active Ingredients of Traditional Chinese Medicines in Poplar Wood
To identify key bioactive components, screening criteria of oral bioavailability (OB) ≥ 5% and drug-likeness (DL) ≥ 0.11 were applied. Among the 16 metabolites, 2,6,10,15-tetramethylheptadecane, cis-Thujopsene, cedrol, patchoulane, and β-cedrene meet both oral bioavailability (OB) ≥ 5% and drug-likeness (DL) ≥ 0.11. These results could serve as a reference for future research on the bioactive components (beneficial to human health) of the volatile substances in the wood of M. sargentiana, P. conradinae, L. limprichtii, C. jensenianum, S. folgneri, and F. lucida. The volatile substances of F. lucida wood contained 11 active medicinal components, among which 2,6,10,15-tetramethyl heptadecane—a key active component—had no corresponding target proteins or diseases. This finding suggested that 2,6,10,15-tetramethylheptadecane, a volatile compound present in the wood of F. lucida, might possess significant health-promoting properties and potential for drug development (Zhang et al. 2008). The presence of plant VOCs and negative air ions has been identified as a significant component of forest therapy. Li et al. (2011) demonstrated that ancient trees were a critical component of forests. This study contributed to the existing body of research on the key active components in the volatile substances of ancient tree wood that affected human health. The wood of M. sargentiana, P. conradinae, L. limprichtii, C. jensenianum, S. folgneri, and F. lucida was notable for its fresh aroma characteristics and high medicinal component content, which was advantageous for human health. These characteristics positioned the aforementioned tree species as promising candidates for the production of healthcare wood (Sales et al. 2020; Kong et al. 2022; Li et al. 2022; Jiang and Fan 2025).
CONCLUSIONS
- In this study, HS–SPME–GC–MS was utilized for the systematic analysis of volatile components in the wood of ancient trees. An analysis of ancient tree species revealed the presence of terpenoid compounds, predominantly monoterpenes (e.g., α-cedrene and β-cedrene) and sesquiterpenes (e.g., (+)-β-Funebrene). These secondary metabolites had the potential to increase the stress resilience of trees through mechanisms such as antibacterial activity and insect repellency. C. fangiana, E. ruber, and C. oleifera contain high levels of aldehyde compounds, including valeraldehyde and hexanal. The two compounds have been shown to release volatile organic compounds, which may contribute to the resistance of plants to herbivorous insects or pathogens. Additionally, they played a role in chemical signal transmission within ecosystems.
- A comprehensive analysis revealed substantial disparities in metabolic characteristics among diverse ancient tree species. D. involucrata, C. oleifera, and E. ruber were found to be particularly abundant in ester compounds, including ethyl acetate, which contributed to their fruity aroma profile and suggests potential for utilization in health care applications. C. fangiana and E. ruber contained a high proportion of aldehydes (with a peak area of up to 67%), which reflected their lipid oxidation properties and provided biomarkers for evaluating the preservation status of wood. M. sargentiana and L. limprichtii produced unique secondary metabolites, such as cedrol, which exhibited antibacterial activity and were valuable for the development of fragrance in the pharmaceutical and chemical engineering industries.
ACKNOWLEDGMENTS
The authors are grateful for the support of the Central Government’s Science and Technology Extension Programs for Forestry and Grassland (G22070663) and the Postgraduate Research Innovation Projects for the year 2025 (2025YCX36).
Conflict of Interest
The present study received no third-party financial support from enterprises, companies, or other entities, and no potential conflicts of interest are present.
Use of Generative AI
The authors declare that no generative AI was used in the preparation of this manuscript.
