Unveiling stand-level airborne biogenic volatile organic compounds (BVOCs): spatiotemporal dynamics and environmental drivers in a Larix gmelinii var. principis-rupprechtii plantation, China

Abstract

Biogenic volatile organic compounds (BVOCs) significantly contribute to forest therapy benefits, yet their spatiotemporal emission patterns and environmental drivers in forest air remain inadequately understood. This study aimed to characterize the spatiotemporal dynamics of BVOCs and their environmental drivers in Larix gmelinii var. principis-rupprechtii plantations. Using open-path sampling with GC-MS, we investigated diurnal BVOCs emission profiles from 6:00 to 20:00 in mid-July, mid-August, and mid-September. Vertical and horizontal distribution patterns were characterized at three canopy strata (upper, middle, lower) and microsites (interior, edge, gap). Simultaneous measurements of illuminance, temperature, humidity, wind speed, particulate matter < 10 μm (PM₁₀), and negative air ion (NAI) were conducted during diurnal sampling to identify environmental correlates. The study found that total BVOCs emission peaked in the upper and middle canopies—the former enriched in decanal and o-cymene, and the latter in nonanal and monoterpenes—while emission was lowest from the chlorotic leaves in the lower. Horizontally, BVOCs concentrations were highest in the forest interior and decreased outward, being lowest in forest gap. Diurnally, BVOCs decrease during midday photosynthetic depression; at dusk cis-3-hexen-1-ol and 2-methylpentanal emerged, and β-pinene were substantially released; nonanal and decanal show U-shaped patterns. Generalized Additive Models (GAM) identified that nonanal variation was primarily driven by illuminance (F=4.46, p = 0.07), followed by PM₁₀ (F = 3.66, p = 0.06), temperature (F = 2.92, p < 0.05), and humidity (F = 2.46, p < 0.05). Partial Least Squares Path Modeling (PLS-PM) revealed that meteorological factors influenced nonanal through both direct (p < 0.05) and indirect (p < 0.05) pathway. The direct effect was substantial (β = 0.37), while the indirect effect—mediated by PM₁₀ and NAI—was weaker (β = 0.14). These findings established spatiotemporal patterns and key environmental drivers of BVOCs in L. gmelinii var. principis-rupprechtii plantations. They suggest elevating walkways to near mid-canopy height and locating core activity areas inside the forest to optimize exposure to beneficial volatiles. Furthermore, scheduling activities around dawn and dusk during the growing season is recommended. From a management perspective, creating diffuse light conditions and elevating air humidity can foster a more conducive microclimate for therapy.

Keywords

Biogenic volatile organic compounds (BVOCs) / Nonanal / Canopy strata / Diurnal variation / Microclimate

References

[1] Aaltonen H, Pumpanen J, Pihlatie M, Hakola H, Hellén H, Kulmala L, Vesala T, Bäck J. Boreal pine forest floor biogenic volatile organic compound emissions peak in early summer and autumn. Agric for Meteor, 2011, 151(6): 682-691.

[2] Akhtar N, Alharthi MF. A comparative study of the performance of new ridge estimators for multicollinearity: insights from simulation and real data application. AIP Adv, 2024, 14(11): 115311.

[3] Alves EG, Harley P, de C Gonçalves JF, da Silva Moura CE, Jardine K. Effects of light and temperature on isoprene emission at different leaf developmental stages of Eschweilera coriacea in central Amazon. Acta Amaz, 2014, 44(1): 9-18.

[4] Anselmo-Moreira F, da Silva Pedrosa G, da Silva IL, do Nascimento A, dos Santos TC, Catharino ELM, Gomes EPC, Borbon A, Fornaro A, de Souza SR. Biogenic volatile organic compound (BVOC) emission profiles from native Atlantic Forest trees: seasonal variation and atmospheric implications in southeastern Brazil. Urban for Urban Green, 2025, 104: 128645.

[5] Api AM, Belsito D, Botelho D, Bruze M, Burton GAJr, Cancellieri MA, Chon H, Dagli ML, Date M, Dekant W, Deodhar C, Fryer AD, Jones L, Joshi K, Kumar M, Lapczynski A, Lavelle M, Lee I, Liebler DC, Moustakas H, Na M, Penning TM, Ritacco G, Romine J, Sadekar N, Schultz TW, Selechnik D, Siddiqi F, Sipes IG, Sullivan G, Thakkar Y, Tokura Y. RIFM fragrance ingredient safety assessment, 2-ethylhexanal, CAS Registry Number 123-05-7. Food Chem Toxicol, 2022, 163: 113058.

