芳樟醇合成调控机制及其生态功能研究

doi:10.16420/j.issn.0513-353x.2024-0845

摘要/Abstract

摘要：

芳樟醇是一种重要的单萜类植物次生代谢物，广泛分布于植物的花、果、叶、茎和根中。其生物合成主要在质体内通过MEP途径完成，受到转录、表观遗传及转录后等多层次的分子调控。作为植物香气的关键成分，芳樟醇在植物与传粉昆虫、植食性昆虫及其天敌之间的生态相互作用中发挥着重要作用。此外，芳樟醇还参与植物对病原体和非生物胁迫的防御响应。本文中综述了芳樟醇的合成路径与调控机制，系统总结了其在吸引传粉者、抗病虫害及应对非生物胁迫等方面的研究进展，探讨其在植物适应复杂环境中的重要意义。同时，对芳樟醇在田间生态环境中的调控效应、单一对映体及衍生物的功能特异性、以及多重胁迫中的协同作用机制研究提出了展望，以期为园艺作物绿色农业应用提供理论支撑和技术指导。

关键词: 芳樟醇, 萜类次生代谢物, 生物合成, 调控机制, 生态功能

Abstract:

Linalool is an important monoterpene secondary metabolite in plants，widely present in flowers，fruits，leaves，stems and roots. Its biosynthesis mainly occurs in plastids via the MEP pathway and is regulated by multiple molecular mechanisms，including transcriptional，epigenetic，and post-transcriptional controls. As a key component of plant aroma，linalool plays a significant role in the ecological interactions between plants，pollinators，herbivorous insects，and their natural enemies. In addition，linalool is involved in plant defense responses against pathogens and abiotic stresses. This review summarizes the pathways and regulatory mechanisms of linalool synthesis，providing a comprehensive overview of its ecological functions in attracting pollinators，resisting pests and diseases，and responding to abiotic stress. The review also explores the significance of linalool in plant adaptation to complex environments. Furthermore，it discusses the regulatory effects of linalool in field ecosystems，the functional specificity of individual enantiomers and their derivatives，and the synergistic mechanisms in response to multiple stresses，with the aim of providing theoretical support and technical guidance for the green agricultural applications of horticultural crops.

Key words: linalool, terpenoid secondary metabolites, biosynthesis, regulation, ecological function

冯玲佳, 刘毓婕, 何林彤, 王新超, 叶萌. 芳樟醇合成调控机制及其生态功能研究[J]. 园艺学报, 2025, 52(8): 2270-2290.

FENG Lingjia, LIU Yujie, HE Lintong, WANG Xinchao, and YE Meng. Research Progress on the Regulation Mechanism of Linalool Synthesis and Its Ecological Functions[J]. Acta Horticulturae Sinica, 2025, 52(8): 2270-2290.

