花香的生物功能及其合成机制研究进展

doi:10.16420/j.issn.0513-353x.2025-0853

摘要/Abstract

摘要：

花香在繁殖与防御中发挥着多功能的生态与生理作用，不仅通过特异性挥发物精准吸引传粉者或抵御生物胁迫，还参与植物体内活性氧清除和信号传导网络，形成“植物—微生物—传粉者”互作网络。在分子层面，花香合成受到多层级精密调控，包括DNA甲基化、非编码RNA等表观遗传机制，转录因子家族对结构基因表达的协同调控，以及泛素化、组蛋白修饰等翻译后蛋白修饰机制，共同实现对花香合成时空动态和胁迫响应的精细控制。研究表明，花香合成与植物衰老和抗逆过程紧密关联，其释放受激素信号网络调控，并在盐胁迫、虫害等环境中表现出重要的生态适应性与防御功能。

关键词: 花香, 逆境, 分子调控

Abstract:

Floral scent serves multifaceted ecological and physiological roles in plant reproduction and defense mechanisms. It not only selectively attracts pollinators through specific volatile compounds and provides resistance against biotic stressors，but also participates in intracellular reactive oxygen species scavenging and signaling transduction，collectively forming a complex“plant-micropollinator-environment”interaction network. At the molecular level，the biosynthesis of floral scent is regulated through sophisticated hierarchical mechanisms，including epigenetic modifications such as DNA methylation and non-coding RNAs，coordinated regulation of structural genes by transcription factor families，as well as post-translational protein modifications including ubiquitination and histone modifications. These regulatory layers collectively ensure precise spatiotemporal control of floral scent production and stress-responsive emission. Research demonstrates that floral scent biosynthesis is intrinsically linked to plant senescence and stress adaptation processes，being modulated by hormonal signaling networks while exhibiting significant ecological adaptability and defensive functions under environmental challenges such as salt stress and herbivory.

Key words: floral scent, stress, molecular regulation

谢雨滢, 施婷婷, 杨秀莲, 王良桂, 岳远征. 花香的生物功能及其合成机制研究进展[J]. 园艺学报, 2026, 53(2): 386-396.

XIE Yuying, SHI Tingting, YANG Xiulian, WANG Lianggui, YUE Yuanzheng. Research Progress on the Biological Functions and Biosynthetic Mechanisms of Floral Scent[J]. Acta Horticulturae Sinica, 2026, 53(2): 386-396.

