植物激素与环境因子调控园艺作物香气合成的研究进展

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

摘要/Abstract

摘要：

园艺作物香气品质是由挥发性有机化合物构成的重要经济性状，其合成受内部调控网络与外部环境因子的协同调控。茉莉酸、乙烯、生长素等植物激素通过激活MYB和bHLH等转录因子形成复杂的调控网络，并通过交叉互作精确调控萜类和苯丙烷类等香气物质的生物合成。钙离子、过氧化氢和一氧化氮等信号分子通过介导激素信号与环境刺激参与调控香气合成基因的表达。光照和温度等环境因子在园艺作物中不仅独立调控其香气合成，还通过光受体与温度传感器等的相互作用产生协同效应。本文中对近年来植物激素、信号分子及环境因子调控园艺作物香气合成的研究进展进行了综述，旨在为植物花香理论研究和高香气园艺植物的培育提供参考依据。

关键词: 植物激素, 信号分子, 环境因子, 香气合成

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

许弘朋, 陆锦萍, 蒋晓楠, 谢伟琳, 刘至柔, 班佳熠, 薛冰冰, 叶若玉, 张可欣, 熊宇辰, 陈佳敏, 宋叶繁, 袁福榜, 王绎乔, 刘倬闻, 高俊平, 梁跃. 植物激素与环境因子调控园艺作物香气合成的研究进展[J]. 园艺学报, 2026, 53(2): 331-358.

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.

图/表 3

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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