[6] Api AM, Belsito D, Botelho D, Bruze M, Burton GA, Cancellieri MA, Chon H, Dagli ML, Date M, Dekant W, Deodhar C, Fryer AD, Jones L, Joshi K, Kumar M, Lapczynski A, Lavelle M, Lee I, Liebler DC, Moustakas H, Na M, Penning TM, Ritacco G, Romine J, Sadekar N, Schultz TW, Selechnik D, Siddiqi F, Sipes IG, Sullivan G, Thakkar Y, Tokura Y. RIFM fragrance ingredient safety assessment, 2-methylpentanal, CAS Registry Number 123–15-9. Food Chem Toxicol, 2022, 163: 113007.

[7] Balahbib A, El Omari N, EL Hachlafi N, Lakhdar F, El Menyiy N, Salhi N, Mrabti HN, Bakrim S, Zengin G, Bouyahya A. Health beneficial and pharmacological properties of p-cymene. Food Chem Toxicol, 2021, 153: 112259.

[8] Bamberger I, Hörtnagl L, Schnitzhofer R, Graus M, Ruuskanen TM, Müller M, Dunkl J, Wohlfahrt G, Hansel A. BVOC fluxes above mountain grassland. Biogeosciences, 2010, 7(5): 1413-1424.

[9] Bradow JM, Connick WJ. Volatile seed germination inhibitors from plant residues. J Chem Ecol, 1990, 16(3): 645-666.

[10] Bunce JA. Effects of humidity on short-term responses of stomatal conductance to an increase in carbon dioxide concentration. Plant Cell Environ, 1998, 21(1): 115-120.

[11] Cai MF, An CJ, Guy C. A scientometric analysis and review of biogenic volatile organic compound emissions: research hotspots, new frontiers, and environmental implications. Renew Sustain Energy Rev, 2021, 149: 111317.

[12] Cai LL, Shi GY, Zhang JS, Du LT, Ni XL, Hu Y, Pang DB, Meng JH. Empirical analysis of the influence mechanism of vegetation and environment on negative air ion in warm temperate forest ecosystems. Environ Pollut, 2024, 363: 125273.

[13] Cao JP, Liu C, Zhang YP. Influence of airborne particles on convective mass transfer of SVOCs on flat surfaces: novel insight and estimation formula. Int J Heat Mass Transf, 2017, 115: 127-136.

[14] Carriero G, Brunetti C, Fares S, Hayes F, Hoshika Y, Mills G, Tattini M, Paoletti E. BVOC responses to realistic nitrogen fertilization and ozone exposure in silver birch. Environ Pollut, 2016, 213: 988-995.

[15] Chen JG, Tang J, Yu XX. Environmental and physiological controls on diurnal and seasonal patterns of biogenic volatile organic compound emissions from five dominant woody species under field conditions. Environ Pollut, 2020, 259: 113955.

[16] Chhikara N, Kour R, Jaglan S, Gupta P, Gat Y, Panghal A. Citrus medica: nutritional, phytochemical composition and health benefits–a review. Food Funct, 2018, 9(4): 1978-1992.

[17] Copeland N, Cape JN, Heal MR. Volatile organic compound emissions from Miscanthus and short rotation coppice willow bioenergy crops. Atmos Environ, 2012, 60: 327-335.

[18] Darbah JNT, Sharkey TD, Calfapietra C, Karnosky DF. Differential response of aspen and birch trees to heat stress under elevated carbon dioxide. Environ Pollut, 2010, 158(4): 1008-1014.

[19] Duan B, Zhang YH, Feng Z, Liu ZG, Tao NG. Octanal enhances disease resistance in postharvest Citrus fruit by the biosynthesis and metabolism of aromatic amino acids. Pestic Biochem Physiol, 2024, 200: 105835.

[20] Duan YF, Lv BY, Zhang CL, Shi LS, Li JT, Liu YJ, Chen QB. Antibacterial activity of the biogenic volatile organic compounds from three species of bamboo. Front Plant Sci, 2024, 15: 1474401.

[21] Duda-Madej A, Viscardi S, Grabarczyk M, Topola E, Kozłowska J, Mączka W, Wińska K. Is camphor the future in supporting therapy for skin infections?. Pharmaceuticals Basel, 2024, 17(6): 715.

[22] El Shiekh RA, Atwa AM, Elgindy AM, Mustafa AM, Senna MM, Alkabbani MA, Ibrahim KM. Therapeutic applications of Eucalyptus essential oils. Inflammopharmacology, 2025, 33(1): 163-182.

[23] El-Saadony MT, Saad AM, Korma SA, Salem HM, Abd El-Mageed TA, Alkafaas SS, Elsalahaty MI, Elkafas SS, Mosa WFA, Ahmed AE, Mathew BT, Albastaki NA, Alkuwaiti AA, El-Tarabily MK, AbuQamar SF, El-Tarabily KA, Ibrahim SA. Garlic bioactive substances and their therapeutic applications for improving human health: a comprehensive review. Front Immunol, 2024, 15: 1277074.

[24] Erickson E, Wakao S, Niyogi KK. Light stress and photoprotection in Chlamydomonas reinhardtii. Plant J, 2015, 82(3): 449-465.

