Research Advances in the Regulation of Aroma Biosynthesis in Horticultural Crops by Plant Hormones and Environmental Factors

doi:10.16420/j.issn.0513-353x.2025-1069

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (2): 331-358.doi: 10.16420/j.issn.0513-353x.2025-1069

Research Advances in the Regulation of Aroma Biosynthesis in Horticultural Crops by Plant Hormones and Environmental Factors

XU Hongpeng¹^,², LU Jinping¹^,², JIANG Xiaonan¹^,², XIE Weilin¹^,², LIU Zhirou¹^,², BAN Jiayi¹^,², XUE Bingbing¹^,², YE Ruoyu¹^,², ZHANG Kexin¹^,², XIONG Yuchen¹^,², CHEN Jiamin¹^,², SONG Yefan¹^,², YUAN Fubang¹^,², WANG Yiqiao¹^,², LIU Zhuowen¹^,², GAO Junping¹^,²(), LIANG Yue¹^,²()

¹ Sanya Institute，China Agricultural University，Sanya，Hainan 572024，China
² College of Horticulture，China Agricultural University，Beijing 100193，China

Received:2025-12-02 Revised:2026-01-11 Online:2026-02-25 Published:2026-02-12

Abstract

Abstract:

The aroma quality of horticultural crops is an important economic trait determined by volatile organic compounds（VOCs）. Its biosynthesis is coordinately regulated by internal regulatory networks and external environmental factors. Phytohormones including jasmonic acid，ethylene，and auxin form complex regulatory networks by activating transcription factors such as MYB and bHLH，and precisely regulate the biosynthesis of aroma substances（e.g.，terpenoids and phenylpropanoids）through signal cross-talk. Signaling molecules，such as calcium ions，hydrogen peroxide，and nitric oxide，participate in regulating the expression of aroma biosynthesis-related genes by mediating hormone signals and environmental stimuli. Environmental factors such as light and temperature not only independently regulate aroma biosynthesis in horticultural crops，but also exert synergistic effects through interactions between photoreceptors and temperature sensors. This review summarizes recent research progress on the regulation of aroma biosynthesis in horticultural crops by phytohormones，signaling molecules，and environmental factors，aiming to provide a theoretical reference for plant aroma research and the breeding of horticultural varieties with high aroma quality.

Key words: phytohormones, signaling molecules, environmental factors, aroma biosynthesis

XU Hongpeng, LU Jinping, JIANG Xiaonan, XIE Weilin, LIU Zhirou, BAN Jiayi, XUE Bingbing, YE Ruoyu, ZHANG Kexin, XIONG Yuchen, CHEN Jiamin, SONG Yefan, YUAN Fubang, WANG Yiqiao, LIU Zhuowen, GAO Junping, LIANG Yue. Research Advances in the Regulation of Aroma Biosynthesis in Horticultural Crops by Plant Hormones and Environmental Factors[J]. Acta Horticulturae Sinica, 2026, 53(2): 331-358.