图/表 3

参考文献 43

[1]	Abbas F, Zhou Y, He J, Ke Y, Qin W, Yu R, Fan Y. 2021. Metabolite and transcriptome profiling analysis revealed that melatonin positively regulates floral scent production in Hedychium coronarium. Front Plant Sci, 12:808899.
[2]	Aratani Y, Uemura T, Hagihara T, Matsui K, Toyota M. 2023. Green leaf volatile sensory calcium transduction in Arabidopsis. Nature Communications, 14 (1):6236. doi: 10.1038/s41467-023-41589-9 pmid: 37848440
[3]	Bao M, Xu Y, Wei G, Bai M, Wang J, Feng L. 2025. The MYC gene RrbHLH105contributes to salt stress-induced geraniol in rose by regulating trehalose-6-phosphate signalling. Plant,Cell & Environment, 48 (3):1947-1962. doi: 10.1111/pce.v48.3 URL
[4]	Beale M H, Birkett M A, Bruce T J A, Chamberlain K, Field L M, Huttly A K, Martin J L, Parker R, Phillips A L, Pickett J A, Prosser I M, Shewry P R, Smart L E, Wadhams L J, Woodcock C M, Zhang Y. 2006. Aphid alarm pheromone produced by transgenic plants affects aphid and parasitoid behavior. Proc Natl Acad Sci U S A, 103 (27):10509-10513. doi: 10.1073/pnas.0603998103 URL
[5]	Boersma M R, Patrick R M, Jillings S L, Shaipulah N F M, Sun P, Haring M A, Dudareva N, Li Y, Schuurink R C. 2022. ODORANT 1 targets multiple metabolic networks in petunia flowers. Plant J, 109 (5):1134-1151. doi: 10.1111/tpj.v109.5 URL
[6]	Burger H, Buttala S, Koch H, Ayasse M, Johnson S D, Stevenson P C. 2024. Nectar cardenolides and floral volatiles mediate a specialized wasp pollination system. Journal of Experimental Biology, 227 (1):jeb246156.
[7]	Cao X M, Wei C Y, Duan W Y, Gao Y, Kuang J F, Liu M C, Chen K S, Klee H, Zhang B. 2021. Transcriptional and epigenetic analysis reveals that NAC transcription factors regulate fruit flavor ester biosynthesis. The Plant Journal, 106 (3):785-800. doi: 10.1111/tpj.15200 pmid: 33595854
[8]	Chen Fang, She Tingting, Zhang Lin, Gao Haotian, Li Guoliang, WANG Jian. 2021. Mechanism of Sm-miR858 negatively regulated a R2R3-MYB transcription factor SmPAP1 in Salvia miltiorrhiza. Bulletin of Botanical Research, 41 (4):522-530. (in Chinese) doi: 10.7525/j.issn.1673-5102.2021.04.007
	陈芳, 佘婷婷, 张琳, 高浩天, 李国梁, 王健. 2021. 丹参体内Sm-miR858对R2R3-MYB转录因子SmPAP1进行靶向负调控作用研究. 植物研究, 41 (4):522-530. doi: 10.7525/j.issn.1673-5102.2021.04.007
[9]	Chen Y Q, Lin R Y, Kong L, Fang N Y, Fan R H, Zhong H.Q. 2024. Identification of floral aroma components and transcriptome analysis of related genes in Camellia japonica L. Journal of Nuclear Agricultural Sciences, 38 (12):2281-2293.
[10]	Cheng L, Tu G, Ma H, Zhang K, Wang X, Zhou H, Gao J, Zhou J, Yu Y, Xu Q. 2024. Alternative splicing of CsbHLH 133 regulates geraniol biosynthesis in tea plants. The Plant Journal, 120 (2):598-614. doi: 10.1111/tpj.v120.2 URL
[11]	Duan H C, Wei L H, Zhang C, Wang Y, Chen L, Lu Z, Chen P R, He C, Jia G. 2017. ALKBH10B is an RNA N6-methyladenosine demethylase affecting Arabidopsis floral transition. Plant Cell, 29 (12):2995-3011. doi: 10.1105/tpc.16.00912 URL
[12]	Feng Y, Guo Z, Zhong J, Liang Y, Zhang P, Sun M. 2023. The LibHLH22 and LibHLH 63 from Lilium‘Siberia’can positively regulate volatile terpenoid biosynthesis. Horticulturae, 9 (4):459. doi: 10.3390/horticulturae9040459 URL
[13]	Fernández-Milmanda G L. 2022. Smells like trouble:β-ocimene primes plant defenses through chromatin remodeling. Plant Physiology, 189 (2):452-453. doi: 10.1093/plphys/kiac109 pmid: 35266541
[14]	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.
[15]	Guo D, Liu Z, Raaijmakers J M, Xu Y, Yang J, Erb M, Zhang J, Zhu Y G, Xu J, Hu L. 2025. Linalool-triggered plant-soil feedback drives defense adaptation in dense maize plantings. Science, 389 (6761):eadv6675.
[16]	Han Y, Lu M, Yue S, Li K, Dong M, Liu L, Wang H, Shang F. 2022. Comparative methylomics and chromatin accessibility analysis in Osmanthus fragrans uncovers regulation of genic transcription and mechanisms of key floral scent production. Hortic Research, 9:uhac096.
[17]	Huang H, Kuo Y W, Chuang Y C, Yang Y P, Huang L M, Jeng M F, Chen W H, Chen H H. 2021. Terpene synthase-b and terpene synthase-e/f genes produce monoterpenes for Phalaenopsis bellina floral scent. Front Plant Sci, 12:700958.
[18]	Jin Z N, Wang J J, Cao X M, Wei C Y, Kuang J F, Chen K S, Zhang B. 2022. Peach fruit PpNAC1 activates PpFAD3-1 transcription to provide ω-3 fatty acids for the synthesis of short-chain flavor volatiles. Horticulture Research, 9:uhac085.
[19]	Jing T, Du W, Gao T, Wu Y, Zhang N, Zhao M, Jin J, Wang J, Schwab W, Wan X, Song C. 2021. Herbivore-induced DMNT catalyzed by CYP82D 47 plays an important role in the induction of JA-dependent herbivore resistance of neighboring tea plants. Plant Cell Environ, 44 (4):1178-1191. doi: 10.1111/pce.v44.4 URL
[20]	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.
[21]	Lan Y, Zhang K, Wang L, Liang X, Liu H X, Zhang X, Jiang N Q, Wu M, Yan H, Xiang Y. 2023. The R2R3-MYB transcription factor OfMYB21 positively regulates linalool biosynthesis in Osmanthus fragrans flowers. International Journal of Biological Macromolecules, 242:126099.