REFERENCES CITED
Chen, Y. C., Li, Z., Zhao, Y. X., Gao, M., Wang, J. Y., Liu, K. W., Wang, X., Wu, L. W., Jiao, Y. L., Xu, Z. L., et al. (2020). “The Litsea genome and the evolution of the laurel family,” Nat. Commun. 11(1), 1-14. https://doi.org/10.1038/s41467-020-15493-5
Dong, S. S., Liu, M., Liu, Y., Chen, F., Yang, T., Chen, L., Zhang, X. T., Guo, X., Fang, D. M., Li, L. Z, et al. (2021). “The genome of Magnolia biondii Pamp. provides insights into the evolution of Magnoliales and biosynthesis of terpenoids,” Hortic. Res. 8(1), 38. https://doi.org/10.1038/s41438-021-00471-9
Gao, Y., Jin, Y.-J., Li, H.-D. and Chen, H.-J. (2005). “Volatile organic compounds and their roles in bacteriostasis in five conifer species,” J. Integr. Plant Biol. 47(4), 499-507. https://doi.org/10.1111/j.1744-7909.2005.00081.x
Hu, S. J., and Zhang, R. M. (2022). “Roles of volatile organic compounds in plant adaptation to stress and physiological ecology,” J. Zhejiang Agric. & For. Univ. 39(6), 1378-1387. https://doi.org/10.11833/j.issn.2095-0756.20220180
Gilhen-Baker, M., Roviello, V., Beresford-Kroeger, D., Roviello, G. N. (2022). “Old growth forests and large old trees as critical organisms connecting ecosystems and human health, A review,” Environ. Chem. Lett. 20(2), 1529-1538. https://doi.org/10.1007/s10311-021-01372-y
Jiang, M., and Fan, Z. L. (2025). “Determination of bioactive compounds and antioxidant activity of four berries,” China Food Addit. 36(10), 69-76. https://doi.org/10.19804/j.issn1006-2513.2025.10.009
Kagawa, D., Jokura, H., Ochiai, R., Tokimitsu, I., and Tsubone, H.(2003). “The sedative effects and mechanism of action of cedrol inhalation with behavioral pharmacological evaluation,” Planta. Med. 69(7), 637-641. https://doi.org/10.1055/s-2003-41114
Kong, Q. B., Jiang, H., Guo, C. Y., Chen, T., Feng, S. L., Ding, C. B., Zhou, L. J. (2022). “Research progress on main chemical constituents and pharmacological activities of Camellia oleifera,” J. Chin. Cereals Oils Assoc. 37(4), 194-202. https://doi.org/10.3969/j.issn.1003-0174.2022.04.029
Li, H. Y., Lv, Q. Y., Liu, A., Wang, J. R., Sun, X. Q., Deng, J., Chen, Q. F., and Wu, Q. (2022). “Comparative metabolomics study of Tartary (Fagopyrum tataricum (L.) Gaertn) and common (Fagopyrum esculentum Moench) buckwheat seeds,” Food Chem. 371(1), 131125.1-131125.11. https://doi.org/10.1016/j.foodchem.2021.131125
Li, M. L., Yin, L.C., Zhang, Y., Su, X. K., Liu, G. H., Wang, X. F., Au, Y., and Wu, X. (2021). “Spatio-temporal dynamics of fractional vegetation coverage based on MODIS-EVI and its driving factors in Southwest China,” Acta Ecol. Sin. 41(3), 1138-1147. https://doi.org/10.5846/stxb201907101451
Li, Q., Otsuka, T., and Kobayashi, M. (2011). “Acute effects of walking in forest environments on cardiovascular and metabolic parameters,” Eur. J. Appl. Physiol. 111(11), 2845-2853. https://doi.org/10.1007/s00421-011-1918-z
Loreto, F., and Schnitzler, J. P. (2010). “Abiotic stresses and induced BVOCs,” Trends Plant Sci. 15(3), 154-166. https://doi.org/10.1016/j.tplants.2009.12.006
Meng, L. S., Zheng, T. T., Wang, S. Q., Chen T. J., Nie, Y., and Xu S. (2025). “Characterization and analysis of characteristic aroma compounds of fresh Dictyophora rubrovalvata in Guizhou based on gas chromatography-ion mobility spectrometry and gas chromatography-mass spectrometry,” Food and Fermentation Industries 51(16), 314-323. https://doi.org/10.13995/j.cnki.11-1802/ts.041636
Meritxell, P. H., Miquel, A. V., Sandra, V., Carolina, G., Clara, P., Vicent, A., Jose L. R., Vicente, N. L., Antonio, G., Alberto, U. (2021). “Plant exposure to herbivore-induced plant volatiles: a sustainable approach through eliciting plant defenses,” J. Pest Sci. 94(4), 1221-1235. https://doi.org/10.1007/s10340-021-01334-x
Nakai, Y., Kido, T., Hashimoto, K., Kase, Y., Sakakibara, I., Higuchi, M., and Sasaki, H. (2003). “Effect of the rhizomes of Atractylodes lancea and its constituents on the delay of gastric emptying,” J. Ethnopharmacol. 84(1), 51-55. https://doi.org/10.1016/s0378-8741(02)00260-x
Ninkuu, V., Zhang, L., Yan, J. P., Fu, Z. C., Yang, T. F., Zeng, H. M. (2021). “Biochemistry of terpenes and recent advances in plant protection,” Int. J. Mol. Sci. 22(11), 5710. https://doi.org/10.3390/ijms22115710
Peng, S. L., Nan P., and Zhong, Y. (2002). “Terpenoids in higher plants and their roles in ecosystems,” Chin. J. Ecol. 21(3), 33-38. https://doi.org/10.13292 / j.1000-4890.2002.0050