Figures/Tables 3

References 172

[1]	Abbas F, Ke Y G, Zhou Y W, Yu Y Y, Waseem M, Ashraf U, Li X Y, Yu R C, Fan Y P. 2021a. Genome-wide analysis of ARF transcription factors reveals HcARF5 expression profile associated with the biosynthesis of β-ocimene synthase in Hedychium coronarium. Plant Cell Reports, 40 (7):1269-1284. doi: 10.1007/s00299-021-02709-1
[2]	Abbas F, Ke Y G, Zhou Y W, Yu Y Y, Waseem M, Ashraf U, Wang C T, Wang X Y, Li X Y, Yue Y C, Yu R C, Fan Y P. 2021b. Genome-wide analysis reveals the potential role of MYB transcription factors in floral scent formation in Hedychium coronarium. Frontiers in Plant Science, 12:623742.
[3]	Alfieri M, Vaccaro M C, Cappetta E, Ambrosone A, De Tommasi N, Leone A. 2018. Coactivation of MEP-biosynthetic genes and accumulation of abietane diterpenes in Salvia sclarea by heterologous expression of WRKY and MYC2 transcription factors. Scientific Reports, 8:11009.
[4]	Alvarez S, Zhu M M, Chen S X. 2010. Proteomics of Arabidopsis redox proteins in response to methyl jasmonate. Journal of Proteomics, 73 (1):30-40. doi: 10.1016/j.jprot.2009.07.005 URL
[5]	Altenburger R, Matile P. 1990. Further observations on rhythmic emission of fragrance in flowers. Planta, 180 (2):194-197. doi: 10.1007/BF00193995 pmid: 24201944
[6]	An X H, Tian Y, Chen K Q, Liu X J, Liu D D, Xie X B, Cheng C G, Cong P H, Hao Y J. 2015. MdMYB9 and MdMYB11 are involved in the regulation of the JA-induced biosynthesis of anthocyanin and proanthocyanidin in apples. Plant and Cell Physiology, 56 (4):650-662. doi: 10.1093/pcp/pcu205 URL
[7]	Asai N, Nishioka T, Takabayashi J, Furuichi T. 2009. Plant volatiles regulate the activities of Ca²⁺-permeable channels and promote cytoplasmic calcium transients in Arabidopsis leaf cells. Plant Signaling & Behavior, 4 (4):294-300.
[8]	Aslam M Z, Lin X, Li X, Yang N, Chen L Q. 2020. Molecular cloning and functional characterization of CpMYC2 and CpBHLH13transcription factors from Wintersweet(Chimonanthus praecox L.). Plants-Basel, 9 (6):785.
[9]	Ayala-Zavala J F, Wang S Y, Wang C Y, González-Aguilar G A. 2005. Methyl jasmonate in conjunction with ethanol treatment increases antioxidant capacity,volatile compounds and postharvest life of strawberry fruit. European Food Research and Technology, 221 (6):731-738. doi: 10.1007/s00217-005-0069-z URL
[10]	Balbontín C, Gaete-Eastman C, Vergara M, Herrera R, Moya-León M A. 2007. Treatment with 1-MCP and the role of ethylene in aroma development of mountain papaya fruit. Postharvest Biology and Technology, 43 (1):67-77. doi: 10.1016/j.postharvbio.2006.08.005 URL
[11]	Bednarczyk D, Skaliter O, Kerzner S, Masci T, Shklarman E, Shor E, Vainstein A. 2025. The homeotic gene PhDEF regulates production of volatiles in petunia flowers by activating EOBI and EOBII. Plant Cell, 37 (2):koaf027.
[12]	Belhadj A, Telef N, Saigne C, Cluzet S, Barrieu F, Hamdi S, Mérillon J M. 2008. Effect of methyl jasmonate in combination with carbohydrates on gene expression of PR proteins,stilbene and anthocyanin accumulation in grapevine cell cultures. Plant Physiology and Biochemistry, 46 (5):493-499. doi: 10.1016/j.plaphy.2007.12.001 URL
[13]	Ben Zvi M M, Florence N Z, Masci T, Ovadis M, Shklarman E, Ben-Meir H, Tzfira T, Dudareva N, Vainstein A. 2008. Interlinking showy traits:co-engineering of scent and colour biosynthesis in flowers. Plant Biotechnology Journal, 6 (4):403-415. doi: 10.1111/pbi.2008.6.issue-4 URL
[14]	Boatright J, Negre F, Chen X, Kish C M, Wood B, Peel G, Orlova I, Gang D, Rhodes D, Dudareva N. 2004. Understanding in vivo benzenoid metabolism in petunia petal tissue. Plant Physiology, 135 (4):1993-2011. doi: 10.1104/pp.104.045468 pmid: 15286288
[15]	Borda A M, Clark D G, Huber D J, Welt B A, Nell T A. 2011. Effects of ethylene on volatile emission and fragrance in cut roses:the relationship between fragrance and vase life. Postharvest Biology and Technology, 59 (3):245-252. doi: 10.1016/j.postharvbio.2010.09.008 URL
[16]	Borda A M, Nell T A, Clark D G. 2007. The relationship between floral fragrance and vase life of cut roses. Acta Horticulturae, 755:235-241.
[17]	Cai H F, An X J, Han S, Jiang L, Yu M L, Ma R J, Yu Z F. 2018. Effect of 1-MCP on the production of volatiles and biosynthesis-related gene expression in peach fruit during cold storage. Postharvest Biology and Technology, 141:50-57. doi: 10.1016/j.postharvbio.2018.03.003 URL
[18]	Cai H F, Han S, Jiang L, Yu M L, Ma R J, Yu Z F. 2019. 1-MCP treatment affects peach fruit aroma metabolism as revealed by transcriptomics and metabolite analyses. Food Research International, 122:573-584. doi: S0963-9969(19)30026-2 pmid: 31229116
[19]	Cai H F, Han S, Yu M L, Ma R J, Yu Z F. 2020. Exogenous nitric oxide fumigation promoted the emission of volatile organic compounds in peach fruit during shelf life after long-term cold storage. Food Research International, 133:109135.
[20]	Cao W, Wang Y, Shi M, Hao X, Zhao W, Wang Y, Ren J, Kai G. 2018. Transcription factor SmWRKY1 positively promotes the biosynthesis of tanshinones in Salvia miltiorrhiza. Frontiers in Plant Science, 9:554. doi: 10.3389/fpls.2018.00554 URL
[21]	Cao Y, Liu L, Ma K, Wang W, Lv H, Gao M, Wang X, Zhang X, Ren S, Zhang N, Guo Y D. 2022. The jasmonate-induced bHLH gene SlJIG functions in terpene biosynthesis and resistance to insects and fungus. Journal of Integrative Plant Biology, 64 (5):1102-1115. doi: 10.1111/jipb.v64.5 URL
[22]	Carvalho S D, Schwieterman M L, Abrahan C E, Colquhoun T A, Folta K M. 2016. Light quality dependent changes in morphology antioxidant capacity and volatile production in Sweet Basil(Ocimum basilicum). Frontiers in Plant Science, 7:1328. doi: 10.3389/fpls.2016.01328 pmid: 27635127
[23]	Chen M, Yan T, Shen Q, Lu X, Pan Q, Huang Y, Tang Y, Fu X, Liu M, Jiang W, Lv Z, Shi P, Ma Y N, Hao X, Zhang L, Li L, Tang K. 2017. GLANDULAR TRICHOME-SPECIFIC WRKY 1 promotes artemisinin biosynthesis in Artemisia annua. New Phytologist, 214 (1):304-316. doi: 10.1111/nph.2017.214.issue-1 URL
[24]	Chen Y, Pang Q, Dai S, Wang Y, Chen S, Yan X. 2011. Proteomic identification of differentially expressed proteins in Arabidopsis in response to methyl jasmonate. Journal of Plant Physiology, 168 (10):995-1008. doi: 10.1016/j.jplph.2011.01.018 URL
[25]	Chen Yuxin, Luo Qiao, Pi Xuena, Wu Mengyue, Hou Jianqi, Wang Yanping, Chan Zhulong, Xiang Lin. 2025. Effects of methyl salicylate and methyl jasmonate on floral volatiles in tulips. Journal of Huazhong Agricultural University, 44 (5):88-97. (in Chinese)
	陈雨馨, 罗巧, 皮学娜, 邬梦玥, 侯健琦, 王艳平, 产祝龙, 向林. 2025. 水杨酸甲酯和茉莉酸甲酯对郁金香花香挥发物的影响. 华中农业大学学报, 44 (5):88-97.
[26]	Chuang Y C, Hung Y C, Tsai W C, Chen W H, Chen H H. 2018. PbbHLH 4 regulates floral monoterpene biosynthesis in Phalaenopsis orchids. Journal of Experimental Botany, 69 (18):4363-4377. doi: 10.1093/jxb/ery246 URL
[27]	Churchill A, Behan J. 2010. Comparison of methods used to study consumer emotions associated with fragrance. Food Quality and Preference, 21 (8):1108-1113. doi: 10.1016/j.foodqual.2010.07.006 URL
[28]	Cna'ani A, Spitzer-Rimon B, Ravid J, Farhi M, Masci T, Aravena-Calvo J, Ovadis M, Vainstein A. 2015. Two showy traits,scent emission and pigmentation,are finely coregulated by the MYB transcription factor PH4 in petunia flowers. New Phytologist, 208 (3):708-714. doi: 10.1111/nph.13534 pmid: 26111005
[29]	Colquhoun T A, Kim J Y, Wedde A E, Levin L A, Schmitt K C, Schuurink R C, Clark D G. 2011. PhMYB4 fine-tunes the floral volatile signature of Petunia × hybrida through PhC4H. Journal of Experimental Botany, 62 (3):1133-1143. doi: 10.1093/jxb/erq342 pmid: 21068208
[30]	Colquhoun T A, Verdonk J C, Schimmel B C, Tieman D M, Underwood B A, Clark D G. 2010. Petunia floral volatile benzenoid/phenylpropanoid genes are regulated in a similar manner. Phytochemistry, 71 (2-3):158-167. doi: 10.1016/j.phytochem.2009.09.036 pmid: 19889429
[31]	De Geyter N, Gholami A, Goormachtig S, Goossens A. 2012. Transcriptional machineries in jasmonate-elicited plant secondary metabolism. Trends in Plant Science, 17 (6):349-359. doi: 10.1016/j.tplants.2012.03.001 pmid: 22459758
[32]	De Paolis A, Caretto S, Quarta A, Di Sansebastiano G P, Sbrocca I, Mita G, Frugis G. 2020. Genome-wide identification of WRKY genes in Artemisia annua:characterization of a putative ortholog of AtWRKY40. Plants, 9 (12):1669. doi: 10.3390/plants9121669 URL
[33]	Ding W, Ouyang Q, Li Y, Shi T, Li L, Yang X, Ji K, Wang L, Yue Y. 2020. Genome-wide investigation of WRKY transcription factors in sweet osmanthus and their potential regulation of aroma synthesis. Tree Physiology, 40 (5):557-572. doi: 10.1093/treephys/tpz129 URL
[34]	Dixon J, Hewett E W. 2001. Exposure to hypoxia conditions alters volatile concentrations of apple cultivars. Journal of the Science of Food and Agriculture, 81 (1):22-29. doi: 10.1002/(ISSN)1097-0010 URL
[35]	Dodd A N, Kudla J, Sanders D. 2010. The language of calcium signaling. Annual Review of Plant Biology, 61:593-620. doi: 10.1146/annurev-arplant-070109-104628 pmid: 20192754
[36]	Dong T Y, Hao T Y, Hakeem A, Ren Y H, Fang J G. 2025. Synergistic variation in abscisic acid and brassinolide treatment signaling component alleviates fruit quality of‘Shine Muscat’grape during cold storage. Food Chemistry, 464:141584.
[37]	Dong T Y, Sun Z Y, Hao T Y, Fang J G. 2025. Melatonin maintains‘Shine Muscat’grape quality during low and high temperature storage. Postharvest Biology and Technology, 230:113807.
[38]	Dong Y, Zhang W, Li J, Wang D, Bai H, Li H, Shi L. 2022. The transcription factor LaMYC4 from lavender regulates volatile terpenoid biosynthesis. BMC Plant Biology, 22 (1):289. doi: 10.1186/s12870-022-03660-3
[39]	Dudareva N, Klempien A, Muhlemann J K, Kaplan I. 2013. Biosynthesis,function and metabolic engineering of plant volatile organic compounds. New Phytologist, 198 (1):16-32. doi: 10.1111/nph.12145 pmid: 23383981
[40]	Farré-Armengol G, Filella I, Llusia J, Peñuelas J. 2013. Floral volatile organic compounds:between attraction and deterrence of visitors under global change. Perspectives in Plant Ecology,Evolution and Systematics, 15 (1):56-67. doi: 10.1016/j.ppees.2012.12.002 URL
[173]	Yang Y Y, Wang S J, Leng P S, Wu J, Hu Z H. 2022b. Calcium and jasmonate signals mediate biosynthesis of the floral fragrance regulated by light quality in snapdragon. Plant Growth Regulation, 97 (1):91-100. doi: 10.1007/s10725-022-00807-y
[174]	Yang Y Y, Zhang X, Han M Z, Hu Z H, Wu J, Ma N, Leng P S, Zhou X F. 2023. LiMYB108 is involved in floral monoterpene biosynthesis induced by light intensity in Lilium‘Siberia’. Plant Cell Reports, 42 (4):763-773. doi: 10.1007/s00299-023-02995-x
[175]	Yang Z, Li Y, Gao F, Jin W, Li S, Kimani S, Yang S, Bao T, Gao X, Wag L. 2020. MYB 21 interacts with MYC2 to control the expression of terpene synthase genes in flowers of Freesia hybrida and Arabidopsis thaliana. Journal of Experimental Botany, 71 (14):4140-4158. doi: 10.1093/jxb/eraa184 URL
[176]	Yeh C W, Zhong H Q, Ho Y F, Tian Z H, Yeh K W. 2022. The diurnal emission of floral scent in Oncidium hybrid orchid is controlled by CIRCADIAN CLOCK ASSOCIATED 1 (CCA1) through the direct regulation on terpene synthase. BMC Plant Biology, 22 (1):472. doi: 10.1186/s12870-022-03850-z
[177]	Yu J, Liu X, Peng Y, Li Q, Han Y. 2022. Identification and characterization of transcription factors involved in geraniol biosynthesis in Rosa chinensis. International Journal of Molecular Sciences, 23 (23):14684. doi: 10.3390/ijms232314684 URL
[178]	Yu Z, Zhang G, Teixeira da Silva JA, Zhao C, Duan J. 2021. The methyl jasmonate-responsive transcription factor DobHLH4 promotes DoTPS10,which is involved in linalool biosynthesis in Dendrobium officinale during floral development. Plant Science, 309:110952.
[179]	Yu Z X, Li J X, Yang C Q, Hu W L, Wang L J, Chen X Y. 2012. The Jasmonate-Responsive AP2/ERF transcription factors AaERF1 and AaERF2 positively regulate artemisinin biosynthesis in Artemisia annua L. Molecular Plant, 5 (2):353-365. doi: 10.1093/mp/ssr087 URL
[180]	Yu Z X, Wang L J, Zhao B, Shan C M, Zhang Y H, Chen D F, Chen X Y. 2015. Progressive regulation of sesquiterpene biosynthesis in Arabidopsis and Patchouli(Pogostemon cablin)by the miR156-Targeted SPL transcription factors. Molecular Plant, 8 (1):98-110. doi: 10.1016/j.molp.2014.11.002 URL
[181]	Yuan X, Ma K F, Zhang M, Wang J, Zhang Q X. 2023. Integration of transcriptome and methylome analyses provides insight into the pathway of floral scent biosynthesis in Prunus mume. Frontiers in Genetics, 12:779557.
[182]	Zhang B, Shen J Y, Wei W W, Xi W P, Xu C J, Ferguson I, Chen K. 2010. Expression of genes associated with aroma formation derived from the fatty acid pathway during peach fruit ripening. Journal of Agricultural and Food Chemistry, 58 (10):6157-6165. doi: 10.1021/jf100172e pmid: 20415420
[183]	Zhang C, Dai Z, Ferrier T, Orduña L, Santiago A, Peris A, Wong D C J, Kappel C, Savoi S, Loyola R, Amato A, Kozak B, Li M, Liang A, Carrasco D, Meyer-Regueiro C, Espinoza C, Hilbert G, Figueroa-Balderas R, Cantu D, Arroyo-Garcia R, Arce-Johnson P, Claudel P, Errandonea D, Rodríguez-Concepción M, Duchêne E, Huang S C, Castellarin S D, Tornielli G B, Barrieu F, Matus J T. 2023. MYB 24 orchestrates terpene and flavonol metabolism as light responses to anthocyanin depletion in variegated grape berries. Plant Cell, 35 (12):4238-4265. doi: 10.1093/plcell/koad228 URL
[184]	Zhang H, Hedhili S, Montiel G, Zhang Y, Chatel G, Pré M, Gantet P, Memelink J. 2011. The basic helix-loop-helix transcription factor CrMYC2 controls the jasmonate-responsive expression of the ORCA genes that regulate alkaloid biosynthesis in Catharanthus roseus. The Plant Journal, 67 (1):61-71. doi: 10.1111/tpj.2011.67.issue-1 URL
[185]	Zhang Jia, Li Yueqing, Shan Xiaotong, Wang Li, Gao Xiang. 2024. The biosynthesis and regulation of floral volatile organic compounds and pigments in Freesia hybrida. Journal of Plant Genetic Resources, 25 (5):679-694. (in Chinese)
	张佳, 李月庆, 单晓彤, 王丽, 高翔. 2024. 香雪兰花色花香物质合成和调控研究进展. 植物遗传资源学报, 25 (5):679-694.
[186]	Zhang Lingjuan. 2024. Effect of exogenous salicylic acid and abscisic acid on the fruit volatile substances of Cerasus humilis[M. D. Dissertation]. Taigu: Shanxi Agricultural University. (in Chinese)
	张羚绢. 2024. 外源水杨酸和脱落酸处理对欧李果实挥发性物质的调控研究[硕士论文]. 太谷: 山西农业大学.
[187]	Zhang T, Sun M, Guo Y, Shi X, Yang Y, Chen J, Zheng T, Han Y, Bao F, Ahmad S. 2018. Overexpression of LiDXS and LiDXR from lily(Lilium‘Siberia’)enhances the terpenoid content in tobacco. Frontiers in Plant Science, 9:909. doi: 10.3389/fpls.2018.00909 URL
[188]	Zhang Z, Tao L, Gao L, Gao Y, Suo J, Yu W, Hu Y, Wei C, Farag M A, Wu J, Song L. 2023. Transcription factors TgbHLH95 and TgbZIP44 cotarget terpene biosynthesis gene TgGPPS in Torreya grandis nuts. Plant Physiology, 193 (3):1161-1176. doi: 10.1093/plphys/kiad385 URL
[189]	Zhang Z, Xu J, Chen Y, Wei J, Wu B. 2019. Nitric oxide treatment maintains postharvest quality of table grapes by mitigation of oxidative damage. Postharvest Biology and Technology, 152:9-18. doi: 10.1016/j.postharvbio.2019.01.015 URL
[190]	Zhao K, Yang W, Zhou Y, Zhang J, Li Y, Ahmad S, Zhang Q. 2017. Comparative transcriptome reveals benzenoid biosynthesis regulation as inducer of floral scent in the woody plant Prunus mume. Frontiers in Plant Science, 8:319. doi: 10.3389/fpls.2017.00319 pmid: 28344586
[191]	Zhao Qian. 2025. Identification and functional analysis of key genes involved in major terpenoid biosynthesis in Herbaceous Peony[Ph. D. Dissertation]. Yangling: Northwest A & F University. (in Chinese)
	赵前. 2025. 芍药花香主要萜类合成关键基因的挖掘及功能解析[博士论文]. 杨凌: 西北农林科技大学.
[192]	Zhao Weishu, Wan Chao, Hu Li, Zhang Qu, Yuan Yuan. 2022. Changes of major terpenoids contents and related genes expression level in jasmine after GAs treatment. Subtropical Plant Science, 51 (4):241-247. (in Chinese)
	赵炜淑, 万超, 胡莉, 张蕖, 袁媛. 2022. 外施GA₃对茉莉花主要萜类物质及其相关基因表达的影响. 亚热带植物科学, 51 (4):241-247.
[193]	Zhao X, Li Y, Shen J, Guo C, Li J, Chen M, Xu H, Li K. 2025. Analysis of the miRNA transcriptome in Aconitum vilmorinianum and its regulation of diterpenoid alkaloid biosynthesis. International Journal of Molecular Sciences, 26 (1):348. doi: 10.3390/ijms26010348 URL
[194]	Zhao Y T, Zhu X, Hou Y Y, Wang X Y, Li X H. 2020. Postharvest nitric oxide treatment delays the senescence of winter jujube(Zizyphus jujuba Mill. cv. Dongzao)fruit during cold storage by regulating reactive oxygen species metabolism. Scientia Horticulturae, 261:109009.
[195]	Zhao Y Y, Tang J X, Song C C, Qi S N, Lin Q, Cui Y, Ling J G, Duan Y Q. 2021. Nitric oxide alleviates chilling injury by regulating the metabolism of lipid and cell wall in cold-storage peach fruit. Plant Physiology and Biochemistry, 169:63-69. doi: 10.1016/j.plaphy.2021.10.039 pmid: 34763202
[196]	Zheng T, Lv J H, Sadeghnezhad E, Cheng J H, Jia H F. 2022. Transcriptomic and metabolomic profiling of strawberry during postharvest cooling and heat storage. Frontiers in Plant Science, 13:1009747.
[197]	Zhou D D, Liu Q, Zhu T, Li T T, Fan G J, Li X J, Wu C E. 2024. Effects of ultraviolet C on the quality and aroma volatile in peach fruit during postharvest storage. Food Chemistry, 456:139906.
[198]	Zhou Jinxin, Wei Tiantian, Yuan Xiaoyu, Feng Liguo, Sheng Lixia. 2019. Effects of exogenous methyl jasmonate treatment on the volatile composition and content of Rosa rugosa. Molecular Plant Breeding, 17 (14):4776-4784. (in Chinese)
	周金鑫, 魏恬恬, 袁晓雨, 冯立国, 生利霞. 2019. 外源茉莉酸甲酯处理对玫瑰花香成分及含量的影响. 分子植物育种, 17 (14):4776-4784.
[199]	Zhou L, Li J, Zeng T, Xu Z, Luo J, Zheng R, Wang Y, Wang C. 2022. TcMYB8,a R3-MYB transcription factor,positively regulates pyrethrin biosynthesis in Tanacetum cinerariifolium. International Journal of Molecular Sciences, 23 (20):12186. doi: 10.3390/ijms232012186 URL
[200]	Zhou Y, Xu Z, Chen X, Zhou J J, Wang S T, Xu Y Q. 2023. Functional characterization of the CfAOC and CfJMT gene promoters related to MelA biosynthesis in Cymbidium faberi. Plant Biotechnology Reports, 17 (2):243-253. doi: 10.1007/s11816-022-00780-y
[201]	Zhou S W, Zhang X Q, Han N X, Sun K Y, Liu X Y, Guo J, Tang Q. 2025. Study of the early regulatory mechanism of the Pichia kudriavzevii-induced resistance response in cherry tomato fruits. Food Bioscience, 66:106167