[22]	Li Zhi, YangGuodong, Duan Yifan, Zhao Youpeng, Cheng Jurong, Wang Xianrong. 2014. Study on major pollinators and their flower-visiting behavior of Osmanthus fragrans. Journal of Nanjing Forestry University(Natural Sciences Edition), 38 (Suppl):47-50. (in Chinese)
	李稚, 杨国栋, 段一凡, 范怡, 赵友朋, 成举荣, 王贤荣. 2014. 桂花访花昆虫及其访花行为观察. 南京林业大学学报(自然科学版), 38 (增刊):47-50. doi: 10.3969/j.issn.1000-2006.2014.S1.010
[23]	Liu Xiaozhou, Fan Yanping, Yu Rangcai, Yue Yuechong, Li Xinyue, Wang Yunwei. 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.
[24]	Liu Y, Wang Q, Abbas F, Zhou Y, He J, Fan Y, Yu R. 2023. Light regulation of LoCOP1 and its role in floral scent biosynthesis in Lilium‘Siberia’. Plants, 12 (10):2004. doi: 10.3390/plants12102004 URL
[25]	Patrick R M, Huang X Q, Dudareva N, Li Y. 2023. Rhythmic histone acetylation acts in concert with day-night oscillation of the floral volatile metabolic network. New Phytologist, 241 (4):1829-1839. doi: 10.1111/nph.v241.4 URL
[26]	Portillo-Estrada M, Ariza-Carricondo C, Ceulemans R. 2020. Outburst of senescence-related VOC emissions from a bioenergy poplar plantation. Plant Physiology and Biochemistry, 148:324-332. doi: S0981-9428(20)30024-3 pmid: 32004916
[27]	Shi S, Zhang S, Wu J, Liu X, Zhang Z. 2022. Identification of long non-coding RNAs involved in floral scent of Rosa hybrida. Frontiers in Plant Science, 13:996474.
[28]	Shor E, Vainstein A. 2024. Petunia PHYTOCHROME INTERACTING FACTOR 4/5 transcriptionally activates key regulators of floral scent. Plant Molecular Biology, 114 (3):66. doi: 10.1007/s11103-024-01455-8 pmid: 38816626
[29]	Silva D M, Ferraz T S O, Domingos-Melo A, Ayasse M, Milet-Pinheiro P. 2025. Chemical composition of floral perfumes in Mormodes (Orchidaceae:implications for selective attraction of euglossine bees. Annals of Botany, 136 (4):doi:10.1093/aob/mcaf133.
[30]	Spitzer-Rimon B, Farhi M, Albo B, Cna’ani A, Ben Zvi M M, Masci T, Edelbaum O, Yu Y X, Shklarman E, Ovadis M, Vainstein A. 2012. The R2R3-MYB-Like regulatory factor EOBI,acting downstream of EOBII,regulates scent production by activating ODO1 and structural scent-related genes in petunia. The Plant Cell, 24 (12):5089-5105. doi: 10.1105/tpc.112.105247 pmid: 23275577
[31]	Tu M, Liu N, He Z S, Dong X M, Gao T Y, Zhu A, 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
[32]	Vlachonasios K E. 2021. Histone acetylation:a requirement for petunia floral scent. Journal of Experimental Botany, 72 (10):3493-3495. doi: 10.1093/jxb/erab092 URL
[33]	Wang F, Liu L, Yu R, Li X, Yu Y, Li X, Yue Y, Fan Y. 2025. Function of floral fragrance-related microRNAs and their targets in Hedychium coronarium. BMC Genomics, 26 (1):430. doi: 10.1186/s12864-025-11583-0
[34]	Wang Jinxuan, Ni Ying, Meng Xin, Leng Zhuo, Ma Bo, Leng Pingsheng, Wu Jing, Hu Zenghui. 2025. Functional characterization of SoCHR35 from Syringa oblata involved in ocimene biosynthesis. Acta Horticulturae Sinica, 52 (12):2655-2665. (in Chinese)
	汪进萱, 倪莹, 孟昕, 冷卓, 马波, 冷平生, 吴静, 胡增辉. 2025. 紫丁香SoCHR35参与罗勒烯合成的功能解析. 园艺学报, 52 (12):2655-2665.
[35]	Wang S, Shi M, Zhang Y, Pan Z, Xie X, Zhang L, Sun P, Feng H, Xue H, Fang C, Zhao J. 2022. The R2R3-MYB transcription factor FaMYB 63 participates in regulation of eugenol production in strawberry. Plant Physiol, 188 (4):2146-2165. doi: 10.1093/plphys/kiac014 URL
[36]	Wei L J, Wei S H, Hu D L, Feng L, Liu Y.Y, Liu H W, Liao W B. 2022. Comprehensive flavor analysis of volatile components during the vase period of cut lily(Lilium spp.‘Manissa’)flowers by HS-SPME/GC-MS combined with E-Nose technology. Frontiers in Plant Science, 13: 822956.
[37]	Wu Q, Huang Q, Guan H, Zhang X, Bao M, Bendahmane M, Fu X. 2022. Comprehensive genome-wide analysis of histone acetylation genes in roses and expression analyses in response to heat stress. Genes (Basel), 13 (6):980. doi: 10.3390/genes13060980 URL
[38]	Yang T, Stoopen G, Thoen M, Wiegers G, Jongsma M A. 2013. Chrysanthemum expressing a linalool synthase gene‘smells good’,but‘tastes bad’to western flower thrips. Plant Biotechnology Journal, 11 (7):875-882. doi: 10.1111/pbi.2013.11.issue-7 URL
[39]	Yang Y Y, Zhang X, Han M Z, Hu Z H, Wu J, Ma N, Leng P S, Zhou X F. 2023. LiMYB 108 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
[40]	Zang N, Li X, Zhang Z, Liu W, Qi L, Yang Y, Sun Q, Yin Z, Wang A. 2024. Transcription factors PuNAC37/PuWRKY74 and E 3 ubiquitin ligase PuRDUF2 inhibit volatile ester synthesis in‘Nanguo’pear. Plant Physiol, 197 (1):kiae635.
[41]	Zhang Shiyuan, Liu Zilin, Xu Jianming, Hu Lingfei. 2025. Regulatory effects and research prospects of green leaf volatiles on plant resistance and soil ecology. Journal of Zhejiang University(Agriculture and Life Sciences), 51 (2):318-326. (in Chinese)
	张诗苑, 刘子琳, 徐建明, 胡凌飞. 2025. 绿叶挥发物对植物抗逆和土壤生态的调控作用与研究展望. 浙江大学学报(农业与生命科学版), 51 (2):318-326.
[42]	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
[43]	Zhou L, Wu S, Chen Y, Huang R, Cheng B, Mao Q, Liu T, Liu Y, Zhao K, Pan H, Yu C, Gao X, Luo L, Zhang Q. 2024. Multi-omics analyzes of Rosa gigantea illuminate tea scent biosynthesis and release mechanisms. Nature Communications, 15:8469. doi: 10.1038/s41467-024-52782-9