[25] Farmaki T, Sanmartín M, Jiménez P, Paneque M, Sanz C, Vancanneyt G, León J, Sánchez-Serrano JJ. Differential distribution of the lipoxygenase pathway enzymes within potato chloroplasts. J Exp Bot, 2007, 58(3): 555-568.

[26] Filella I, Peñuelas J, Llusià J. Dynamics of the enhanced emissions of monoterpenes and methyl salicylate, and decreased uptake of formaldehyde, by Quercus ilex leaves after application of jasmonic acid. New Phytol, 2006, 169(1): 135-144.

[27] Gao Y, Jin YJ, Li HD, Chen HJ. Volatile organic compounds and their roles in bacteriostasis in five conifer species. J Integr Plant Biol, 2005, 47(4): 499-507.

[28] Greenberg JP, Asensio D, Turnipseed A, Guenther AB, Karl T, Gochis D. Contribution of leaf and needle litter to whole ecosystem BVOC fluxes. Atmos Environ, 2012, 59: 302-311.

[29] Guenther AB, Zimmerman PR, Harley PC, Monson RK, Fall R. Isoprene and monoterpene emission rate variability: model evaluations and sensitivity analyses. J Geophys Res Atmos, 1993, 98(D7): 12609-12617.

[30] Guidolotti G, Calfapietra C, Loreto F. The relationship between isoprene emission, CO₂ assimilation and water use efficiency across a range of poplar genotypes. Physiol Plant, 2011, 142(3): 297-304.

[31] Hagiwara T, Ishihara MI, Takabayashi J, Hiura T, Shiojiri K. Effective distance of volatile cues for plant–plant communication in beech. Ecol Evol, 2021, 11(18): 12445-12452.

[32] Hammerbacher A, Coutinho TA, Gershenzon J. Roles of plant volatiles in defence against microbial pathogens and microbial exploitation of volatiles. Plant Cell Environ, 2019, 42(10): 2827-2843.

[33] Hidalgo-Fuentes B, de JoséJesús- E, de J Cabrera-Hidalgo A, Sandoval-Castilla O, Espinosa-Solares T, González-Reza RM, Zambrano-Zaragoza ML, Liceaga AM, Aguilar-Toalá JE. Plant-based fermented beverages: nutritional composition, sensory properties, and health benefits. Foods, 2024, 13(6): 844.

[34] Hirose S, Satake A. Theoretical analyses for the evolution of biogenic volatile organic compounds (BVOC) emission strategy. Ecol Evol, 2024, 14(7): e11548.

[35] Holopainen JK, Gershenzon J. Multiple stress factors and the emission of plant VOCs. Trends Plant Sci, 2010, 15(3): 176-184.

[36] Hu ZH, Shen YB, Su XH. Saturated aldehydes C6–C10 emitted from ashleaf maple (Acer negundo L.) leaves at different levels of light intensity, O₂, and CO₂. J Plant Biol, 2009, 52(4): 289-297.

[37] Hu ZH, Leng PS, Shen YB, Wang WH. Emissions of saturated C6–C10 aldehydes from poplar (Populus simonii × P. pyramidalis ‘Opera 8277’) cuttings at different levels of light intensity. J for Res, 2011, 22(2): 233-238.

[38] Huang L, McGaughey G, McDonald-Buller E, Kimura Y, Allen DT. Quantifying regional, seasonal and interannual contributions of environmental factors on isoprene and monoterpene emissions estimates over eastern Texas. Atmos Environ, 2015, 106: 120-128.

[39] Huang JB, Hartmann H, Hellén H, Wisthaler A, Perreca E, Weinhold A, Rücker A, van Dam NM, Gershenzon J, Trumbore S, Behrendt T. New perspectives on CO₂, temperature, and light effects on BVOC emissions using online measurements by PTR-MS and cavity ring-down spectroscopy. Environ Sci Technol, 2018, 52(23): 13811-13823.

[40] Huang RX, Zhang TN, Ge XG, Cao YH, Li ZC, Zhou BZ. Emission trade-off between isoprene and other BVOC components in Pinus massoniana saplings may be regulated by content of chlorophylls, starch and NSCs under drought stress. Int J Mol Sci, 2023, 24(10): 8946.

[41] Ibrahim MH, Jaafar HZE. Reduced photoinhibition under low irradiance enhanced Kacip Fatimah (Labisia pumila Benth) secondary metabolites, phenyl alanine lyase and antioxidant activity. Int J Mol Sci, 2012, 13(5): 5290-5306.

[42] Ieda T, Kitamori Y, Mochida M, Hirata R, Hirano T, Inukai K, Fujinuma Y, Kawamura K. Diurnal variations and vertical gradients of biogenic volatile and semi-volatile organic compounds at the Tomakomai larch forest station in Japan. Tellus B Chem Phys Meteorol, 2006, 58(3): 177-186.