[202]	Zou Jingjing, Cai Xuan, Zeng Xiangling, Zheng Riru, Wang Caiyun. 2017. Changes of aroma-active compounds in different cultivars of Osmanthus fragrans during flowering. Acta Horticulturae Sinica, 44 (8):1517-1534. (in Chinese)
	邹晶晶, 蔡璇, 曾祥玲, 郑日如, 王彩云. 2017. 桂花不同品种开花过程中香气活性物质的变化. 园艺学报, 44 (8):1517-1534. doi: 10.16420/j.issn.0513-353x.2017-0050 URL
[203]	Zvi M M B, Shklarman E, Masci T, Kalev H, Debener T, Shafir S, Ovadis M, Vainstein A. 2012. PAP 1 transcription factor enhances production of phenylpropanoid and terpenoid scent compounds in rose flowers. New Phytologist, 195 (2):335-345. doi: 10.1111/nph.2012.195.issue-2 URL
[41]	Farré-Armengol G, Filella I, Llusià J, Niinemets Ü, Peñuelas J. 2015. Optimum temperature for floral terpene emissions tracks the mean temperature of the flowering season. Functional Plant Biology, 42 (9):851-857. doi: 10.1071/FP14279 pmid: 32480727
[42]	Fan X X, Lu N, Xu W S, Zhuang Y F, Jin J, Mao X J, Ren N. 2023. Response of flavor substances in tomato fruit to light spectrum and daily light integral. Plants-Basel, 12 (15):2832.
[43]	Farneti B, Paucek I, Khomenko I, Pennisi G, Pistillo A, Pastore C, Crepaldi A, Gianquinto G, Biasioli F, Orsini F, Spinelli F. 2026. Early postharvest application of LED lighting enhances strawberry quality during storage. Postharvest Biology and Technology, 232:114018.
[44]	Fu X, Peng B, Hassani D, Xie L, Liu H, Li Y, Chen T, Liu P, Tang Y, Li L, Zhao J, Sun X, Tang K. 2021. AaWRKY9 contributes to light- and jasmonate-mediated to regulate the biosynthesis of artemisinin in Artemisia annua. New Phytologist, 231 (5):1858-1874. doi: 10.1111/nph.v231.5 URL
[45]	Fu Qi, Wang Dan, Jing Weikun, Zhang Hao, Wang Huichun, Jian Hongying, Qiu Xianqin, Wang Qigang, Tang Kaixue, Yan Huijun. 2025. Function of rose carotenoid cleavage dioxygenase gene RcCCD4 in floral scent synthesis. Acta Horticulturae Sinica, 52 (3):623-634. (in Chinese)
	付琪, 王丹, 景维坤, 张颢, 王慧纯, 蹇洪英, 邱显钦, 王其刚, 唐开学, 晏慧君. 2025. 月季类胡萝卜素裂解双加氧酶基因RcCCD4在花香合成中的功能. 园艺学报, 52 (3):623-634. doi: 10.16420/j.issn.0513-353x.2024-0382 URL
[46]	Gao J, Chen Y, Gao M, Wu L, Zhao Y, Wang Y. 2023. LcWRKY17,a WRKY transcription factor from Litsea cubeba,effectively promotes monoterpene synthesis. International Journal of Molecular Sciences, 24 (8):7210. doi: 10.3390/ijms24087210 URL
[47]	Gao Ting. 2021. E3 Ligase HcPUB30 mediates HcWRKY1 to participate in regulation of floral scent of Hedychium coronarium[M. D. Dissertation]. Guangzhou: South China Agricultural University. (in Chinese)
	高婷. 2021. 姜花E3泛素连接酶HcPUB30介导HcWRKY1参与花香调控的机理研究[硕士论文]. 广州: 华南农业大学.
[48]	Gao Yang, Ye Nan, Kong Xiangjin, Ma Yinghu, Wu Juandi, Han Shunyu, Li Wei. 2022. Effect of exogenous application of methyl jasmonate and methyl dihydrojasmonate on the aroma compounds of Pinot Noir grapes and wine. Food and Fermentation Sciences & Technology, 58 (6):1-8. (in Chinese)
	高阳, 冶楠, 孔祥锦, 马英虎, 吴娟弟, 韩舜愈, 李蔚. 2022. 外源喷施茉莉酸甲酯类对‘黑比诺’葡萄及葡萄酒香气品质的影响. 食品与发酵科技, 58 (6):1-8.
[49]	Gong Z, Luo Y, Zhang W, Jian W, Zhang L, Gao X, Hu X, Yuan Y, Wu M, Xu X, Zheng X, Wu G, Li Z, Li Z, Deng W. 2021. A SIMYB75-centred cascade regulates trichome formation and transcriptional sesquiterpene accumulation in tomato. Journal of Experimental Botany, 72 (10):3806-3820. doi: 10.1093/jxb/erab086 URL
[50]	Guo Bing’ao. 2024. Functional analysis of Osmanthus fragrans WRKY21 transcription factor in the regulation of CCD4 gene expression and floral production[M. D. Dissertation]. Kaifeng: Henan University. (in Chinese)
	郭炳澳. 2024. 桂花OfWRKY21转录因子调控OfCCD4基因表达和花香产生的功能分析[硕士论文]. 开封: 河南大学.
[51]	Han J, Li T, Wang X, Zhang X, Bai X, Shao H, Wang S, Hu Z, Wu J, Leng P. 2022. AmMYB24 regulates floral terpenoid biosynthesis induced by blue light in snapdragon flowers. Frontiers in Plant Science, 13:885168.
[52]	Han X Y, Wang X Y, Shen C, Mo Y W, Tian R G, Mao L C, Luo Z S, Yang H Y. 2022. Exogenous ABA promotes aroma biosynthesis of postharvest kiwifruit after low-temperature storage. Planta, 255 (4):82. doi: 10.1007/s00425-022-03855-w pmid: 35257207
[53]	He X, Wang H, Yang J, Deng K, Wang T. 2018. RNA sequencing on Amomum villosum Lour. induced by MeJA identifies the genes of WRKY and terpene synthases involved in terpene biosynthesis. Genome, 61 (2):91-102. doi: 10.1139/gen-2017-0142 pmid: 29338341
[54]	Hong G J, Xue X Y, Mao Y B, Wang L J, Chen X Y. 2012. Arabidopsis MYC 2 interacts with DELLA proteins in regulating sesquiterpene synthase gene expression. Plant Cell, 24 (6):2635-2648. doi: 10.1105/tpc.112.098749 URL
[55]	Hu Y, Song A, Guan Z, Zhang X, Sun H, Wang Y, Yu Q, Fu X, Fang W, Chen F. 2023. CmWRKY41 activates CmHMGR2 and CmFPPS2to positively regulate sesquiterpenes synthesis in Chrysanthemum morifolium. Plant Physiology and Biochemistry, 196:821-829. doi: 10.1016/j.plaphy.2023.02.036 URL
[56]	Hu Z H, Li T J, Zheng J, Yang K, He X F, Leng P S. 2015. Ca²⁺ signal contributing to the synthesis and emission of monoterpenes regulated by light intensity in Lilium‘Siberia’. Plant Physiology and Biochemistry, 91:1-9. doi: 10.1016/j.plaphy.2015.03.005 URL
[57]	Hu Z H, Zhang H X, Leng P S, Zhao J, Wang W H, Wang S D. 2013. The emission of floral scent from Lilium‘Siberia’in response to light intensity and temperature. Acta Physiologiae Plantarum, 35 (5):1691-1700. doi: 10.1007/s11738-012-1211-8 URL
[58]	Huang J Q, Wen J Q, Wu F, Zhou P, Zhang J J, Wang L X, Li H L. 2025. Comparative transcriptomic analysis of loquat floral fragrance and hormone synthesis regulation across developmental stages in petals and stamens. Frontiers in Plant Science, 16:1574771.
[59]	Hüve K, Bichele I, Rasulov B, Niinemets U. 2011. When it is too hot for photosynthesis:heat-induced instability of photosynthesis in relation to respiratory burst cell permeability changes and H₂O₂ formation. Plant Cell and Environment, 34 (1):113-126. doi: 10.1111/pce.2011.34.issue-1 URL
[60]	Islam M R, Mitcham E J. 2024. Changes in raspberry sensory quality after harvest as affected by CO₂ atmospheres. Food Packaging and Shelf Life, 45:101335.
[61]	Ji Y, Xiao J, Shen Y, Ma D, Li Z, Pu G, Li X, Huang L, Liu B, Ye H, Wang H. 2014. Cloning and characterization of AabHLH1,a bHLH transcription factor that positively regulates artemisinin biosynthesis in Artemisia annua. Plant and Cell Physiology, 55 (9):1592-1604. doi: 10.1093/pcp/pcu090 URL
[62]	Jia H, Xie Z, Wang C, Shangguan L, Qian N, Cui M, Liu Z, Zheng T, Wang M, Fang J. 2017. Abscisic acid,sucrose,and auxin coordinately regulate berry ripening process of the Fujiminori grape. Functional & Integrative Genomics, 17 (4):441-457.
[63]	Jiang Y Y, Ren Y T, Zheng X M, Wang L H, Wang Y X, Wu C E, Qiao Y, Ma L L, Bai C M, Du H S, Zuo J H, Zheng Y Y. 2025. Pulsed light treatment mediated ethylene and abscisic acid synthesis effectively inhibited bell pepper turning red and softening. Food Chemistry, 499:147303.
[64]	Jimenez-Garcia S N, Vázquez-Cruz M A, Miranda-Lopez R, Garcia-Mier L, Guevara-González R G, Feregrino-Perez A A. 2018. Effect of elicitors as stimulating substances on sensory quality traits in color sweet bell pepper(Capsicum annuum L. cv. Fascinato and Orangela)grown under greenhouse conditions. Polish Journal of Food and Nutrition Sciences, 68 (4):359-365. doi: 10.2478/pjfns-2018-0003 URL
[65]	Jin N, Zhang D, Jin L, Wang S Y, Yang X T, Lei Y Z, Meng X, Xu Z Q, Sun J H, Lyu J, Yu J H. 2023. Controlling water deficiency as an abiotic stress factor to improve tomato nutritional and flavour quality. Food Chemistry-X, 19:100756.
[66]	Kawoosa T, Singh H, Kumar A, Sharma S K, Devi K, Dutt S, Vats S K, Sharma M, Ahuja P S, Kumar S. 2010. Light and temperature regulated terpene biosynthesis:hepatoprotective monoterpene picroside accumulation in Picrorhiza kurrooa. Functional & integrative genomics, 10 (3):393-404.
[67]	Ke Y G, Abbas F, Zhou Y W, Yu R C, Fan Y P. 2021. Auxin-responsive R2R3-MYB transcription factors HcMYB1 and HcMYB2activate volatile biosynthesis in Hedychium coronarium flowers. Frontiers in Plant Science, 12:710826.
[68]	Ke Yanguo. 2019. The regulation of floral scent metabolism by auxin signal and Auxin-dependent factors in Hedychium coronarium[Ph. D. Dissertation]. Guangzhou: South China Agricultural University. (in Chinese)
	柯艳果. 2019. 生长素信号及其应答因子对姜花花香物质代谢的调控研究[博士论文]. 广州: 华南农业大学.
[69]	Keinänen M, Oldham N J, Baldwin I T. 2001. Rapid HPLC screening of jasmonate-induced increases in tobacco alkaloids,phenolics,and diterpene glycosides in Nicotiana attenuata. Journal of Agricultural and Food Chemistry, 49 (8):3553-3558. doi: 10.1021/jf010200+ pmid: 11513627
[70]	Kim H J, Chen F, Wang X, Rajapakse N C. 2006. Effect of methyl jasmonate on secondary metabolites of sweet basil(Ocimum basilicum L.). Journal of Agricultural and Food Chemistry, 54 (6):2327-2332. doi: 10.1021/jf051979g URL
[71]	Kivimäenpää M, Riikonen J, Valolahti H, Elina H, Holopainen JK, Holopainen T. 2022. Effects of elevated ozone and warming on terpenoid emissions and concentrations of Norway spruce depend on needle phenology and age. Tree physiology, 42 (8):1570-1586. doi: 10.1093/treephys/tpac019 URL
[72]	Knudsen J T, Eriksson R, Gershenzon J, Stahl B. 2006. Diversity and distribution of floral scent. Botanical Review, 72 (1):1-120. doi: 10.1663/0006-8101(2006)72[1:DADOFS]2.0.CO;2 URL
[73]	Kothari S K, Virendra S, Kamla S. 1987. Effect of rates and methods of phosphorus application on herb and oil yields and nutrient concentrations in Japanese mint(Mentha arvensis L.). Journal of Agricultural Science, 108 (3):691-693.
[74]	Lan Xi, Zhang Lehua, Zhang Jinzheng, Cui Hongxia, Jiang Chuangdao, Shi Lei. 2012. Research progress of rhododendron breeding. Acta Horticulturae Sinica, 39 (9):1829-1838. (in Chinese)
	兰熙, 张乐华, 张金政, 崔洪霞, 姜闯道, 石雷. 2012. 杜鹃花属植物育种研究进展. 园艺学报, 39 (9):1829-1838.
[75]	Lan Y A, Zhang K M, Wang L N, Liang X Y, Liu H X, Zhang X Y, Jiang N Q, Wu M, Yan H W, Xiang Y. 2023. The R2R3-MYB transcription factor OfMYB21 positively regulates linalool biosynthesis in Osmanthus fragrans flowers. International Journal of Biological Macromolecules, 249:126099.
[76]	Lecourieux D, Mazars C, Pauly N, Ranjeva R, Pugin A. 2002. Analysis and effects of cytosolic free calcium increases in response to elicitors in Nicotiana plumbaginifolia cells. Plant Cell, 14 (11):2627-2641. doi: 10.1105/tpc.005579 URL
[77]	Ledderhof D, Reynolds A G, Manin L, Brown R. 2014. Influence of water status on sensory profiles of Ontario Pinot noir wines. LWT-Food Science and Technology, 57 (1):65-82. doi: 10.1016/j.lwt.2013.12.010 URL
[78]	Li C Y, Leopold A L, Sander G W, Shanks J V, Zhao L, Gibson S I. 2015. CrBPF1 overexpression alters transcript levels of terpenoid indole alkaloid biosynthetic and regulatory genes. Frontiers in Plant Science, 6:818.
[79]	Li D, Yang J S, Dai Z B, Chen Y J, Shao Z, Wang C H, Jin X, Wang Y H, Feng L D. 2024. Prohexadione-calcium improves grape quality by regulating endogenous hormones,sugar and acid metabolism and related enzyme activities in grape berries. BMC Plant Biology, 24 (1):122. doi: 10.1186/s12870-024-04803-4
[80]	Li H Z, Song K K, Zhang X H, Hou J, Yang L. 2025. Aroma formation in jasmine tea:insights from metabolomics and microbial community analyses during multiple scenting processes. Lwt-Food Science and Technology, 233:118540.
[81]	Li Haiyan, Li Huogen, Yang Xiulian, Yue Yuanzheng, Xu Chen. 2018. Advances studies on the synthesis and regulation of floral substances in plant. Molecular Plant Breeding, 16 (1):123-129. (in Chinese)
	李海燕, 李火根, 杨秀莲, 岳远征, 徐晨, 丁文杰, 王良桂. 2018. 植物花香物质合成与调控研究进展. 分子植物育种, 16 (1):123-129.
[82]	Li L X, Fang Y, Li D, Zhu Z H, Zhang Y, Tang Z Y, Li T, Chen X S, Feng S Q. 2023. Transcription factors MdMYC2 and MdMYB85 interact with ester aroma synthesis gene MdAAT1 in apple. Plant Physiology, 193 (4):2442-2458. doi: 10.1093/plphys/kiad459 pmid: 37590971
[83]	Li T, Xu Y X, Zhang L C, Ji Y L, Tan D M, Yuan H, Wang A D. 2017. The Jasmonate-activated transcription factor MdMYC2 regulates ETHYLENE RESPONSE FACTOR and ethylene biosynthetic genes to promote ethylene biosynthesis during apple fruit ripening. Plant Cell, 29 (6):1316-1334. doi: 10.1105/tpc.17.00349 URL
[84]	Li X, Xu Y Y, Shen S L, Yin X R, Klee H, Zhang B, Chen K S. 2017. Transcription factor CitERF71 activates the terpene synthase gene CitTPS16involved in the synthesis of E-geraniol in sweet orange fruit. Journal of Experimental Botany, 68 (17):4929-4938. doi: 10.1093/jxb/erx316 URL
[85]	Li Y, Li R, Shang J, Zhao K, Sui Y, Liu Z, Yan H, Bao M, Liang M, Zhao Q, Yuan YW, Ning G. 2025. A de novo-originated gene drives rose scent diversification. Cell, 188 (22):6121-6137. doi: 10.1016/j.cell.2025.08.011 URL
[86]	Li Y Q, Bao T T, Zhang J, Li H J, Shan X T, Yan H J, Kimani S, Zhang L S, Gao X. 2025. The coordinated interaction or regulation between floral pigments and volatile organic compounds. Horticultural Plant Journal, 11 (2):463-485. doi: 10.1016/j.hpj.2024.01.002 URL
[87]	Liu F, Xiao Z N, Yang L, Chen Q, Shao L, Liu J X, Yu Y X. 2017. PhERF6 interacting with EOBI negatively regulates fragrance biosynthesis in petunia flowers. New Phytologist, 215 (4):1490-1502. doi: 10.1111/nph.2017.215.issue-4 URL
[88]	Liu Mengxi, Tian Ruiping, Zeng Qilong, Yu Hong, Wei Jiguang. 2021. Effects of spraying methyl jasmonate on volatile components of ripe fruits of Vaccinium corymbosum‘Guleoast’. Journal of Plant Resources and Environment, 30 (2):71-73. (in Chinese)
	刘梦溪, 田瑞平, 曾其龙, 於虹, 韦继光. 2021. 喷施茉莉酸甲酯对高丛越橘品种‘海岸’成熟果实挥发性成分的影响. 植物资源与环境学报, 30 (2):71-73.
[89]	Liu Xiaozhou, Fan Yanping, Yu Rangcai, Yue Yuechong, Li Xinyue, Yu Yunyi. 2018. Effect of ethylene and 1-MCP on the terpenoid fragrance of Hedychium coronarium and the expressions of related genes. Journal of South China Agricultural University, 39 (4):68-72. (in Chinese)
	刘晓洲, 范燕萍, 余让才, 岳跃冲, 李昕悦, 玉云祎. 2018. 乙烯和1-MCP对白姜花萜类香气及相关基因表达的影响. 华南农业大学学报, 39 (4):68-72.
[90]	Lu H Y, Luo Z S, Li D, Jiang Y H, Li L. 2021. FaMYB11 promotes the accumulation of volatile esters by regulating FaLOX5 during strawberry(Fragaria × ananassa)ripening. Postharvest Biology and Technology, 178:111560.
[91]	Lu H Y, Luo Z S, Wang L, Liu W S, Li D, Belwal T, Xu Y Q, Li L. 2020. FaMYB9 is involved in the regulation of C6 volatile biosynthesis in strawberry. Plant Science, 293:110422.
[92]	Luo M, Zhou X, Hao Y, Sun H, Zhou Q, Sun Y, Ji S J. 2021. Methyl jasmonate pretreatment improves aroma quality of cold-stored‘Nanguo’pears by promoting ester biosynthesis. Food Chemistry, 338:127846.
[93]	Ma C, Li R, Sun Y, Zhang M, Li S, Xu Y, Song J, Li J, Qi J, Wang L, Wu J. 2023. ZmMYC2s play important roles in maize responses to simulated herbivory and jasmonate. Journal of Integrative Plant Biology, 65 (4):1041-1058. doi: 10.1111/jipb.v65.4 URL
[94]	Magalhaes H C R, Alves E G, Garruti D D, Massaretto I L, Purgatto E. 2021. Effect of postharvest methyl jasmonate and ethylene treatments on the biosynthesis of volatile compounds of hot pepper fruits. Scientia horticulturae, 289:110477.
[95]	Mandaokar A, Thines B, Shin B, Lange B M, Choi G, Koo Y J, Yoo Y J, Choi Y D, Choi G, Browse J. 2006. Transcriptional regulators of stamen development in Arabidopsis identified by transcriptional profiling. The Plant Journal, 46 (6):984-1008. doi: 10.1111/tpj.2006.46.issue-6 URL
[96]	Matschi S, Hake K, Herde M, Hause B, Romeis T. 2015. The calcium-dependent protein kinase CPK 28 regulates development by inducing growth phase-specific spatially restricted alterations in jasmonic acid levels independent of defense responses in Arabidopsis. Plant Cell, 27 (3):591-606. doi: 10.1105/tpc.15.00024 URL
[97]	Matsui K, Fukutomi S, Wilkinson J, Hiatt B, Knauf V, Kajwara T. 2001. Effect of overexpression of fatty acid 9-hydroperoxide lyase in tomatoes (Lycopersicon esculentum Mill.). Journal of Agricultural and Food Chemistry, 49 (11):5418-5424. doi: 10.1021/jf010607e pmid: 11714337
[98]	Mcdonald R E, Mccollum T G, Baldwin E A. 1996. Prestorage heat treatments influence free sterols and flavor volatiles of tomatoes stored at chilling temperature. Journal of the American Society for Horticultural Science, 121 (3):531-536. doi: 10.21273/JASHS.121.3.531 URL
[99]	Medina-Puche L, Cumplido-Laso G, Amil-Ruiz F, Hoffmann T, Ring L, Rodríguez-Franco A, Caballero J L, Schwab W, Muñoz-Blanco J, Blanco-Portales R. 2014. MYB 10 plays a major role in the regulation of flavonoid/phenylpropanoid metabolism during ripening of Fragaria × ananassa fruits. Journal of Experimental Botany, 65 (2):401-417. doi: 10.1093/jxb/ert377 pmid: 24277278
[100]	Medina-Puche L, Molina-Hidalgo F J, Boersma M, Schuurink R C, López-Vidriero I, Solano R, Franco-Zorrilla J M, Caballero J L, Blanco-Portales R, Muñoz-Blanco J. 2015. An R2R3-MYB transcription factor regulates eugenol production in ripe strawberry fruit receptacles. Plant Physiology, 168 (2):598-614. doi: 10.1104/pp.114.252908 pmid: 25931522
[101]	Mertens J, Van Moerkercke A, Vanden Bossche R, Pollier J, Goossens A. 2016. Clade IVa basic helix-loop-helix transcription factors form part of a conserved jasmonate signaling circuit for the regulation of bioactive plant terpenoid biosynthesis. Plant and Cell Physiology, 57 (12):2564-2575. pmid: 27694525
[102]	Neill S J, Desikan R, Hancock J T. 2003. Nitric oxide signalling in plants. New Phytologist, 159 (1):11-35. doi: 10.1046/j.1469-8137.2003.00804.x pmid: 33873677
[103]	Niu Tongfei, Xue Xian, Guo Lili, Yu Min, Zhang Chenjie, Xu Xin’ao, Li Ruiya, Hou Xiaogai. 2023. Effects of exogenous methyl jasmonate on volatile components and content of Paeonia suffruticosa‘Luoyanghong’in greenhouse. Scientia Silvae Sinicae, 59 (5):53-60. (in Chinese).
	牛童非, 薛娴, 郭丽丽, 郁敏, 张晨洁, 徐鑫傲, 李瑞雅, 侯小改. 2023. 外源茉莉酸甲酯对温室牡丹‘洛阳红’挥发性成分及含量的影响. 林业科学, 59 (5):53-60.
[104]	Ortiz A, Echeverría G, Graell J, Lara I. 2009. Calcium dips enhance volatile emission of cold-stored‘Fuji Kiku-8’apples. Journal of Agricultural and Food Chemistry, 57 (11):4931-4938. doi: 10.1021/jf9003576 pmid: 19449874
[105]	Ortiz A, Echeverría G, Graell J, Lara I. 2010a. The emission of flavour-contributing volatile esters by‘Golden Reinders’apples is improved after mid-term storage by postharvest calcium treatment. Postharvest Biology and Technology, 57 (2):114-123. doi: 10.1016/j.postharvbio.2010.03.006 URL