调控类型 Regulate type	植物种类 Plant species	转录因子/调控因子 Transcription factor/Regulator	影响的花香物质/代谢产物 Affected floral volatiles/Metabolites	参考文献 Reference
MYB转录因子家族 MYB Transcription factor family	桂花Osmanthus fragrans	OfMYB21	芳樟醇Linalool	Lan et al.,2023
	草莓 Fragaria × ananassa	FaMYB63（R2R3-MYB）	丁香酚Eugenol	Wang et al.,2022
	西藏虎头兰Cymbidium tracyanum	CtMYB2、CtMYB3	萜类花香物质 Terpenoid floral volatiles	Tu et al.,2025
	百合Lilium brownii	LiMYB108	单萜类花香物质 Monoterpenoid floral volatiles	Yang et al.,2023
	除虫菊Tanacetum cinerariifolium	TcMYB8	除虫菊酯（次生代谢产物） Pyrethrins（Secondary metabolites）	Zhou et al.,2022
	矮牵牛Petunia hybrida	EOBI（R2R3-MYB）	甲基苯甲酸酯、水杨酸甲酯、异丁香酚等 Methyl benzoate，methyl salicylate，isoeugenol，etc.	Spitzer et al.,2012
bHLH转录因子家族 bHLH Transcription factor family	茶树Camellia sinensis	CsbHLH133；CsbHLH133-AS	香叶醇Geraniol	Cheng et al.,2024
	玫瑰Rosa rugosa	RrbHLH105	香叶醇Geraniol	Bao et al.,2025
	月季Rosa chinensis	bHLH（未命名） bHLH（Unidentified）	苯丙素类香气成分（丁香酚、异丁香酚） Phenylpropanoid aroma constituents（e.g.，Eugenol and Isoeugenol）	Zhou et al.,2024
	百合Lilium brownii	LibHLH22、LibHLH63	萜类花香物质 Terpenoid floral volatiles	Feng et al.,2023
AP2/ERF及其他转录因子 AP2/ERF as well as additional transcription factors	西藏虎头兰Cymbidium tracyanum	CtAP2/ERF1、CtAP2/ERF4；CtbZIP1（bZIP）	萜类化合物 Terpenoids	Tu et al.,2025
	山茶花amellia japonica	MYB、bHLH家族成员（推测） MYB、bHLH（Candidate family member）	芳樟醇、苯丙素类香气成分 Linalool and phenylpropanoid fragrance compounds	Chen et al.,2024
	矮牵牛Petunia hybrida	PhPIF4/5；EOBII	甲基苯甲酸酯类挥发物 Methyl benzoate-type volatiles	Shor & Vainstein,2024