[43] Jardine K, Harley P, Karl T, Guenther A, Lerdau M, Mak JE. Plant physiological and environmental controls over the exchange of acetaldehyde between forest canopies and the atmosphere. Biogeosciences, 2008, 5(6): 1559-1572.

[44] Jia ZC, Zhou RY, Jiao JJ, Pan CY, Chen ZH, Huang YC, Zhou YF, Zhou GM. Trends and influencing factors of summer air quality changes in four forest types. Forests, 2025, 16(5): 833.

[45] Jing XX, Lun XX, Fan C, Ma WF. Emission patterns of biogenic volatile organic compounds from dominant forest species in Beijing, China. J Environ Sci, 2020, 95: 73-81.

[46] Kang W, Choi D, Park T. Decanal protects against UVB-induced photoaging in human dermal fibroblasts via the cAMP pathway. Nutrients, 2020, 12(5): 1214.

[47] Kang J, Choi J, Lee K. Development of an evaluation index for forest therapy environments. Int J Environ Res Public Health, 2024, 21(2): 136.

[48] Karlsson T, Klemedtsson L, Rinnan R, Holst T. Leaf-scale study of biogenic volatile organic compound emissions from willow (Salix spp.) short rotation coppices covering two growing seasons. Atmosphere, 2021, 12(11): 1427.

[49] Kegge W, Pierik R. Biogenic volatile organic compounds and plant competition. Trends Plant Sci, 2010, 15(3): 126-132.

[50] Kesselmeier J, Staudt M. Biogenic volatile organic compounds (VOC): an overview on emission, physiology and ecology. J Atmos Chem, 1999, 33(1): 23-88.

[51] Khruengsai S, Phalangrit KA, Sripahco T, Kaewfoo M, Sittikarn N, Poshyachinda S, Pongpiachan S, Pripdeevech P. Diurnal variations of biogenic volatile organic compounds and their role in secondary pollutant formation in the Huai Hong Khrai subtropical forest, Thailand. Environ Pollut, 2025, 372: 126044.

[52] Kim JC, Dinh TV, Oh HK, Son YS, Ahn JW, Song KY, Choi IY, Park CR, Szulejko JE, Kim KH. The potential benefits of therapeutic treatment using gaseous terpenes at ambient low levels. Appl Sci, 2019, 9(21): 4507.

[53] Kim M, Sowndhararajan K, Choi HJ, Park SJ, Kim S. Olfactory stimulation effect of aldehydes, nonanal, and decanal on the human electroencephalographic activity, according to nostril variation. Biomedicines, 2019, 7(3): 57.

[54] Kim YJ, Shin YM, Lee H, Moon SY, Kim TJ, Park SU, Seo WD, Kim JK. Comparative analysis of primary and secondary metabolites in wheat seedlings (Triticum aestivum L.) cultivated under varying photosynthetic photon flux densities and growth periods. J Sci Food Agric, 2025, 105(13): 7260-7269.

[55] Kirešová S, Guzan M. Determining the correlation between particulate matter PM₁₀ and meteorological factors. Eng, 2022, 3(3): 343-363.

[56] J Knutsson 2024 Volatile organic compound emissions and secondary organic aerosols formation during clearcutting of a boreal forest at Norunda, Sweden, Gothenburg University

[57] La Mer VK. The calculation of thermodynamic quantities from hysteresis data. J Colloid Interface Sci, 1967, 23(2): 297-301.

[58] Lappalainen HK, Sevanto S, Bäck J, Ruuskanen TM, Kolari P, Taipale R, Rinne J, Kulmala M, Hari P. Day-time concentrations of biogenic volatile organic compounds in a boreal forest canopy and their relation to environmental and biological factors. Atmos Chem Phys, 2009, 9(15): 5447-5459.

[59] Lee HJ, Hyun EA, Yoon WJ, Kim BH, Rhee MH, Kang HK, Cho JY, Yoo ES. In vitro anti-inflammatory and anti-oxidative effects of Cinnamomum camphora extracts. J Ethnopharmacol, 2006, 103(2): 208-216.

[60] Leonowicz G, Trzebuniak KF, Zimak-Piekarczyk P, Ślesak I, Mysliwa-Kurdziel B. The activity of superoxide dismutases (SOD_s) at the early stages of wheat deetiolation. PLoS ONE, 2018, 13(3): e0194678.

[61] Li HL, Yang PF, Liu LY, Gong BB, Zhang ZF, Ma WL, MacDonald RW, Nikolaev AN, Li YF. Steady-state based model of airborne particle/gas and settled dust/gas partitioning for semivolatile organic compounds in the indoor environment. Environ Sci Technol, 2022, 56(12): 8373-8383.

[62] Li LJ, Wang XZ, Jia ZK. Effects of environmental factors on the temporal and spatial variations in branch and leaf CO₂ efflux of Larix gmelinii var. principis-rupprechtii Mayr. J for Res, 2023, 34(4): 1007-1019.