[106]	Ortiz A, Graell J, López M L, Echeverría G, Lara I. 2010b. Volatile ester-synthesising capacity in‘Tardibelle’peach fruit in response to controlled atmosphere and 1-MCP treatment. Food Chemistry, 123 (3):698-704. doi: 10.1016/j.foodchem.2010.05.037 URL
[107]	Patrick R M, Huang X Q, Dudareva N, Li Y. 2021. Dynamic histone acetylation in floral volatile synthesis and emission in petunia flowers. Journal of Experimental Botany, 72 (10):3704-3722. doi: 10.1093/jxb/erab072 pmid: 33606881
[108]	Peng X, Wang B, Wang X L, Ni B B, Zuo Z J. 2020. Variations in aroma and specific flavor in strawberry under different colored light-quality selective plastic film. Flavour and Fragrance Journal, 35 (3):350-359. doi: 10.1002/ffj.3569
[109]	Petrovic L, Popovic N, Todorovic M, Matekalo D, Milutinovic M, Nakarada D, Bozunovi J, Zivkovic J N, Dmitrovic S, Siler B, Banjanac T, Filipovic B, Gasic U, Mojovic M, Misic D, Skoric M. 2025. Metabolomic and transcriptomic responses to waterlogging stress and hydrogen peroxide elicitation:impact on the production of iridoids and phenolics in leaves of two Nepeta species. Industrial Crops and Products, 235:121762.
[110]	Pirbalouti A G, Rahimmalek M, Elikaei-Nejhad L, Hamedi B. 2014. Essential oil compositions of summer savory under foliar application of jasmonic acid and salicylic acid. Journal of Essential Oil Research, 26 (5):342-347. doi: 10.1080/10412905.2014.922508 URL
[111]	Qiu D, Pan X, Wilson I W, Li F, Liu M, Teng W, Zhang B. 2009. High throughput sequencing technology reveals that the taxoid elicitor methyl jasmonate regulates microRNA expression in Chinese yew(axus chinensis). Gene, 436 (1-2):37-44. doi: 10.1016/j.gene.2009.01.006 URL
[112]	Ramya M, Lee S Y, An H R, Park P M, Kim N S, Park P H. 2019. MYB 1 transcription factor regulation through floral scent in Cymbidium cultivar ‘Sael Bit’. Phytochemistry Letters, 32:181-187. doi: 10.1016/j.phytol.2019.06.007 URL
[113]	Reddy V A, Wang Q, Dhar N, Kumar N, Venkatesh P N, Rajan C, Panicker D, Sridhar V, Mao H Z, Sarojam R. 2017. Spearmint R2R3-MYB transcription factor MsMYB negatively regulates monoterpene production and suppresses the expression of geranyl diphosphate synthase large subunit(MsGPPS. LSU). Plant Biotechnology Journal, 15 (9):1105-1119. doi: 10.1111/pbi.2017.15.issue-9 URL
[114]	Reeves P H, Ellis C M, Ploense S E, Wu M F, Yadav V, Tholl D, Chételat A, Haupt I, Kennerley B J, Hodgens C, Farmer E E, Nagpal P, Reed J W. 2012. A regulatory network for coordinated flower maturation. PLoS Genetics, 8 (2):e1002506. doi: 10.1371/journal.pgen.1002506 URL
[115]	Ren Tingting, Zhang Tingxiu, Zhan Liang, Zhang Zhichang, Ren Yiran, Ji Xinglong, Wang Peipei, Liu Yuanxia, Meng Qingfu, Liu Gengsen, Fang Jinggui, Leng Xiangpeng. 2025. Effect of different light quality on berry quality of‘Shine Muscat’grape. Acta Horticulturae Sinica, 52 (9):2464-2476. (in Chinese) doi: 10.16420/j.issn.0513-353x.2024-0877 URL
	任婷婷, 张廷秀, 战良, 张志昌, 任怡然, 季兴龙, 王培培, 刘源霞, 孟庆福, 刘更森, 房经贵, 冷翔鹏. 2025. 不同光质补光对设施栽培‘阳光玫瑰’葡萄果实品质的影响. 园艺学报, 52 (9):2464-2476. doi: 10.16420/j.issn.0513-353x.2024-0877 URL
[116]	Rubio-Rodríguez E, Vera-Reyes I, Rodríguez-Hernández A A, López-Laredo A R, Ramos-Valdivia A C, Trejo-Tapia G. 2023. Mixed elicitation with salicylic acid and hydrogen peroxide modulates the phenolic and iridoid pathways in Castilleja tenuiflora plants. Scientific Reports, 258 (1):20.
[117]	Sarid O, Zaccai M. 2016. Changes in mood states are induced by smelling familiar and exotic fragrances. Frontiers in Psychology, 7:1724.
[118]	Schade F, Legge R L, Thompson J E. 2001. Fragrance volatiles of developing and senescing carnation flowers. Phytochemistry, 56 (7):703-710. pmid: 11314956
[119]	Sexton R, Stopford A P, Moodie W T, Porter A E A. 2005. Aroma production from cut sweet pea flowers(Lathyrus odoratus):the role of ethylene. Physiologia Plantarum, 124 (3):381-389. doi: 10.1111/ppl.2005.124.issue-3 URL
[120]	Shan Q W, Wang Y P, Li J, Zhang Y, Chen K L, Liang Z, Zhang K, Liu J X, Xi J J, Qiu J L, Gao C X. 2013. Targeted genome modification of crop plants using a CRISPR-Cas system. Nature Biotechnology, 31 (8):686-688. doi: 10.1038/nbt.2650 pmid: 23929338
[121]	Shang Tong. 2023. Changes of aroma content of Rosa rugosa in response to MeJA treatment and related key gene analysis[M. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	尚彤. 2023. 玫瑰芳香物质响应茉莉酸甲酯处理的变化及关键基因挖掘[硕士论文]. 泰安: 山东农业大学.
[122]	Shen S L, Yin X R, Zhang B, Xie X L, Jiang Q, Grierson D, Chen K S. 2016. CitAP2.10 activation of the terpene synthase CsTPS1 is associated with the synthesis of (+)-valencene in‘Newhall’orange. Journal of Experimental Botany, 67 (14):4105-4115. doi: 10.1093/jxb/erw189 URL
[123]	Shi S C, Duan G Y, Li D D, Wu J, Liu X T, Hong B, Yi M F, Zhang Z. 2018. Two-dimensional analysis provides molecular insight into flower scent of Lilium‘Siberia’. Scientific Reports, 8:5352. doi: 10.1038/s41598-018-23588-9
[124]	Shi Xuejun. 2020. BTH regulates monoterpene volatile in Lilium‘Siberia’by regulating the content of H₂O₂[M. D. Dissertation]. Beijing: Beijing Forestry University. (in Chinese)
	石雪珺. 2020. BTH通过调控H₂O₂含量调节百合'西伯利亚'单萜类挥发物释放[硕士论文]. 北京: 北京林业大学.
[125]	Shi Y, Wang Y B, Ma Y, Xu Y, Zhao X Y, Zhang C. 2020. Red light exposure delays appearance and aroma deterioration of fresh-cut watermelon during retail display. Journal of Food Quality, 2020:1840648.
[126]	Shu G P, Tang Y L, Nagdy M M, Zeng L J, Zhang F Y, Chen M, Mao J X, Liao Z H. 2025. AabZlP1 positively regulates the biosynthesis of monoterpenes and sesquiterpenes in Artemisia annua. Industrial Crops and Products, 236:122024.
[127]	Skaliter O, Bednarczyk D, Shor E, Shklarman E, Manasherova E, Aravena-Calvo J, Kerzner S, Cnaani A, Jasinska W, Masci T, Dvir G, Edelbaum O, Rimon B, Brotman Y, Cohen H, Vainstein A. 2023. The R2R3-MYB transcription factor EVER controls the emission of petunia floral volatiles by regulating epicuticular wax biosynthesis in the petal epidermis. Plant Cell, 36 (1):174-193. doi: 10.1093/plcell/koad251 URL
[128]	Skibbe M, Qu N, Galis I, Baldwin I T. 2008. Induced plant defenses in the natural environment:Nicotiana attenuata WRKY3 and WRKY 6 coordinate responses to herbivory. Plant Cell, 20 (7):1984-2000. doi: 10.1105/tpc.108.058594 URL
[129]	Song F Y, Sun Z K, Guo H J, Chen J L, Jin X W. 2024. Low temperature conditioning and nitric oxide treatment reduced chilling injury and altered fatty acid metabolism of flat peaches. Postharvest Biology and Technology, 218:113191.
[130]	Souza S R, Blande J D, Holopainen J K. 2013. Pre-exposure to nitric oxide modulates the effect of ozone on oxidative defenses and volatile emissions in lima bean. Environmental Pollution, 179:111-119. doi: 10.1016/j.envpol.2013.03.065 pmid: 23669460
[131]	Soyk S, Müller N A, Park S J, Schmalenbach I, Jiang K, Hayama R, Zhang L, Van Eck J, Jiménez-Gómez J M, Lippman Z B. 2017. Variation in the flowering gene SELF PRUNING 5G promotes day-neutrality and early yield in tomato. Nature Genetics, 49 (1):162-168. doi: 10.1038/ng.3733
[132]	Staudt M, Lhoutellier L. 2011. Monoterpene and sesquiterpene emissions from Quercus coccifera exhibit interacting responses to light and temperature. Source Title, 8 (9):2757-2771.
[133]	Sun Xiaoling, Dong Wenxia, Cai Xiaoming, Gui Lianyou, Chen Zongmao. 2016. Variation in tea-plant volatiles induced by exogenous application of different concentrations of methyl jasmonate. Chinese Journal of Applied Entomology, 53 (3):499-506. (in Chinese)
	孙晓玲, 董文霞, 蔡晓明, 桂连友, 陈宗懋. 2016. 外用不同浓度茉莉酸甲酯诱导的茶树挥发物的种类和时序变化. 应用昆虫学报, 53 (3):499-506.
[134]	Tiliwa E S, Boateng I D, Zhou C S, Xu B G. 2023. Role of drying technologies on the drying kinetics,physical quality,aroma,and enzymatic activity of pineapple slices. Sustainability, 15 (22):15906. doi: 10.3390/su152215906 URL
[135]	Tobaruela E D, Meza S L R, Massaretto I L, Pascoal G B, Freschi L, Purgatto E. 2024. Metabolomics reveals how ethylene-auxin interaction delays tomato ripening maintaining fruit nutritional and sensorial quality. International Journal of Food Science and Technology, 59 (6):4315-4325. doi: 10.1111/ijfs.v59.6 URL
[136]	Tu M L, Liu N Y W, He Z S, Dong X M, Gao T Y, Zhu A D, Yang J B, Zhang S B. 2025. Integrative omics reveals mechanisms of biosynthesis and regulation of floral scent in Cymbidium tracyanum. Plant Biotechnology Journal, 23 (6):2162-2181. doi: 10.1111/pbi.v23.6 URL
[137]	Underwood B A, Tieman D M, Shibuya K, Dexter R J, Loucas H M, Simkin A J, Sims C A, Schmelz E A, Klee H J, Clark D G. 2005. Ethylene-regulated floral volatile synthesis in Petunia corollas. Plant Physiology, 138 (1):255-266. pmid: 15849311
[138]	van der Fits L, Memelink J. 2000. ORCA 3 a jasmonate-responsive transcriptional regulator of plant primary and secondary metabolism. Science, 289 (5477):295-297. doi: 10.1126/science.289.5477.295 pmid: 10894776
[139]	van Moerkercke A, Haring M A, Schuurink R C. 2011. The transcription factor emission of benzenoids II activates the MYB ODORANT 1 promoter at a MYB binding site specific for fragrant petunias. Plant Journal, 67 (5):917-928. doi: 10.1111/tpj.2011.67.issue-5 URL
[140]	van Moerkercke A, Steensma P, Gariboldi I, Espoz J, Purnama P C, Schweizer F, Miettinen K, Vanden Bossche R, De Clercq R, Memelink J, Goossens A. 2016. The basic helix-loop-helix transcription factor BIS 2 is essential for monoterpenoid indole alkaloid production in the medicinal plant Catharanthus roseus. The Plant Journal, 88 (1):3-12. doi: 10.1111/tpj.13230 pmid: 27342401
[141]	van Moerkercke A, Steensma P, Schweizer F, Pollier J, Gariboldi I, Payne R, Bossche R V, Miettinen K, Espoz J, Purnama P C, Kellner F, Seppänen-Laakso T, O'Connor S E, Rischer H, Memelink J, Goossens A. 2015. The bHLH transcription factor BIS1 controls the iridoid branch of the monoterpenoid indole alkaloid pathway in Catharanthus roseus. Proceedings of the National Academy of Sciences of the United States of America, 112 (26):8130-8135.
[142]	Verdonk J C, Haring M A, Tunen A J van, Schuurink R C. 2005. ODORANT 1 regulates fragrance biosynthesis in petunia flowers. Plant Cell, 17 (5):1612-1624. doi: 10.1105/tpc.104.028837 pmid: 15805488
[143]	Wang C, Xing J, Chin C K, Ho C T, Martin C E. 2001. Modification of fatty acids changes the flavor volatiles in tomato leaves. Phytochemistry, 58 (2):227-232. doi: 10.1016/s0031-9422(01)00233-3 pmid: 11551543
[144]	Wang Chutian. 2021. The Regulation of floral scent formation by ABA and ABA signaling components in Hedychium coronarium[Ph. D. Dissertation]. Guangzhou: South China Agricultural University. (in Chinese)
	王楚天. 2021. ABA及其信号元件对姜花花香物质合成的调控[博士论文]. 广州: 华南农业大学.
[145]	Wang F J, Wang C L, Liu P Q, Lei C L, Hao W, Gao Y, Liu Y G, Zhao K J. 2016. Enhanced rice blast resistance by CRISPR/Cas9-targeted mutagenesis of the ERF transcription factor gene OsERF922. PLoS ONE, 11 (4):e0154027. doi: 10.1371/journal.pone.0154027 URL
[146]	Wang J F, VanderWeide J, Yan Y F, Tindjau R, Pico J, Deluc L, Zandberg W F, Castellarin S D. 2022. Impact of hormone applications on ripening-related metabolites in Gewurztraminer grapes(Vitis vinifera L.):The key role of jasmonates in terpene modulation. Food Chemistry, 388:132948.
[147]	Wang Q, Liu H, Zhang M, Liu S, Hao Y, Zhang Y. 2020. MdMYC2 and MdERFpositively co-regulate α-farnesene biosynthesis in apple. Frontiers in Plant Science, 11:512844.
[148]	Wang Qi, Liu Guangda, Su Lei. 2020. Effects of salicylic acid and methyl jasmonate on expression of terpenoid secondary metabolites related genes of miniature rose. Molecular Plant Breeding, 18 (3):797-803. (in Chinese)
	王启, 刘广达, 苏蕾. 2020. 水杨酸和茉莉酸甲酯对微型月季萜类次生代谢产物相关基因表达的影响. 分子植物育种, 18 (3):797-803.
[149]	Wang S S, Saito T, Ohkawa K, Ohara H, Suktawee S, Ikeura H, Kondo S. 2018. Abscisic acid is involved in aromatic ester biosynthesis related with ethylene in green apples. Journal of Plant Physiology, 221:85-93. doi: S0176-1617(17)30295-X pmid: 29268086
[150]	Wang S Y, Bunce J A. 2004. Elevated carbon dioxide affects fruit flavor in field-grown strawberries(Fragaria × ananassa Duch). Journal of the Science of Food and Agriculture, 84 (12):1464-1468. doi: 10.1002/jsfa.v84:12 URL
[151]	Wang Wenjing, Lü Sijia, Wang Qinghao, He Fan, Wu Yueyan. 2021. Research advance on the metabolism and regulation of plant floral fragrance. Molecular Plant Breeding, 19 (22):7612-7617. (in Chinese)
	王文静, 吕思佳, 汪庆昊, 何凡, 吴月燕. 2021. 植物花香物质代谢与调控研究进展. 分子植物育种, 19 (22):7612-7617.
[152]	Wang X B, Zhang C L, Miao Y L, Deng L, Zhang B, Meng J R, Wang Y, Pan L, Niu L, Liu H, Cui G C, Wang Z Q, Zeng W F. 2022. Interaction between PpERF5 and PpERF7 enhances peach fruit aroma by upregulating PpLOX4 expression. Plant Physiology and Biochemistry, 185:378-389. doi: 10.1016/j.plaphy.2022.06.024 URL
[153]	Wang Z, Yuan Y, Dong R, Zeng R, Zhao X, Xu Y, Gao J, Hong B, Gu Z. 2025. Salicylic acid regulates biosynthesis of floral fragrance (E)-β-farnesene via NPR3-WRKY 1 module in chrysanthemum. Molecular Horticulture, 5 (1):52. doi: 10.1186/s43897-025-00174-y
[154]	Wei C, Liu H, Cao X, Zhang M, Li X, Chen K, Zhang B. 2021. Synthesis of flavour-related linalool is regulated by PpbHLH1 and associated with changes in DNA methylation during peach fruit ripening. Plant Biotechnology Journal, 19 (10):2082-2096. doi: 10.1111/pbi.v19.10 URL
[155]	Wei C Y, Li M T, Cao X M, Jin Z N, Zhang C, Xu M, Chen K S, Zhang B. 2022. Linalool synthesis related PpTPS1 and PpTPS3 are activated by transcription factor PpERF61 whose expression is associated with DNA methylation during peach fruit ripening. Plant Science, 317:111200.
[156]	Wei Y, Meng N, Wang Y, Cheng J, Duan C, Pan Q. 2023. Transcription factor VvWRKY70 inhibits both norisoprenoid and flavonol biosynthesis in grape. Plant Physiology, 193 (3):2055-2070. doi: 10.1093/plphys/kiad423 pmid: 37471439
[157]	Wen Dian. 2022. Effects of high temperature and Spodoptera litura on terpenoid biosynthesis in Chrysanthemum nankingense[M. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	文典. 2022. 高温、斜纹夜蛾侵害对菊花脑萜类物质生物合成影响的研究[硕士论文]. 南京: 南京农业大学.
[158]	Wendt L M, Ludwig V, Thewes F R, Soldateli F J, Batista C B, Fukui C M, dos Santos G G, Katsurayama J M, Brackmann A, Both V. 2024. Effect of dynamic controlled atmosphere on volatile compound profile and quality of pears. Scientia Horticulturae, 328:112910.
[159]	Wu Miao. 2017. Characterization of OfWRKY3,a transcription factor that positively regulates the carotenoid cleavage dioxygenase gene OfCCD4in Osmanthus fragrans[M. D. Dissertation]. Kaifeng: Henan University. (in Chinese)
	吴淼. 2017. 桂花WRKY3转录因子调控CCD4基因功能的验证[硕士论文]. 开封: 河南大学.
[160]	Wu Q, Tao X Y, Ai X Z, Luo Z S, Mao L C, Ying T J, Li L. 2018. Contribution of abscisic acid to aromatic volatiles in cherry tomato(Solanum lycopersicum L.) fruit during postharvest ripening. Plant Physiology and Biochemistry, 130:205-214. doi: 10.1016/j.plaphy.2018.06.039 URL
[161]	Wu Qi, Fu Yuchen, Leng Pingsheng, Hu Zenghui. 2017. Effects of different treatment time of methyl jasmonate(MeJA)on the floral scent of Lilium‘Siberia’. Beijing: China Forestry Publishing House:315-320. (in Chinese)
	吴琦, 付宇辰, 冷平生, 胡增辉. 2017. 茉莉酸甲酯(MeJA)处理时长对‘西伯利亚’百合花香的影响. 北京: 中国林业出版社:315-320.
[162]	Wu Qi, Fu Yuchen, Yan Zifei, Wang Shaojie, Leng Pingsheng, Hu Zenghui. 2018. Effect of foliar application of methyl jasmonate on the floral scent of lily. Jiangsu Agricultural Sciences, 46 (6):100-104. (in Chinese)
	吴琦, 付宇辰, 闫子飞, 王少杰, 冷平生, 胡增辉. 2018. 喷施茉莉酸甲酯对百合花香的影响. 江苏农业科学, 46 (6):100-104.
[163]	Xi W, Jiang M Y, Zhu L L, Zeng X M, Ju H, Yang Q L, Zhang T Y, Wang C Y, Zheng R R. 2025. OfWRKY33 binds to the promoter of key linalool synthase gene OfTPS7 to stimulate linalool synthesis in Osmanthus fragrans flowers. Horticulture Research, 12 (9):uhaf155.
[164]	Xu J, van Herwijnen Z O, Dräger D B, Sui C, Haring M A, Schuurink R C. 2018. SlMYC1 regulates type VI glandular trichome formation and terpene biosynthesis in tomato glandular cells. Plant Cell, 30 (12):2988-3005. doi: 10.1105/tpc.18.00571 URL
[165]	Xu Shanbin, Zheng Hongliang, Liu Lifeng, Bu Qingyun, Li Xiufeng, Zou Detang. 2022. Efficient creation of long-grain fragrant rice using CRISPR/Cas 9 technology. Chinese Journal of Rice Science, 34 (5):406-412. (in Chinese)
	徐善斌, 郑洪亮, 刘利锋, 卜庆云, 李秀峰, 邹德堂. 2022. 利用CRISPR/Cas9技术高效创制长粒香型水稻. 中国水稻科学, 34 (5):406-412.
[166]	Xu X, Zhu X, Jiang F, Li Q, Zhang A, Zhang H, Li J. 2024. Mechanism of abscisic acid in promoting softening of postharvest‘Docteur Jules Guyot’pear(Pyrus communis L.). Frontiers in Plant Science, 15:1502623.
[167]	Xu Y H, Wang J W, Wang S, Wang J Y, Chen X Y. 2004. Characterization of GaWRKY1,a cotton transcription factor that regulates the sesquiterpene synthase gene (+)-delta-cadinene synthase-A. Plant Physiology, 135 (2):507-515. doi: 10.1104/pp.104.038612 URL
[168]	Xue W Y, Liu N, Lu P F, Yang Y Y, Chen S X. 2024. Photosynthesis and fatty acid metabolism reveals the effects of shading treatment on the fruit aroma quality of cucumber(Cucumis sativus L.). Scientia Horticulturae, 337:113508.
[169]	Yan X, Ding W, Wu X, Wang L, Yang X, Yue Y. 2022. Insights into the MYB-related transcription factors involved in regulating floral aroma synthesis in sweet osmanthus. Frontiers in Plant Science, 13:765213.
[170]	Yan X M, Zhang L, Wang J, Liao P, Zhang Y, Zhang R, Kai G Y. 2009. Molecular characterization and expression of 1-deoxy-d-xylulose 5-phosphate reductoisomerase(DXR)gene from Salvia miltiorrhiza. Acta Physiologiae Plantarum, 31 (5):1015-1022. doi: 10.1007/s11738-009-0320-5 URL
[171]	Yang J H, Chen Y C, Gao M, Zhao Y X, Wang Y D. 2025. LcMYB46,a R2R3-MYB transcription factor,promotes the accumulation of citral by activating TPS-CYP cascade in Litsea cubeba. Journal of Agricultural and Food Chemistry, 73 (25):15796-15808. doi: 10.1021/acs.jafc.5c00678 URL
[172]	Yang Y Y, Ma B, Li Y Y, Han M Z, Wu J, Zhou X F, Tian J, Wang W H, Leng P S, Hu Z H. 2022a. Transcriptome analysis identifies key gene LiMYB305 involved in monoterpene biosynthesis in Lilium‘Siberia’. Frontiers in Plant Science, 13:1021576.