Pichersky, E., and Gershenzon, J. (2002). “The formation and function of plant volatiles: perfumes for pollinator attraction and defense,” Curr. Opin. Plant Biol. 5(3), 237-243. https://doi.org/10.1016/s1369-5266(02)00251-0
Rashaduz, Z., Akanksha J., Almuth H., Jonathan G., and Dineshkumar K. (2024). “A rationale for chemical defense mixtures in spruce oleoresin: Most monoterpenes are highly toxic to either bark beetles or to their symbiotic fungi, but not both,” New Phytol. 1(1), 1-21. https://doi.org/10.21203/rs.3.rs-4919445/v1
Ru, J. L., Li, P., Wang, J. N., Zhou, W., Li, B. H., Huang, C., Li, P. D., Guo, Z. H., Tao, W. Y. Yang, Y. F., et al. (2014). “TCMSP: a database of systems pharmacology for drug discovery from herbal medicines,” J. Cheminform. 6(1), 1-6. https://doi.org/10.1186/1758-2946-6-13
Sales, A., Felipe, O. L., and Bicas, J. L. (2020). “Production, properties, and applications of α-terpineol,” Food Bioproc. Tech. 13(8), 1261-1279. https://doi.org/10.1007/s11947-020-02461-6
Schreiner, L., Bauer, P., and Buettnerm, A. (2018). “Resolving the smell of wood identification of odour-active compounds in Scots pine (Pinus sylvestris L.),” Sci. Rep. 8(1), 1-9. https://doi.org/10.1038/s41598-018-26626-8
Shen, L., Chu Y. G., Zhang W. X., Zhang J., Zhang T. Q., Li Z. H., Ding C. J., and Su X. H. (2024). “Comparative analysis of volatile compounds in the wood of Populus koreana and Populus ussuriensis,” Scientia Silvae Sinicae 60(01), 103-110. https://doi.org/10.11707/j.1001-7488.LYKX20220443
Sotaro, H., and Akiko, S. (2024). “Theoretical analyses for the evolution of biogenic volatile organic compounds (BVOC) emission strategy,” BMC Ecol. Evol. 14(7), 1-13. https://doi.org/10.1002/ece3.11548
Tak, J. H., and Isman, M. B. (2015). “Enhanced cuticular penetration as the mechanism for synergy of insecticidal constituents of rosemary essential oil in Trichoplusia ni,” Sci. Rep. 5(1), 12690. https://doi.org/10.1038/srep12690
Tays, G., Arthur A., Larisse P., Julio O., Mathania F., Javanyr J., Robson V. and Isac M. (2025) “Monoterpenes in vascular function: A review of bioactivity and mechanisms of action,” Int. J. Mol. Sci. 26(18), 9243. https://doi.org/10.3390/ijms26189243
Turlings, T. C. J., and Erb, M. (2018). “Tritrophic interactions mediated by herbivore induced plant volatiles: Mechanisms, ecological relevance, and application potential,” Annu. Rev. Entomol. 7(63), 433-452. https://doi.org/10.1146/annurev-ento-020117-043507
Umeno, K., Hori, E., Tsubota, M., Shojaku, H., Miwa, T., Nagashima, Y., Yada, Y., Suzuki, T., Ono, T., and Nishijo, H. (2008). “Effects of direct cedrol inhalation into the lower airway on autonomic nervous activity in totally laryngectomized subjects,” Br. J. Clin. Pharmacol. 65(2), 188-196. https://doi.org/10.1111/j.1365-2125.2007.02992.x
Xie, D. Z., Ma, H. Y., Lai, W. T., Zhang, H. L., Yuan, F. J., and Qian, Y. L. (2022). “Released characteristics of volatile organic compounds in Cunninghamia lanceolata,” Subtrop. Plant Sci. 51(03), 178-183. https://doi.org/10.3969/j.issn.1009-7791.2022.03.003
Yan, Y. F., Yang, C. J., Shang, X. F., Zhao, Z. M., Liu, Y. Q., Zhou, R., Liu, H., Wu, T. L., Zhao, W. B., Wang, Y. L., et al. (2020). “Bioassay- guided isolation of two antifungal compounds from Magnolia officinalis, and the mechanism of action of honokiol,” Pestic. Biochem. Physiol. 170(170), 12-13. https://doi.org/10.1016/j.pestbp.2020.104705
Yu, J. F., He, Q., Zhang, X. T., Yao, L. H., Wang, X. M. (2020). “Sampling and test method of volatile organic compounds (VOCs) from wood drying,” Appl. Chem. Ind. 49(08), 1941-1945. https://doi.org/10.16581/j.cnki.issn.1671-3206.2020.08.002
Zeng, B., Shen, J., Wang, Q. F., Dong, H. J. (2021). “Analysis of odorants in Cinnamomum burmannii wood with different moisture contents,” Sci. Silv. Sin. 57(4), 133-141. https://doi.org/10. 11707/j.1001-7488.20210414
Zhang, Y. Y., Jia, X. N., Cao, Y. X., and Wang, J. X. (2008). “Studies on the chemical constituents of Abelmoschus manihot,” Northwest Pharm. J. 23(2), 80-82. https://doi.org/10.3969/j.issn.1004-2407.2008.02.010
Zhu-Salzman, K., Bi, J. L., and Liu, T. X. (2005). “Molecular strategies of plant defense and insect counter-defense,” Insect Sci. 12(1), 3-15. https://doi.org/10.1111/j.1672-9609.2005.00002.x
Article submitted: March 5, 2026; Peer review completed: March 29, 2026; Revisions accepted: April 5, 2026; Published: April 15, 2026.
DOI: 10.15376/biores.21.2.4780-4791