[63] Lin Y, Lun XX, Tang W, Zhang ZZ, Jing XX, Fan C, Wang Q. Characteristics and chemical reactivity of biogenic volatile organic compounds from dominant forest species in the Jing-Jin-Ji area, China. Forest Ecosystems, 2021, 8: 52.

[64] Liu Y, Li L, An JY, Huang L, Yan RS, Huang C, Wang HL, Wang Q, Wang M, Zhang W. Estimation of biogenic VOC emissions and its impact on ozone formation over the Yangtze River Delta region, China. Atmos Environ, 2018, 186: 113-128.

[65] Liu LP, Seyler BC, Liu HF, Zhou L, Chen DY, Liu S, Yan C, Yang FM, Song DL, Tan QW, Jia FJ, Feng C, Wang QW, Li YC. Biogenic volatile organic compound emission patterns and secondary pollutant formation potentials of dominant greening trees in Chengdu, Southwest China. J Environ Sci, 2022, 114: 179-193.

[66] Llusia J, Peñuelas J, Guenther A, Rapparini F. Seasonal variations in terpene emission factors of dominant species in four ecosystems in NE Spain. Atmos Environ, 2013, 70: 149-158.

[67] Loreto F, Pinelli P, Manes F, Kollist H. Impact of ozone on monoterpene emissions and evidence for an isoprene-like antioxidant action of monoterpenes emitted by Quercus ilex leaves. Tree Physiol, 2004, 24(4): 361-367.

[68] Loreto F, Centritto M, Barta C, Calfapietra C, Fares S, Monson RK. The relationship between isoprene emission rate and dark respiration rate in white poplar (Populus alba L.) leaves. Plant Cell Environ, 2007, 30(5): 662-669.

[69] Lu DL, Zhu JJ, Zhang GQ, Fang S, Sun YR, Zhu CY, Zhang JX, Wang GG. A forest gap is not forever: towards an objective standard to determine when a gap is considered closed in temperate forests. Agric for Meteorol, 2023, 340: 109598.

[70] Marsol-Vall A, Kortesniemi M, Karhu ST, Kallio H, Yang BR. Profiles of volatile compounds in blackcurrant (Ribes nigrum) cultivars with a special focus on the influence of growth latitude and weather conditions. J Agric Food Chem, 2018, 66(28): 7485-7495.

[71] Mehra A, Krechmer JE, Lambe A, Sarkar C, Williams L, Khalaj F, Guenther A, Jayne J, Coe H, Worsnop D, Faiola C, Canagaratna M. Oligomer and highly oxygenated organic molecule formation from oxidation of oxygenated monoterpenes emitted by California sage plants. Atmos Chem Phys, 2020, 20(18): 10953-10965.

[72] Mohammed HA. Phytochemical analysis, antioxidant potential, and cytotoxicity evaluation of traditionally used Artemisia absinthium L. (wormwood) growing in the central region of Saudi Arabia. Plants, 2022, 11(8): 1028.

[73] Moliterno AAC, Jakuš R, Modlinger R, Unelius CR, Schlyter F, Jirošová A. Field effects of oxygenated monoterpenes and estragole combined with pheromone on attraction of Ips typographus and its natural enemies. Front for Glob Change, 2023, 6: 1292581.

[74] Morrison EC, Drewer J, Heal MR. A comparison of isoprene and monoterpene emission rates from the perennial bioenergy crops short-rotation coppice willow and Miscanthus and the annual arable crops wheat and oilseed rape. GCB Bioenergy, 2016, 8(1): 211-225.

[75] Mozaffar A, Schoon N, Bachy A, Digrado A, Heinesch B, Aubinet M, Fauconnier ML, Delaplace P, du Jardin P, Amelynck C. Biogenic volatile organic compound emissions from senescent maize leaves and a comparison with other leaf developmental stages. Atmos Environ, 2018, 176: 71-81.

[76] Nie CN, Gao Y, Du X, Bian JL, Li H, Zhang X, Wang CM, Li SY. Characterization of the effect of cis-3-hexen-1-ol on green tea aroma. Sci Rep, 2020, 10: 15506.

[77] Oku H, Inafuku M, Takamine T, Nagamine M, Saitoh S, Fukuta M. Temperature threshold of isoprene emission from tropical trees, Ficus virgata and Ficus septica. Chemosphere, 2014, 95: 268-273.

[78] Paduch R, Kandefer-Szerszeń M, Trytek M, Fiedurek J. Terpenes: substances useful in human healthcare. Arch Immunol Ther Exp, 2007, 55(5): 315-327.

[79] Petersen R, Holst T, Mölder M, Kljun N, Rinne J. Vertical distribution of sources and sinks of volatile organic compounds within a boreal forest canopy. Atmos Chem Phys, 2023, 23(13): 7839-7858.