转录因子家族 Transcription factor family		基因 Gene		功能描述 Functional description	物种 Species		参考文献 Reference
MYB		AtMYB21/24		调控苯丙素合成相关基因，促进芳香物质产生 Enhancement of aromatic volatile production via the regulation of phenylpropanoid biosynthetic genes	拟南芥 Arabidopsis thaliana		Reeves et al.,2012
		OfMYB1R-70，OfMYB1R-114，OfMYB1R-201		调控苯丙烷/苯丙酮等香气物质合成，影响β-紫罗酮等挥发物含量Regulation of phenylpropanoid/phenylpropanone-derived aroma compounds biosynthesis，and influencing the emission of volatiles such as β-ionone	桂花 Osmanthus fragrans		Yan et al.,2022
		CtMYB2		同时激活CtTPS3和CtTPS8的表达 Activating the expression of CtTPS3 and CtTPS8	西藏虎头兰Cymbidium tracyanum		Tu et al.,2025
		CtMYB3		同时激活 CtTPS2、CtTPS3和CtTPS8 的表达 Activating the expression of CtTPS2，CtTPS3 and CtTPS8	西藏虎头兰Cymbidium tracyanum		Tu et al.,2025
		ODORANT1		直接调控莽草酸途径中关键酶基因表达 Directly modulates the expression of key enzymes in the shikimate pathway	矮牵牛 Petunia hybrida		Verdonk et al.,2005
		FaEOBII		通过激活肉桂醇脱氢酶（FaCAD1）和丁香酚合成酶（FaEGS2）的表达，调控挥发性丁香酚的产生 Regulating volatile eugenol production by activating the expression of cinnamyl-alcohol dehydrogenase（FaCAD1）and eugenol synthase（FaEGS2）	草莓 Fragaria × ananassa		Medina-Puche et al.,2015
		FhMYB21L1		直接结合FhTPS1启动子中的MYBCORE位点，激活FhTPS1的表达，促进芳樟醇的释放 Directly binds to the MYBCORE cis-element in the FhTPS1 promoter，thereby activating FhTPS1 expression and promoting linalool emission	香雪兰 Freesia hybrida		张佳等,2024
		FhMYB21L2			香雪兰 Freesia hybrida		张佳等,2024
		PhMYB4		作为苯丙烷类途径中肉桂酸-4-羟基酶的阻滞剂，控制矮牵牛花中由对香豆酸衍生的挥发物的碳通量，如异丁香酚和丁香酚 Acting as a repressor of cinnamate-4-hydroxylase in the phenylpropanoid pathway，and controlling the carbon flux toward p-coumaric acid-derived volatiles，such as isoeugenol and eugenol in petunia flowers	矮牵牛 Petunia hybrida		Colquhoun et al.,2011
		CrBPF1		增加吲哚和萜类生物合成途径中基因的转录水平 Elevates the transcript levels of genes in the indole and terpenoid biosynthetic pathways	长春花 Catharanthus roseus		Li et al.,2015
		AtMYB21		激活倍半萜合酶基因的表达 Activating the expression of sesquiterpene synthase genes	拟南芥 Arabidopsis thaliana		Yang et al.,2020
		PAP1		增加挥发性苯丙素/苯类化合物的含量 Increases the abundance of volatile phenylpropanoid/benzenoid compounds	拟南芥 Arabidopsis thaliana		Ben et al.,2008
				增加萜类化合物中气味化合物水平 Elevates the levels of odor-active terpenoid compounds	月季 Rosa chinensis		Zvi et al.,2012
		MYB1		与代谢物通量、产生黄酮醇/花青素的苯丙烷途径有关 Related to metabolic flux through the phenylpropanoid pathway that produces flavonols and anthocyanins.	蕙兰 Cymbidium faberi		Ramya et al.,2019
		FaEOBII		调节挥发性苯丙烷通路基因表达 Regulates the expression of genes within the volatile phenylpropanoid pathway	草莓 Fragaria × ananassa		Medina-Puche et al.,2015
		FaMYB10		调节类黄酮/苯丙烷途径的早期和晚期生物合成基因 Modulates both early- and late-biosynthetic genes in the flavonoid/phenylpropanoid pathway	草莓 Fragaria × ananassa		Medina-Puche et al.,2015
		AtMYB21		激活倍半萜合酶基因的表达 Activates the expression of sesquiterpene synthase genes	拟南芥 Arabidopsis thaliana		Yang et al.,2020
MYB	HcMYB7，HcMYB8，HcMYB79，HcMYB145，HcMYB248		调控花香生物合成基因的表达 Regulates the expression of floral scent biosynthetic genes			姜花Hedychium coronarium		Abbas et al.,2021b
	MdMYB85		MdMYC2和MdMYB85与MdAAT1的启动子区域相互作用，从而增强其转录活性，促进苹果酯香气合成 MdMYC2 and MdMYB85 interact with the promoter region of MdAAT1 to enhance its transcriptional activity，thereby promoting ester-derived aroma biosynthesis in apple			苹果 Malus × domestica		Li et al.,2023
	MsMYB		单萜生物合成的新型负调节因子 A novel negative regulator of monoterpene biosynthesis			留兰香 Mentha spicata		Reddy et al.,2017
	CCA1		上调萜烯合酶表达，影响蝴蝶兰花香的昼夜节律发射 Upregulates the expression of terpene biosynthsis genes，and influences the diurnal emission of floral volatiles in Phalaenopsis			蝴蝶兰 Phalaenopsis aphrodite		Yeh et al.,2022
	OfMYB21		正向调控芳樟醇生物合成 Positively regulates linalool biosynthesis			桂花 Osmanthus fragrans		Lan et al.,2023
	ODO1		调控莽草酸途径前体的合成 Regulates the precursor biosynthesis in shikimate pathway			矮牵牛 Petunia hybrida		van Moerkercke et al.,2011
	PhMYB4		抑制C4H转录，间接控制矮牵牛花组织中挥发性苯类/苯丙烷类化合物产生的平衡Repression of C4H transcription，indirectly controls the balance of volatile benzenoid/phenylpropanoid production in petunia tissues			矮牵牛 Petunia hybrida		Colquhoun et al.,2011
	PH4		PH4基因的沉默导致花苯丙醇挥发性物质排放显着减少 Silencing of PH4 markedly reduces floral phenylpropanoid- alcohol volatile emission			矮牵牛 Petunia hybrida		Cna'ani et al.,2015
	SlMYB75		调节倍半萜的积累 Regulates sesquiterpene accumulation			番茄 Solanum lycopersicum		Gong et al.,2021
	TcMYB8		调节萜类化合物的生物合成 Modulates terpenoid biosynthesis			除虫菊Tanacetum cinerariifolium		Zhou et al.,2022
	VviMYB24		与TPS和HYH的启动子结合，调节单萜和黄酮醇的生物合成 Binds to the promoter regions of both TPS genes and HYH to modulate monoterpene and flavonol biosynthesis			欧亚种葡萄 Vitis vinifera		Zhang et al.,2023
	AmMYB24		调控蓝光诱导的萜类物质合成 Regulates blue-light-induced terpenoid biosynthesis			金鱼草 Antirrhinum majus		Han et al.,2022
	PbbHLH4		激活单萜合成基因GDPS Activates the monoterpene-biosynthetic gene GDPS			蝴蝶兰 Phalaenopsis aphrodite		Chuang et al.,2018
	DobHLH4		芳樟醇合成的正向调控因子，促进萜类物质积累 A positive regulator of linalool synthesis that promotes terpenoid accumulation			铁皮石斛Dendrobium officinale		Yu et al.,2021
	CfbHLH		直接结合CfAOC和CfJMT启动子，促进茉莉酸甲酯合成 Directly binds to the CfAOC and CfJMT promoters to promote methyl jasmonate biosynthesis			蕙兰 Cymbidium faberi		Zhou et al.,2023
bHLH	AaMYC2		激活CYP71AV1和DBR2的转录，导致青蒿素含量增加 Activates the transcription of CYP71AV1 and DBR2，increasing accumulation of artemisinin			青蒿 Artemisia annua		Shen et al.,2016
	AabHLH1		正向调控青蒿素（倍半萜内酯）的生物合成 Positively regulates artemisinin（sesquiterpene lactone）biosynthesis			青蒿 Artemisia annua		Ji et al.,2014
	AtMYC2		参与GA介导的倍半萜合酶基因的诱导 Participates in gibberellin-mediated induction of sesquiterpene synthase genes expression			拟南芥 Arabidopsis thaliana		Hong et al.,2012
	CrMYC2		调节萜类物质生物合成Regulates terpenoid biosynthesis			长春花Catharanthus roseus		Zhang et al.,2011
	BIS1		调控单萜物质生物合成Regulates monoterpene biosynthesis			长春花Catharanthus roseus		van Moerkercke et al.,2015,2016
	BIS2		调控萜类物质合成Regulates terpenoid biosynthesis			长春花Catharanthus roseus		van Moerkercke et al.,2015,2016
	MtTSAR1，MtTSAR2		提高单萜吲哚生物碱（MIA）通路环烯醚萜分支的基因的表达 Enhances the expression of genes within the iridoid branch of the monoterpene indole alkaloid（MIA） pathway			长春花Catharanthus roseus		Mertens et al.,2016
	CpMYC2，CpbHLH13		参与单萜（芳樟醇）和倍半萜（β-石竹烯）的生物合成 Participates in the biosynthesis of monoterpenes（linalool）and sesquiterpenes（β-caryophyllene）			腊梅 Chimonanthus praecox		Aslam et al.,2020
bHLH	MYC2		花香产生的候选基因Candidate genes for floral scent production			百合属Lilium spp.		Shi et al.,2018
	LaMYC4		LaMYC4调节挥发性萜类生物合成 LaMYC4 regulates volatile terpenoid biosynthesis			薰衣草 Lavandula angustifolia		Dong et al.,2022
	MdMYC2		调节苹果果实中的α-法呢烯生物合成 Regulates α-farnesene biosynthesis in apple fruit			苹果 Malus × domestica		Wang et al.,2020
	PbbHLH4		调控单萜生物合成Regulates monoterpene biosynthesis			蝴蝶兰 Phalaenopsis aphrodite		Chuang et al.,2018
	PpbHLH1		提高芳樟醇的产量Enhances linalool yield			桃 Prunus persica		Wei et al.,2021
	SlMYC1		调节萜烯生物合成Regulates terpene biosynthesis			番茄Solanum lycopersicum		Xu et al.,2018
	JIG1		MYC2-SlJIG模块在萜烯生物合成发挥作用 The MYC2-SlJIG module functions in terpene biosynthesis			番茄 Solanum lycopersicum		Cao et al.,2022
	TgbHLH95		TgbHLH95-TgbZIP44复合物结合TgGPPS启动子，上调香榧果实收获后萜烯生物合成基因 The TgbHLH95-TgbZIP44 complex binds the TgGPPS promoter and upregulates post-harvest terpene biosynthesis genes in Torreya grandis nuts			香榧 Torreya grandis		Zhang et al.,2023
	OfWRKY3		直接结合花瓣香气关键酶OfCCD4启动子的W-box序列，有效调控该酶的转录活性 Directly binds the W-box element in the promoter of the petal-aroma key enzyme OfCCD4 and effectively modulates its transcriptional activity			桂花 Osmanthus fragrans		吴淼,2017
	OfWRKY33		诱导MEP途径基因OfDXS1表达，促进芳樟醇的生物合成 Induces expression of OfDXS1 and promotes linalool biosynthesis			桂花 Osmanthus fragrans		Xi et al.,2025
	OfWRKY21		显著提升丁酸1-苯乙基酯、苯甲酰胺、苯丙酸甲酯等挥发性芳香物质的积累水平 Significantly enhances the accumulation of volatile aromatics，including 1-phenylethyl butyrate，benzamide，and methyl phenylpropanoate			桂花 Osmanthus fragrans		郭炳澳,2024
WRKY	HcWRKY1		促进β-石竹烯、柠檬烯和α-蒎烯等花朵挥发性物质的生物合成 Promotes the biosynthesis of floral volatiles such as β-caryophyllene，limonene，and α-pinene			姜花 Hedychium coronarium		高婷,2025
	AvWRKY61，AvWRKY28，AvWRKY40		参与萜类物质合成Participates in terpenoid biosynthesis			砂仁Amomum villosum		He et al.,2018
	AtWRKY40		激活MEP途径中多个基因的转录 Activates the transcription of multiple genes in the MEP pathway			欧丹参Salvia sclarea		Alfieri et al.,2018
	CmWRKY41		激活 CmHMGR2 和 CmFPPS2，正向调控菊花中倍半萜的合成 Activates the expression of CmHMGR2 and CmFPPS2 to positively regulate sesquiterpene biosynthesis in Chrysanthemum			杭白菊Chrysanthemum morifolium		Hu et al.,2023
	NaWRKY3，NaWRKY6		激活茉莉酸（JA）的生物合成，提高烟草中的挥发性物质含量 Activation of jasmonic acid（JA）biosynthesis to increase the volatile content in tobacco			烟草 Tobacco		Skibbe et al.,2008
	WRKY17		促进单萜合成Promotes monoterpene biosynthesis			山胡椒Lindera glauca		Gao et al.,2023
	OfWRKY7，OfWRKY19，OfWRKY36		与单萜类物质合成有关 Involved in monoterpene biosynthesis			桂花 Osmanthus fragrans		Ding et al.,2020
	WRKY70		调控单萜类化合物香叶醇的生物合成 Regulates the biosynthesis of the monoterpenoid geraniol			月季 Rosa chinensis		Yu et al.,2022
	SmWRKY1		参与丹参酮生物合成的调控，激活MEP通路中的SmDXR表达Participates in the regulation of tanshinone biosynthesis by activating SmDXR in the MEP pathway			丹参 Salvia miltiorrhiza		Cao et al.,2018
	AaGSW1		特异性WRKY转录因子，是青蒿素生物合成途径中的正调节因子 A specific WRKY transcription factor that functions as a positive regulator of artemisinin biosynthesis			青蒿 Artemisia annua		Chen et al.,2017
	AaWRKY9		直接与AaDBR2和AaGSW1的启动子结合，正向调节青蒿素生物合成 Directly binds to the promoters of AaDBR2 and AaGSW1 to positively regulate artemisinin biosynthesis			青蒿Artemisia annua		Fu et al.,2021
	AaWRKY40		参与萜类代谢 Involved in terpenoid metabolism			青蒿 Artemisia annua		De Paolis et al.,2020
WRKY	CiAP2.10		促进倍半萜（+）-valencene的合成 Promotes the biosynthesis of the sesquiterpene（+）-valencene			橘Citrus reticulata Blanco		Shen et al.,2016
	CitERF71		控制柑橘类水果中E-香叶醇的产生 Controls the production of E-geraniol in Citrus fruit			甜橙Citrus sinensis		Li et al.,2017
	MdERF3		调节苹果果实中的α-法呢烯生物合成 Modulates α-farnesene biosynthesis in apple fruits			苹果Malus pumila		Wang et al.,2020
AP2/ERF	PpERF61		提高 PpTPS1和PpTPS3 的表达和芳香醇含量 Enhances the expression of PpTPS1 and PpTPS3 and the content of aromatic alcohols			桃Prunus persica		Wei et al.,2022
	PhERF6		通过竞争EOBI蛋白的c-myb 结构域与花香相关基因的启动子的结合，负向调节矮牵牛花的挥发性物质产生 Negatively regulates the production of floral volatiles in petunia by competing with the EOBI protein for binding to the c-myb domain in the promoters of scent-related genes			矮牵牛 Petunia hybrida		Liu et al.,2017
	PpERF5，PpERF7		上调PpLOX4表达从而促进香气的生物合成 Upregulates PpLOX4 expression and thereby promotes aroma biosynthesis			桃Prunus persica		Wang et al.,2022