调控类型 Regulate type	植物种类 Plant species	转录因子/调控因子 Transcription factor/Regulator	影响的花香物质/代谢产物 Affected floral volatiles/Metabolites	参考文献 Reference
MYB转录因子家族 MYB Transcription factor family	桂花Osmanthus fragrans	OfMYB21	芳樟醇Linalool	Lan et al.,2023
	草莓 Fragaria × ananassa	FaMYB63（R2R3-MYB）	丁香酚Eugenol	Wang et al.,2022
	西藏虎头兰Cymbidium tracyanum	CtMYB2、CtMYB3	萜类花香物质 Terpenoid floral volatiles	Tu et al.,2025
	百合Lilium brownii	LiMYB108	单萜类花香物质 Monoterpenoid floral volatiles	Yang et al.,2023
	除虫菊Tanacetum cinerariifolium	TcMYB8	除虫菊酯（次生代谢产物） Pyrethrins（Secondary metabolites）	Zhou et al.,2022
	矮牵牛Petunia hybrida	EOBI（R2R3-MYB）	甲基苯甲酸酯、水杨酸甲酯、异丁香酚等 Methyl benzoate，methyl salicylate，isoeugenol，etc.	Spitzer et al.,2012
bHLH转录因子家族 bHLH Transcription factor family	茶树Camellia sinensis	CsbHLH133；CsbHLH133-AS	香叶醇Geraniol	Cheng et al.,2024
	玫瑰Rosa rugosa	RrbHLH105	香叶醇Geraniol	Bao et al.,2025
	月季Rosa chinensis	bHLH（未命名） bHLH（Unidentified）	苯丙素类香气成分（丁香酚、异丁香酚） Phenylpropanoid aroma constituents（e.g.，Eugenol and Isoeugenol）	Zhou et al.,2024
	百合Lilium brownii	LibHLH22、LibHLH63	萜类花香物质 Terpenoid floral volatiles	Feng et al.,2023
AP2/ERF及其他转录因子 AP2/ERF as well as additional transcription factors	西藏虎头兰Cymbidium tracyanum	CtAP2/ERF1、CtAP2/ERF4；CtbZIP1（bZIP）	萜类化合物 Terpenoids	Tu et al.,2025
	山茶花amellia japonica	MYB、bHLH家族成员（推测） MYB、bHLH（Candidate family member）	芳樟醇、苯丙素类香气成分 Linalool and phenylpropanoid fragrance compounds	Chen et al.,2024
	矮牵牛Petunia hybrida	PhPIF4/5；EOBII	甲基苯甲酸酯类挥发物 Methyl benzoate-type volatiles	Shor & Vainstein,2024