[80] Pinheiro-Neto FR, Lopes EM, Acha BT, da Silva Gomes L, Dias WA, dos Reis Filho AC, de Sousa Leal B, do Nascimento Rodrigues DC, do Nascimento Silva J, Dittz D, Ferreira PMP, de Castro Almeida FR. α-Phellandrene exhibits antinociceptive and tumor-reducing effects in a mouse model of oncologic pain. Toxicol Appl Pharmacol, 2021, 418: 115497.

[81] Potosnak MJ. Including the interactive effect of elevated CO₂ concentration and leaf temperature in global models of isoprene emission. Plant Cell Environ, 2014, 37(8): 1723-1726.

[82] Ravindra K, Rattan P, Mor S, Aggarwal AN. Generalized additive models: building evidence of air pollution, climate change and human health. Environ Int, 2019, 132: 104987.

[83] Razo-Belmán R, Castañeda-Rodríguez R, Heil M, Ozuna C. Nonanal, a volatile organic compound, improves the techno-functional, nutritional, and nutraceutical properties of the common bean (Phaseolus vulgaris L.). Food Chem Adv, 2024, 4: 100707.

[84] Ren L, Huo JX, Xiang X, Pan YP, Li YQ, Wang YY, Meng DH, Yu C, Chen Y, Xu ZC, Huang YM. Environmental conditions are the dominant factor influencing stability of terrestrial ecosystems on the Tibetan Plateau. Commun Earth Environ, 2023, 4: 196.

[85] Rosenstiel TN, Potosnak MJ, Griffin KL, Fall R, Monson RK. Increased CO₂ uncouples growth from isoprene emission in an agriforest ecosystem. Nature, 2003, 421(6920): 256-259.

[86] Rosenstiel TN, Ebbets AL, Khatri WC, Fall R, Monson RK. Induction of poplar leaf nitrate reductase: a test of extrachloroplastic control of isoprene emission rate. Plant Biol, 2004, 6(1): 12-21.

[87] Sacchi R, Caporaso N, Squadrilli GA, Paduano A, Ambrosino ML, Cavella S, Genovese A. Sensory profile, biophenolic and volatile compounds of an artisanal ice cream (‘gelato’) functionalised using extra virgin olive oil. Int J Gastron Food Sci, 2019, 18: 100173.

[88] Salehi B, Upadhyay S, Erdogan Orhan I, Kumar Jugran A, Jayaweera LDS, Dias DA, Sharopov F, Taheri Y, Martins N, Baghalpour N, Cho CW, Sharifi-Rad J. Therapeutic potential of α- and β-pinene: a miracle gift of nature. Biomolecules, 2019, 9(11): 738.

[89] Sanadze GA. Biogenic isoprene emission as expression of dissipativity, a fundamental cell property. Russ J Plant Physiol, 2017, 64(2): 133-140.

[90] Satake A, Hagiwara T, Nagano AJ, Yamaguchi N, Sekimoto K, Shiojiri K, Sudo K. Plant molecular phenology and climate feedbacks mediated by BVOCs. Annu Rev Plant Biol, 2024, 75: 605-627.

[91] Sayyari M, Babalar M, Kalantari S, Martínez-Romero D, Guillén F, Serrano M, Valero D. Vapour treatments with methyl salicylate or methyl jasmonate alleviated chilling injury and enhanced antioxidant potential during postharvest storage of pomegranates. Food Chem, 2011, 124(3): 964-970.

[92] Schulze BC, Wallace HW, Flynn JH, Lefer BL, Erickson MH, Jobson BT, Dusanter S, Griffith SM, Hansen RF, Stevens PS, VanReken T, Griffin RJ. Differences in BVOC oxidation and SOA formation above and below the forest canopy. Atmos Chem Phys, 2017, 17(3): 1805-1828.

[93] GH Seol, KY Kim Eucalyptol and its role in chronic diseases. In: Drug discovery from mother nature. Springer International Publishing, pp 389–398. https://doi.org/10.1007/978-3-319-41342-6_18 2016

[94] Serradilla MJ, Martín A, Hernandez A, López-Corrales M, Lozano M, de Guía Córdoba M. Effect of the commercial ripening stage and postharvest storage on microbial and aroma changes of ‘ambrunés’ sweet cherries. J Agric Food Chem, 2010, 58(16): 9157-9163.

[95] Shahrajabian MH, Sun WL. The important nutritional and wonderful health benefits of cashew (Anacardium occidentale L.). Nat Prod J, 2023, 13(4): e270422204127.

[96] Shi GY, Zhou Y, Sang YQ, Zhang JS, Meng P, Cai LL, Pei SY, Wang Y. Influence of environmental factors on negative air ion using random forest algorithm. Chin J Agrometeorol, 2021, 42(5): 390-401. (in Chinese).

[97] Shimada Y, Kumazoe M, Otsuka Y, Tetsuzen R, Fujimura Y, Tachibana H. Neuroprotective effect of isovaleraldehyde accompanied with upregulation of BDNF and CREB phosphorylation via the PKA pathway. J Nat Med, 2024, 78(1): 208-215.