转录因子家族 Transcription factor family		基因 Gene		功能描述 Functional description	物种 Species		参考文献 Reference
MYB		AtMYB21/24		调控苯丙素合成相关基因，促进芳香物质产生 Enhancement of aromatic volatile production via the regulation of phenylpropanoid biosynthetic genes	拟南芥 Arabidopsis thaliana		Reeves et al.,2012
		OfMYB1R-70，OfMYB1R-114，OfMYB1R-201		调控苯丙烷/苯丙酮等香气物质合成，影响β-紫罗酮等挥发物含量Regulation of phenylpropanoid/phenylpropanone-derived aroma compounds biosynthesis，and influencing the emission of volatiles such as β-ionone	桂花 Osmanthus fragrans		Yan et al.,2022
		CtMYB2		同时激活CtTPS3和CtTPS8的表达 Activating the expression of CtTPS3 and CtTPS8	西藏虎头兰Cymbidium tracyanum		Tu et al.,2025
		CtMYB3		同时激活 CtTPS2、CtTPS3和CtTPS8 的表达 Activating the expression of CtTPS2，CtTPS3 and CtTPS8	西藏虎头兰Cymbidium tracyanum		Tu et al.,2025
		ODORANT1		直接调控莽草酸途径中关键酶基因表达 Directly modulates the expression of key enzymes in the shikimate pathway	矮牵牛 Petunia hybrida		Verdonk et al.,2005
		FaEOBII		通过激活肉桂醇脱氢酶（FaCAD1）和丁香酚合成酶（FaEGS2）的表达，调控挥发性丁香酚的产生 Regulating volatile eugenol production by activating the expression of cinnamyl-alcohol dehydrogenase（FaCAD1）and eugenol synthase（FaEGS2）	草莓 Fragaria × ananassa		Medina-Puche et al.,2015
		FhMYB21L1		直接结合FhTPS1启动子中的MYBCORE位点，激活FhTPS1的表达，促进芳樟醇的释放 Directly binds to the MYBCORE cis-element in the FhTPS1 promoter，thereby activating FhTPS1 expression and promoting linalool emission	香雪兰 Freesia hybrida		张佳等,2024
		FhMYB21L2			香雪兰 Freesia hybrida		张佳等,2024
		PhMYB4		作为苯丙烷类途径中肉桂酸-4-羟基酶的阻滞剂，控制矮牵牛花中由对香豆酸衍生的挥发物的碳通量，如异丁香酚和丁香酚 Acting as a repressor of cinnamate-4-hydroxylase in the phenylpropanoid pathway，and controlling the carbon flux toward p-coumaric acid-derived volatiles，such as isoeugenol and eugenol in petunia flowers	矮牵牛 Petunia hybrida		Colquhoun et al.,2011
		CrBPF1		增加吲哚和萜类生物合成途径中基因的转录水平 Elevates the transcript levels of genes in the indole and terpenoid biosynthetic pathways	长春花 Catharanthus roseus		Li et al.,2015
		AtMYB21		激活倍半萜合酶基因的表达 Activating the expression of sesquiterpene synthase genes	拟南芥 Arabidopsis thaliana		Yang et al.,2020
		PAP1		增加挥发性苯丙素/苯类化合物的含量 Increases the abundance of volatile phenylpropanoid/benzenoid compounds	拟南芥 Arabidopsis thaliana		Ben et al.,2008
				增加萜类化合物中气味化合物水平 Elevates the levels of odor-active terpenoid compounds	月季 Rosa chinensis		Zvi et al.,2012
		MYB1		与代谢物通量、产生黄酮醇/花青素的苯丙烷途径有关 Related to metabolic flux through the phenylpropanoid pathway that produces flavonols and anthocyanins.	蕙兰 Cymbidium faberi		Ramya et al.,2019
		FaEOBII		调节挥发性苯丙烷通路基因表达 Regulates the expression of genes within the volatile phenylpropanoid pathway	草莓 Fragaria × ananassa		Medina-Puche et al.,2015
		FaMYB10		调节类黄酮/苯丙烷途径的早期和晚期生物合成基因 Modulates both early- and late-biosynthetic genes in the flavonoid/phenylpropanoid pathway	草莓 Fragaria × ananassa		Medina-Puche et al.,2015
		AtMYB21		激活倍半萜合酶基因的表达 Activates the expression of sesquiterpene synthase genes	拟南芥 Arabidopsis thaliana		Yang et al.,2020
MYB	HcMYB7，HcMYB8，HcMYB79，HcMYB145，HcMYB248		调控花香生物合成基因的表达 Regulates the expression of floral scent biosynthetic genes			姜花Hedychium coronarium		Abbas et al.,2021b
	MdMYB85		MdMYC2和MdMYB85与MdAAT1的启动子区域相互作用，从而增强其转录活性，促进苹果酯香气合成 MdMYC2 and MdMYB85 interact with the promoter region of MdAAT1 to enhance its transcriptional activity，thereby promoting ester-derived aroma biosynthesis in apple			苹果 Malus × domestica		Li et al.,2023
	MsMYB		单萜生物合成的新型负调节因子 A novel negative regulator of monoterpene biosynthesis			留兰香 Mentha spicata		Reddy et al.,2017
	CCA1		上调萜烯合酶表达，影响蝴蝶兰花香的昼夜节律发射 Upregulates the expression of terpene biosynthsis genes，and influences the diurnal emission of floral volatiles in Phalaenopsis			蝴蝶兰 Phalaenopsis aphrodite		Yeh et al.,2022
	OfMYB21		正向调控芳樟醇生物合成 Positively regulates linalool biosynthesis			桂花 Osmanthus fragrans		Lan et al.,2023
	ODO1		调控莽草酸途径前体的合成 Regulates the precursor biosynthesis in shikimate pathway			矮牵牛 Petunia hybrida		van Moerkercke et al.,2011
	PhMYB4		抑制C4H转录，间接控制矮牵牛花组织中挥发性苯类/苯丙烷类化合物产生的平衡Repression of C4H transcription，indirectly controls the balance of volatile benzenoid/phenylpropanoid production in petunia tissues			矮牵牛 Petunia hybrida		Colquhoun et al.,2011
	PH4		PH4基因的沉默导致花苯丙醇挥发性物质排放显着减少 Silencing of PH4 markedly reduces floral phenylpropanoid- alcohol volatile emission			矮牵牛 Petunia hybrida		Cna'ani et al.,2015
	SlMYB75		调节倍半萜的积累 Regulates sesquiterpene accumulation			番茄 Solanum lycopersicum		Gong et al.,2021
	TcMYB8		调节萜类化合物的生物合成 Modulates terpenoid biosynthesis			除虫菊Tanacetum cinerariifolium		Zhou et al.,2022
	VviMYB24		与TPS和HYH的启动子结合，调节单萜和黄酮醇的生物合成 Binds to the promoter regions of both TPS genes and HYH to modulate monoterpene and flavonol biosynthesis			欧亚种葡萄 Vitis vinifera		Zhang et al.,2023
	AmMYB24		调控蓝光诱导的萜类物质合成 Regulates blue-light-induced terpenoid biosynthesis			金鱼草 Antirrhinum majus		Han et al.,2022
	PbbHLH4		激活单萜合成基因GDPS Activates the monoterpene-biosynthetic gene GDPS			蝴蝶兰 Phalaenopsis aphrodite		Chuang et al.,2018
	DobHLH4		芳樟醇合成的正向调控因子，促进萜类物质积累 A positive regulator of linalool synthesis that promotes terpenoid accumulation			铁皮石斛Dendrobium officinale		Yu et al.,2021
	CfbHLH		直接结合CfAOC和CfJMT启动子，促进茉莉酸甲酯合成 Directly binds to the CfAOC and CfJMT promoters to promote methyl jasmonate biosynthesis			蕙兰 Cymbidium faberi		Zhou et al.,2023
bHLH	AaMYC2		激活CYP71AV1和DBR2的转录，导致青蒿素含量增加 Activates the transcription of CYP71AV1 and DBR2，increasing accumulation of artemisinin			青蒿 Artemisia annua		Shen et al.,2016
	AabHLH1		正向调控青蒿素（倍半萜内酯）的生物合成 Positively regulates artemisinin（sesquiterpene lactone）biosynthesis			青蒿 Artemisia annua		Ji et al.,2014
	AtMYC2		参与GA介导的倍半萜合酶基因的诱导 Participates in gibberellin-mediated induction of sesquiterpene synthase genes expression			拟南芥 Arabidopsis thaliana		Hong et al.,2012
	CrMYC2		调节萜类物质生物合成Regulates terpenoid biosynthesis			长春花Catharanthus roseus		Zhang et al.,2011
	BIS1		调控单萜物质生物合成Regulates monoterpene biosynthesis			长春花Catharanthus roseus		van Moerkercke et al.,2015,2016
	BIS2		调控萜类物质合成Regulates terpenoid biosynthesis			长春花Catharanthus roseus		van Moerkercke et al.,2015,2016
	MtTSAR1，MtTSAR2		提高单萜吲哚生物碱（MIA）通路环烯醚萜分支的基因的表达 Enhances the expression of genes within the iridoid branch of the monoterpene indole alkaloid（MIA） pathway			长春花Catharanthus roseus		Mertens et al.,2016
	CpMYC2，CpbHLH13		参与单萜（芳樟醇）和倍半萜（β-石竹烯）的生物合成 Participates in the biosynthesis of monoterpenes（linalool）and sesquiterpenes（β-caryophyllene）			腊梅 Chimonanthus praecox		Aslam et al.,2020
bHLH	MYC2		花香产生的候选基因Candidate genes for floral scent production			百合属Lilium spp.		Shi et al.,2018
	LaMYC4		LaMYC4调节挥发性萜类生物合成 LaMYC4 regulates volatile terpenoid biosynthesis			薰衣草 Lavandula angustifolia		Dong et al.,2022
	MdMYC2		调节苹果果实中的α-法呢烯生物合成 Regulates α-farnesene biosynthesis in apple fruit			苹果 Malus × domestica		Wang et al.,2020
	PbbHLH4		调控单萜生物合成Regulates monoterpene biosynthesis			蝴蝶兰 Phalaenopsis aphrodite		Chuang et al.,2018
	PpbHLH1		提高芳樟醇的产量Enhances linalool yield			桃 Prunus persica		Wei et al.,2021
	SlMYC1		调节萜烯生物合成Regulates terpene biosynthesis			番茄Solanum lycopersicum		Xu et al.,2018
	JIG1		MYC2-SlJIG模块在萜烯生物合成发挥作用 The MYC2-SlJIG module functions in terpene biosynthesis			番茄 Solanum lycopersicum		Cao et al.,2022
	TgbHLH95		TgbHLH95-TgbZIP44复合物结合TgGPPS启动子，上调香榧果实收获后萜烯生物合成基因 The TgbHLH95-TgbZIP44 complex binds the TgGPPS promoter and upregulates post-harvest terpene biosynthesis genes in Torreya grandis nuts			香榧 Torreya grandis		Zhang et al.,2023
	OfWRKY3		直接结合花瓣香气关键酶OfCCD4启动子的W-box序列，有效调控该酶的转录活性 Directly binds the W-box element in the promoter of the petal-aroma key enzyme OfCCD4 and effectively modulates its transcriptional activity			桂花 Osmanthus fragrans		吴淼,2017
	OfWRKY33		诱导MEP途径基因OfDXS1表达，促进芳樟醇的生物合成 Induces expression of OfDXS1 and promotes linalool biosynthesis			桂花 Osmanthus fragrans		Xi et al.,2025
	OfWRKY21		显著提升丁酸1-苯乙基酯、苯甲酰胺、苯丙酸甲酯等挥发性芳香物质的积累水平 Significantly enhances the accumulation of volatile aromatics，including 1-phenylethyl butyrate，benzamide，and methyl phenylpropanoate			桂花 Osmanthus fragrans		郭炳澳,2024
WRKY	HcWRKY1		促进β-石竹烯、柠檬烯和α-蒎烯等花朵挥发性物质的生物合成 Promotes the biosynthesis of floral volatiles such as β-caryophyllene，limonene，and α-pinene			姜花 Hedychium coronarium		高婷,2025
	AvWRKY61，AvWRKY28，AvWRKY40		参与萜类物质合成Participates in terpenoid biosynthesis			砂仁Amomum villosum		He et al.,2018
	AtWRKY40		激活MEP途径中多个基因的转录 Activates the transcription of multiple genes in the MEP pathway			欧丹参Salvia sclarea		Alfieri et al.,2018
	CmWRKY41		激活 CmHMGR2 和 CmFPPS2，正向调控菊花中倍半萜的合成 Activates the expression of CmHMGR2 and CmFPPS2 to positively regulate sesquiterpene biosynthesis in Chrysanthemum			杭白菊Chrysanthemum morifolium		Hu et al.,2023
	NaWRKY3，NaWRKY6		激活茉莉酸（JA）的生物合成，提高烟草中的挥发性物质含量 Activation of jasmonic acid（JA）biosynthesis to increase the volatile content in tobacco			烟草 Tobacco		Skibbe et al.,2008
	WRKY17		促进单萜合成Promotes monoterpene biosynthesis			山胡椒Lindera glauca		Gao et al.,2023
	OfWRKY7，OfWRKY19，OfWRKY36		与单萜类物质合成有关 Involved in monoterpene biosynthesis			桂花 Osmanthus fragrans		Ding et al.,2020
	WRKY70		调控单萜类化合物香叶醇的生物合成 Regulates the biosynthesis of the monoterpenoid geraniol			月季 Rosa chinensis		Yu et al.,2022
	SmWRKY1		参与丹参酮生物合成的调控，激活MEP通路中的SmDXR表达Participates in the regulation of tanshinone biosynthesis by activating SmDXR in the MEP pathway			丹参 Salvia miltiorrhiza		Cao et al.,2018
	AaGSW1		特异性WRKY转录因子，是青蒿素生物合成途径中的正调节因子 A specific WRKY transcription factor that functions as a positive regulator of artemisinin biosynthesis			青蒿 Artemisia annua		Chen et al.,2017
	AaWRKY9		直接与AaDBR2和AaGSW1的启动子结合，正向调节青蒿素生物合成 Directly binds to the promoters of AaDBR2 and AaGSW1 to positively regulate artemisinin biosynthesis			青蒿Artemisia annua		Fu et al.,2021
	AaWRKY40		参与萜类代谢 Involved in terpenoid metabolism			青蒿 Artemisia annua		De Paolis et al.,2020
WRKY	CiAP2.10		促进倍半萜（+）-valencene的合成 Promotes the biosynthesis of the sesquiterpene（+）-valencene			橘Citrus reticulata Blanco		Shen et al.,2016
	CitERF71		控制柑橘类水果中E-香叶醇的产生 Controls the production of E-geraniol in Citrus fruit			甜橙Citrus sinensis		Li et al.,2017
	MdERF3		调节苹果果实中的α-法呢烯生物合成 Modulates α-farnesene biosynthesis in apple fruits			苹果Malus pumila		Wang et al.,2020
AP2/ERF	PpERF61		提高 PpTPS1和PpTPS3 的表达和芳香醇含量 Enhances the expression of PpTPS1 and PpTPS3 and the content of aromatic alcohols			桃Prunus persica		Wei et al.,2022
	PhERF6		通过竞争EOBI蛋白的c-myb 结构域与花香相关基因的启动子的结合，负向调节矮牵牛花的挥发性物质产生 Negatively regulates the production of floral volatiles in petunia by competing with the EOBI protein for binding to the c-myb domain in the promoters of scent-related genes			矮牵牛 Petunia hybrida		Liu et al.,2017
	PpERF5，PpERF7		上调PpLOX4表达从而促进香气的生物合成 Upregulates PpLOX4 expression and thereby promotes aroma biosynthesis			桃Prunus persica		Wang et al.,2022

激素 Hormone	物种 Species	应用方式 Application method	植物组织 Tissue	主要调控作用 Major regulatory effect	参考文献 Reference
茉莉酸Jasmonic Acid	玫瑰 Rosa rugosa	喷施 Spraying	花蕾、叶片 Lower bud，leaf	玫瑰半开期花瓣中的芳香物质总含量均提高 The total content of aromatic compounds of the petal is significantly elevated in Rosa rugosa at the half-open stage	尚彤等,2023
	‘南果’梨 Pyrus ussuriensis‘Nanguo’	浸泡 Immersion	果实 Fruit	提高了挥发性酯和不饱和脂肪酸的含量以及醇酰基转移酶、醇脱氢酶和LOX的活性 Elevates volatile ester and unsaturated fatty-acid contents and activates the expression of alcohol acyltransferase，alcohol dehydrogenase and LOX	Luo et al.,2021
	‘黑比诺’葡萄Vitis vinifera‘Pinot Noir’	喷施 Spraying	叶片 Leaf	显著增加癸酸乙酯、乙酸异戊酯、辛酸乙酯和1-辛醇含量 Significantly increases the content of ethyl decanoate，isoamyl acetate，ethyl octanoate，and 1-octanol	高阳等,2022
茉莉酸Jasmonic Acid	越橘 Vaccinium vitis-idaea	喷施 Spraying	果实 Fruit	提高果实的短链脂肪族衍生物含量，丰富果实香气层次 Enhances the content of short-chain aliphatic derivatives in fruit，thereby enriching the aroma profile	刘梦溪等,2021
	红山茶Camellia japonica var. rubra	喷施 Spraying	叶片 Leaf	诱导茶树释放挥发物引诱害虫天敌，提高茶树防御能力 Induces the emission of volatiles to attract natural enemies of pests in tea plants，thereby enhancing defense capability in tea	孙晓玲等,2016
	百合 Lilium brownii	喷施 Spraying	花瓣 Petal	挥发物总释放量显著升高 Significantly increasing the total volatile emission	吴琦等,2018
	玫瑰 Rosa rugosa	喷施 Spraying	花瓣 Petal	花蕾期、半开期和盛开期的玫瑰花香气成分释放总量均明显增加 The total emission of rose floral volatiles increases markedly at the bud，half-open，and full-bloom stages	周金鑫等,2019
	牡丹‘洛阳红’ Paeonia suffruticosa ‘Luoyang Hong’	喷施 Spraying	花瓣 Petal	2,4-二叔丁基苯酚和茉莉酸甲酯含量升高 Significantly increasing the content of 2,4-di-tert-butylphenol and methyl jasmonate	牛童非等,2023
赤霉素Gibberellin	茉莉花 Jasminum sambac	喷施 Spraying	花蕾 Flower bud	芳香成分芳樟醇和法尼烯含量升高 Significantly increasing the level of the aroma compounds linalool and farnesene	赵炜淑等,2022
生长素Auxin	白姜花 Hedychium coronarium‘White’	瓶插Vase-insertion	短截花枝 Cut flowering shoot	花香挥发性物质释放量升高 Promoting the emission of floral volatile compounds	柯艳果,2019
	葡萄Vitis vinifera	浸泡Immersion	果实 Fruit	花香挥发性物质释放量升高 Promoting the emission of floral volatile compounds	Jia et al.,2017
	姜花Hedychium coronarium	瓶插 Vase-insertion	短截花枝 Cut flowering shoot	上调苯甲酸甲酯生物合成的关键酶HcBSMT2和芳樟醇生物合成的关键酶HcTPS5的表达 Up-regulates the expression of HcBSMT2，a key enzyme in methyl benzoate biosynthesis，and HcTPS，a key enzyme in linalool biosynthesis	Ke et al.,2021
水杨酸Salicylic acid	郁金香 Tulipa gesneriana	喷施 Spraying	花苞 Floral bud	花香挥发性物质释放量明显提高 The emission of floral volatiles increased markedly	陈雨馨等,2025
	菊花 Chrysanthemum morifolium	喷施 Spraying	花瓣 Petal	通过CmWRKY1-CmNPR3转录调控模块，在花发育过程中特异性激活CmEβFS的表达，从而促进(E)-β-法尼烯的生物合成 The CmWRKY1-CmNPR3 transcriptional module specifically activates CmEβFS expression during flower development under SA treatmen，thereby promoting (E)-β-farnesene biosynthesis	Wang et al.,2025
	微型月季 Rosa chinensis‘Mini’	喷施 Spraying	叶片 Leaf	提高萜类物质合成相关基因的表达水平 Elevates the expression level of genes related to terpenoid biosynthesis	王启等,2020
	欧李‘农大6号’ Prunus humilis‘Nongda 6’	喷施 Spraying	果实 Fruit	果实不同时期均显著提高了果实的挥发物含量 Significantly increases fruit volatile content at all developmental stages	张羚绢,2024
脱落酸 Abscisic acid	番茄 Solanum lycopersicum	浸泡 Immersion	果实 Fruit	提高果实中亚油酸和亚麻酸等不饱和脂肪酸的含量，增加 β-大马酮、苯甲醛、苄腈等香气物质的积累 Elevates linoleic and α-linolenic acid contents and promotes the accumulation of aroma compounds such as β-damascenone，benzaldehyde，and benzonitrile	Wu et al.,2018
脱落酸 Abscisic acid	姜花 Hedychium coronarium	瓶插 Vase-insertion	短截花枝 Cut flowering shoot	促进姜花花香物质合成 Promotes floral scent biosynthesis in Hedychium coronarium	王楚天,2021
乙烯Ethylene	桂花 Osmanthus fragrans	盘插 Tray-insertion	花序 Inflorescence	抑制芳樟醇、二氢芳樟醇等单萜类化合物释放 Suppresses the emission of monoterpenoids，such as linalool and dihydrolinalool	邹晶晶等,2017
	白姜花 Hedychium coronarium‘White’	瓶插 Tray-insertion	短截花枝 Cut flowering shoot	乙烯处理提高白姜花萜类化合物（如罗勒烯和法尼烯）的挥发量；1-MCP处理抑制萜类化合物的挥发量 Ethylene treatment increases the emission of terpenoids such as ocimene and farnesene from Hedychium coronarium flowers，whereas 1-MCP suppresses terpenoid emission	刘晓洲等,2018
	矮牵牛 Petunia hybrida	瓶插 Vase-insertion	短截花枝 Cut flowering shoot	降低苯甲酸甲酯和其他挥发性有机化合物的排放 Reduces the emission of methyl benzoate and other volatile organic compounds	Underwood et al.,2005