[98] Stamatiou R, Anagnostopoulou M, Ioannidou-Kabouri K, Rapti C, Lazou A. Camphene as a protective agent in myocardial ischemia/reperfusion injury. Antioxidants, 2024, 13(4): 405.

[99] Thangaleela S, Sivamaruthi BS, Kesika P, Tiyajamorn T, Bharathi M, Chaiyasut C. A narrative review on the bioactivity and health benefits of alpha-phellandrene. Sci Pharm, 2022, 90(4): 57.

[100] Tian GJ, Qiao Z, Xu XL. Characteristics of particulate matter (PM₁₀) and its relationship with meteorological factors during 2001–2012 in Beijing. Environ Pollut, 2014, 192: 266-274.

[101] Touhami D, Mofikoya AO, Girling RD, Langford B, Misztal PK, Pfrang C. Atmospheric degradation of ecologically important biogenic volatiles: investigating the ozonolysis of (E)-β-ocimene, isomers of α and β-farnesene, α-terpinene and 6-methyl-5-hepten-2-one, and their gas-phase products. J Chem Ecol, 2024, 50(3): 129-142.

[102] Tudor C, Muntean PAL, Socaci SA, Dulf FV, Bunea A, Rugină D, Pintea AM. Stability of sea buckthorn, blackcurrant, linseed and olive oil-in-water emulsions against thermally induced oxidation. Bull Univ Agric Sci Vet Med Cluj-Napoca Food Sci Technol, 2025, 82(2): 37-50.

[103] Ullah A, Gao DM, Wu FZ. Common mycorrhizal network: the predominant socialist and capitalist responses of possible plant–plant and plant–microbe interactions for sustainable agriculture. Front Microbiol, 2024, 15: 1183024.

[104] Wang SS, Yang Y, Trishchenko AP, Barr AG, Black TA, McCaughey H. Modeling the response of canopy stomatal conductance to humidity. J Hydrometeorol, 2009, 10(2): 521-532.

[105] Wang H, Wang B, Niu X, Song QF, Li MW, Luo YY, Liang LD, Du PF, Peng W. Study on the change of negative air ion concentration and its influencing factors at different spatio-temporal scales. Glob Ecol Conserv, 2020, 23: e01008.

[106] Wang T, Deng LJ, Huang SJ, Xiong B, Ihtisham M, Zheng ZD, Zheng W, Qin ZY, Zhang MF, Sun GC, Wang J, Wang ZH. Genetic relationship, SPAD reading, and soluble sugar content as indices for evaluating the graft compatibility of Citrus interstocks. Biology, 2022, 11(11): 1639.

[107] Wang XY, Zhang YS, Tan YR, Tan Y, Bai JH, Gu DS, Ma ZZ, Du JH, Han ZY. Effects of light on the emissions of biogenic isoprene and monoterpenes: a review. Atmos Pollut Res, 2022, 13(5): 101397.

[108] Wen LC, Cao JM, Li W, Guo YF. Changes in volatile profile and related gene expression during senescence of tobacco leaves. J Sci Food Agric, 2023, 103(13): 6540-6552.

[109] Winters AJ, Adams MA, Bleby TM, Rennenberg H, Steigner D, Steinbrecher R, Kreuzwieser J. Emissions of isoprene, monoterpene and short-chained carbonyl compounds from Eucalyptus spp. in southern Australia. Atmos Environ, 2009, 43(19): 3035-3043.

[110] Woo J, Lee CJ. Sleep-enhancing effects of phytoncide via behavioral, electrophysiological, and molecular modeling approaches. Exp Neurobiol, 2020, 29(2): 120-129.

[111] Woo J, Yang H, Yoon M, Gadhe CG, Pae AN, Cho S, Lee CJ. 3-carene, a phytoncide from pine tree has a sleep-enhancing effect by targeting the GABA_A-benzodiazepine receptors. Exp Neurobiol, 2019, 28(5): 593-601.

[112] Wood SN, Goude Y, Shaw S. Generalized additive models for large data sets. J R Stat Soc Ser C Appl Stat, 2015, 64(1): 139-155.

[113] Wu J, Wang Q, Xu CY, Lun XX, Wang LX, Gao YS, Huang L, Zhang Q, Li LJ, Liu BX, Liu HX, Xu LJ. Biogenic volatile organic compounds in forest therapy base: a source of air pollutants or a healthcare function?. Sci Total Environ, 2024, 931: 172944.

[114] Xu C, Zhang YX, Wu D, Wu S, Xin XB, Pei SX. Variation characteristics of biogenic volatile organic compounds in three typical kinds of forests in Beijing. J Cent South Univ for Technol, 2023, 43(5): 130-139. in Chinese).

[115] Yan QL, Zhu JJ, Zhang JP, Yu LZ, Hu ZB. Spatial distribution pattern of soil seed bank in canopy gaps of various sizes in temperate secondary forests, Northeast China. Plant Soil, 2010, 329(1): 469-480.

[116] Yáñez-Serrano AM, Corbera J, Portillo-Estrada M, Janssens IA, Llusià J, Filella I, Peñuelas J, Preece C, Sabater F, Fernández-Martínez M. Drivers of biogenic volatile organic compound emissions in hygrophytic bryophytes. Sci Total Environ, 2024, 946: 174293.

[117] Yang WZ, Cao J, Wu Y, Kong FL, Li LY. Review on plant terpenoid emissions worldwide and in China. Sci Total Environ, 2021, 787: 147454.

[118] Yang H, Shen L, Li YR, Wu SQ, Zhang XY, Yang XP. Non-targeted metabolomic analysis reveals the mechanism of quality formation of Citrus flower-green tea. J Sci Food Agric, 2024, 104(10): 5807-5815.

[119] Yao B, He B, Peng JH, Song X, Zhao R, Sun Y, Zhang YF. The comprehensive review of eucalyptol: synthesis, metabolism, and therapeutic applications in disease treatment. Mol Biol Rep, 2025, 52(1): 346.

[120] Yenisoy-Karakaş S, Dörter M, Odabasi M. Intraday and interday variations of 69 volatile organic compounds (BVOCs and AVOCs) and their source profiles at a semi-urban site. Sci Total Environ, 2020, 723: 138028.

[121] Yuan HB, Wang L, Zhu JL, Ren JK, Liu RP, Li XQ, Chen Y, Hou H, Tu HT. Inhibitory effect of 2-methylvaleraldehyde on Botrytis cinerea and its application in postharvest strawberry. Postharvest Biol Technol, 2025, 230: 113775.

[122] Zhang JJ, Zhang PY. Study on photochemical degradation of nonanal-an odorant in gas phase by vacuum ultraviolet. Chin J Environ Eng, 2010, 4(6): 1368-1372. (in Chinese).

[123] Zhang CY, Wu ZN, Li ZH, Li HF, Lin JM. Inhibition effect of negative air ions on adsorption between volatile organic compounds and environmental particulate matter. Langmuir, 2020, 36(18): 5078-5083.

[124] Zhang BW, Jia YQ, Bai GK, Han HJ, Yang WZ, Xie WX, Li LY. Characterizing BVOC emissions of common plant species in northern China using real world measurements: towards strategic species selection to minimize ozone forming potential of urban greening. Urban for Urban Green, 2024, 96: 128341.

[125] Zhang J, Li M, Cheng RM, Ge ZX, Liu C, Zhang ZD. Stand structure and functional traits determine productivity of Larix principis-rupprechtii forests. Forests, 2024, 15(6): 924.

[126] Zhong X, Qi JH, Li HT, Dong LJ, Gao DM. Seasonal distribution of microbial activity in bioaerosols in the outdoor environment of the Qingdao coastal region. Atmos Environ, 2016, 140: 506-513.

[127] Zhou PT, Ganzeveld L, Taipale D, Rannik Ü, Rantala P, Rissanen MP, Chen DA, Boy M. Boreal forest BVOC exchange: emissions versus in-canopy sinks. Atmos Chem Phys, 2017, 17(23): 14309-14332.

[128] Zhou X, Chen Q, Sharma RP, Wang YH, He P, Guo JP, Lei YC, Fu LY. A climate sensitive mixed-effects diameter class mortality model for Prince Rupprecht larch (Larix gmelinii var. principis-rupprechtii) in Northern China. For Ecol Manage, 2021, 491: 119091.

[129] Zhou ZY, Xian JC, Wei WK, Xu C, Yang JF, Zhan RT, Ma DM. Volatile metabolic profiling and functional characterization of four terpene synthases reveal terpenoid diversity in different tissues of Chrysanthemum indicum L. Phytochemistry, 2021, 185: 112687.

[130] Zhou QQ, Song K, Zhang Y, Wan ZC, Hu K, Zhang ZR, Lu SH, Xie SD, Hu M, Guo S. Profiles of ambient semi- and intermediate- volatility organic compounds (S/IVOCs) in winter of urban Chengdu. Sci Total Environ, 2024, 957: 177480.

[131] Zhu JJ, Lu DL, Zhang WD. Effects of gaps on regeneration of woody plants: a meta-analysis. J for Res, 2014, 25(3): 501-510.

[132] Zhu JJ, Zhang GQ, Wang GG, Yan QL, Lu DL, Li XF, Zheng X. On the size of forest gaps: can their lower and upper limits be objectively defined?. Agric for Meteor, 2015, 213: 64-76.

[133] Zhu SX, Wang RF, Wang QY, Lei T, Cui GF. Coniferous and broad-leaved mixed forest has the optimal forest therapy environment among stand types in Xinjiang. Ecol Indic, 2024, 169: 112950.

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