激素 Hormone	物种 Species	应用方式 Application method	植物组织 Tissue	主要调控作用 Major regulatory effect	参考文献 Reference
茉莉酸Jasmonic Acid	玫瑰 Rosa rugosa	喷施 Spraying	花蕾、叶片 Lower bud，leaf	玫瑰半开期花瓣中的芳香物质总含量均提高 The total content of aromatic compounds of the petal is significantly elevated in Rosa rugosa at the half-open stage	尚彤等,2023
	‘南果’梨 Pyrus ussuriensis‘Nanguo’	浸泡 Immersion	果实 Fruit	提高了挥发性酯和不饱和脂肪酸的含量以及醇酰基转移酶、醇脱氢酶和LOX的活性 Elevates volatile ester and unsaturated fatty-acid contents and activates the expression of alcohol acyltransferase，alcohol dehydrogenase and LOX	Luo et al.,2021
	‘黑比诺’葡萄Vitis vinifera‘Pinot Noir’	喷施 Spraying	叶片 Leaf	显著增加癸酸乙酯、乙酸异戊酯、辛酸乙酯和1-辛醇含量 Significantly increases the content of ethyl decanoate，isoamyl acetate，ethyl octanoate，and 1-octanol	高阳等,2022
茉莉酸Jasmonic Acid	越橘 Vaccinium vitis-idaea	喷施 Spraying	果实 Fruit	提高果实的短链脂肪族衍生物含量，丰富果实香气层次 Enhances the content of short-chain aliphatic derivatives in fruit，thereby enriching the aroma profile	刘梦溪等,2021
	红山茶Camellia japonica var. rubra	喷施 Spraying	叶片 Leaf	诱导茶树释放挥发物引诱害虫天敌，提高茶树防御能力 Induces the emission of volatiles to attract natural enemies of pests in tea plants，thereby enhancing defense capability in tea	孙晓玲等,2016
	百合 Lilium brownii	喷施 Spraying	花瓣 Petal	挥发物总释放量显著升高 Significantly increasing the total volatile emission	吴琦等,2018
	玫瑰 Rosa rugosa	喷施 Spraying	花瓣 Petal	花蕾期、半开期和盛开期的玫瑰花香气成分释放总量均明显增加 The total emission of rose floral volatiles increases markedly at the bud，half-open，and full-bloom stages	周金鑫等,2019
	牡丹‘洛阳红’ Paeonia suffruticosa ‘Luoyang Hong’	喷施 Spraying	花瓣 Petal	2,4-二叔丁基苯酚和茉莉酸甲酯含量升高 Significantly increasing the content of 2,4-di-tert-butylphenol and methyl jasmonate	牛童非等,2023
赤霉素Gibberellin	茉莉花 Jasminum sambac	喷施 Spraying	花蕾 Flower bud	芳香成分芳樟醇和法尼烯含量升高 Significantly increasing the level of the aroma compounds linalool and farnesene	赵炜淑等,2022
生长素Auxin	白姜花 Hedychium coronarium‘White’	瓶插Vase-insertion	短截花枝 Cut flowering shoot	花香挥发性物质释放量升高 Promoting the emission of floral volatile compounds	柯艳果,2019
	葡萄Vitis vinifera	浸泡Immersion	果实 Fruit	花香挥发性物质释放量升高 Promoting the emission of floral volatile compounds	Jia et al.,2017
	姜花Hedychium coronarium	瓶插 Vase-insertion	短截花枝 Cut flowering shoot	上调苯甲酸甲酯生物合成的关键酶HcBSMT2和芳樟醇生物合成的关键酶HcTPS5的表达 Up-regulates the expression of HcBSMT2，a key enzyme in methyl benzoate biosynthesis，and HcTPS，a key enzyme in linalool biosynthesis	Ke et al.,2021
水杨酸Salicylic acid	郁金香 Tulipa gesneriana	喷施 Spraying	花苞 Floral bud	花香挥发性物质释放量明显提高 The emission of floral volatiles increased markedly	陈雨馨等,2025
	菊花 Chrysanthemum morifolium	喷施 Spraying	花瓣 Petal	通过CmWRKY1-CmNPR3转录调控模块，在花发育过程中特异性激活CmEβFS的表达，从而促进(E)-β-法尼烯的生物合成 The CmWRKY1-CmNPR3 transcriptional module specifically activates CmEβFS expression during flower development under SA treatmen，thereby promoting (E)-β-farnesene biosynthesis	Wang et al.,2025
	微型月季 Rosa chinensis‘Mini’	喷施 Spraying	叶片 Leaf	提高萜类物质合成相关基因的表达水平 Elevates the expression level of genes related to terpenoid biosynthesis	王启等,2020
	欧李‘农大6号’ Prunus humilis‘Nongda 6’	喷施 Spraying	果实 Fruit	果实不同时期均显著提高了果实的挥发物含量 Significantly increases fruit volatile content at all developmental stages	张羚绢,2024
脱落酸 Abscisic acid	番茄 Solanum lycopersicum	浸泡 Immersion	果实 Fruit	提高果实中亚油酸和亚麻酸等不饱和脂肪酸的含量，增加 β-大马酮、苯甲醛、苄腈等香气物质的积累 Elevates linoleic and α-linolenic acid contents and promotes the accumulation of aroma compounds such as β-damascenone，benzaldehyde，and benzonitrile	Wu et al.,2018
脱落酸 Abscisic acid	姜花 Hedychium coronarium	瓶插 Vase-insertion	短截花枝 Cut flowering shoot	促进姜花花香物质合成 Promotes floral scent biosynthesis in Hedychium coronarium	王楚天,2021
乙烯Ethylene	桂花 Osmanthus fragrans	盘插 Tray-insertion	花序 Inflorescence	抑制芳樟醇、二氢芳樟醇等单萜类化合物释放 Suppresses the emission of monoterpenoids，such as linalool and dihydrolinalool	邹晶晶等,2017
	白姜花 Hedychium coronarium‘White’	瓶插 Tray-insertion	短截花枝 Cut flowering shoot	乙烯处理提高白姜花萜类化合物（如罗勒烯和法尼烯）的挥发量；1-MCP处理抑制萜类化合物的挥发量 Ethylene treatment increases the emission of terpenoids such as ocimene and farnesene from Hedychium coronarium flowers，whereas 1-MCP suppresses terpenoid emission	刘晓洲等,2018
	矮牵牛 Petunia hybrida	瓶插 Vase-insertion	短截花枝 Cut flowering shoot	降低苯甲酸甲酯和其他挥发性有机化合物的排放 Reduces the emission of methyl benzoate and other volatile organic compounds	Underwood et al.,2005

Research Advances in the Regulation of Aroma Biosynthesis in Horticultural Crops by Plant Hormones and Environmental Factors

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