图/表 4

参考文献 112

[1]	Abbas F, Ke Y G, Yu R C, Fan Y P. 2018. Functional characterization and expression analysis of two terpene synthases involved in floral scent formation in Lilium‘Siberia’. Planta, 249 (1):71-93.
[2]	Achhami B B, Reddy G V P, Hofland M L, Sherman J D, Peterson R K D, Weaver D K. 2021. Plant volatiles and oviposition behavior in the selection of barley cultivars by wheat stem sawfly(Hymenoptera:Cephidae). Environmental Entomology, 50 (4):940-947.
[3]	Aharoni A, Giri A P, Deuerlein S, Griepink F, de Kogel W J, Verstappen F W A, Verhoeven H A, Jongsma M A, Schwab W, Bouwmeester H J. 2003. Terpenoid metabolism in wild-type and transgenic arabidopsis plants. Plant Cell, 15 (12):2866-2884.
[4]	Aharoni A, Giri A P, Verstappen F W A, Bertea C M, Sevenier R, Sun Z K, Jongsma M A, Schwab W, Bouwmeester H J. 2004. Gain and loss of fruit flavor compounds produced by wild and cultivated strawberry species. Plant Cell, 16 (11):3110-3131.
[5]	Altúzar A, Malo E, González-Hernández H, Rojas J. 2007. Electrophysiological and behavioural responses of Scyphophorus acupunctatus(Col.,Curculionidae)to Agave tequilana volatiles. Journal of Applied Entomology,131:121-127.
[6]	Aprotosoaie A C, Hăncianu M, Costache I I, Miron A. 2014. Linalool:a review on a key odorant molecule with valuable biological properties. Flavour and Fragrance Journal, 29 (4):193-219.
[7]	Arimura G-i, Kost C, Boland W. 2005. Herbivore-induced,indirect plant defences. Biochimica et Biophysica Acta, 1734 (2):91-111.
[113]	Taniguchi S, Hosokawa-Shinonaga Y, Tamaoki D, Yamada S, Akimitsu K, Gomi K. 2014. Jasmonate induction of the monoterpene linalool confers resistance to rice bacterial blight and its biosynthesis is regulated by JAZ protein in rice. Plant Cell and Environment, 37 (2):451-461.
[114]	Terry I, Moore C J, Walter G H, Forster P I, Roemer R B, Donaldson J D, Machin P J. 2004. Association of cone thermogenesis and volatiles with pollinator specificity in Macrozamia cycads. Plant Systematics and Evolution, 243 (3):233-247.
[115]	Theis N. 2006. Fragrance of Canada thistle(Cirsium arvense)attracts both floral herbivores and pollinators. Journal of Chemical Ecology, 32 (5):917-927.
[116]	Ulland S, Ian E, Borg-Karlson A K, Mustaparta H. 2006. Discrimination between enantiomers of linalool by olfactory receptor neurons in the cabbage moth Mamestra brassicae(L.). Chemical Senses, 31 (4):325-334.
[117]	van Schie C C N, Haring M A, Schuurink R C. 2007. Tomato linalool synthase is induced in trichomes by jasmonic acid. Plant Molecular Biology, 64 (3):251-263.
[118]	Vaughn S F, Spencer G F. 1991. Volatile monoterpenes inhibit potato tuber sprouting. American Potato Journal, 68 (12):821-831.
[119]	Vranová E, Coman D, Gruissem W. 2012. Structure and dynamics of the isoprenoid pathway network. Molecular Plant, 5 (2):318-333.
[120]	Wang Peiyun, Li Ziang, Bai Yang, Yang Ping, Yin Chengpeng, Li Chuanrong, Zhang Xinwen, Song Xiuhua. 2024. Cloning and functional verification of linalool synthase gene PsTPS14 in tree peony‘High Noon’. Acta Horticulturae Sinica, 51 (6):1273-1283. (in Chinese)
	王佩云, 李子昂, 白杨, 杨萍, 尹承芃, 李传荣, 张馨文, 宋秀华. 2024. ‘海黄’牡丹芳樟醇合酶基因PsTPS14的克隆及功能验证. 园艺学报, 51 (6):1273-1283.
[121]	Wang Qifang, Wang Xiaoyun, Li Haoshen, Yang Xiaoyu, Zhang Ruimin, Gong Biao, Li Xiuming, Shi Qinghua. 2023. Effects of linalool on Botrytis cinerea growth and control of tomato gray mold. Chinese Journal of Applied Ecology, 34 (1):213-220. (in Chinese)
[8]	Balducci M G, Martins D J, Johnson S D. 2019. Pollination of the long-spurred African terrestrial orchid Bonatea steudneri by long-tongued hawkmoths,notably Xanthopan morganii. Plant Systematics and Evolution, 305 (9):765-775.
[9]	Bergman M E, Davis B, Phillips M A. 2019. Medically useful plant trpenoids:biosynthesis,occurrence,and mechanism of action. Molecules, 24 (21):3961.
[10]	Bhargav P, Chaurasia S, Kumar A, Srivastava G, Pant Y, Chanotiya C S, Ghosh S. 2023. Unraveling the terpene synthase family and characterization of BsTPS 2 contributing to(S)-(+)-linalool biosynthesis in Boswellia. Plant Molecular Biology, 113 (4-5):219-236.
[11]	Bloch K, Chaykin S, Phillips A H, de Waard A. 1959. Mevalonic acid pyrophosphate and isopentenylpyrophosphate. Journal of Biological Chemistry, 234 (10):2595-2604.
[12]	Boachon B, Junker R R, Miesch L, Bassard J E, Höfer R, Caillieaudeaux R, Seidel D E, Lesot A, Heinrich C, Ginglinger J F, Allouche L, Vincent B, Wahyuni D S, Paetz C, Beran F, Miesch M, Schneider B, Leiss K, Werck-Reichhart D. 2015. CYP76C1(cytochrome P450)-mediated linalool metabolism and the formation of volatile and soluble linalool oxides in Arabidopsis flowers:a strategy for defense against floral antagonists. Plant Cell, 27 (10):2972-2990.
[13]	Bonnländer B, Cappuccio R, Liverani F S, Winterhalter P. 2006. Analysis of enantiomeric linalool ratio in green and roasted coffee. Flavour and Fragrance Journal, 21 (4):637-641.
[121]	王启方, 王晓云, 李浩森, 杨晓玉, 张锐敏, 巩彪, 李秀明, 史庆华. 2023. 芳樟醇对灰葡萄孢生长的影响及对番茄灰霉病的防控效果. 应用生态学报, 34 (1):213-220.
[122]	Wang W, Wang M Y, Zeng Y L, Chen X Y, Wang X Y, Barrington A M, Tao J M, Atkinson R G, Nieuwenhuizen N J. 2023. The terpene synthase(TPS)gene family in kiwifruit shows high functional redundancy and a subset of TPS likely fulfil overlapping functions in fruit flavour,floral bouquet and defence. Molecular Horticulture, 3 (1):9.
[123]	Wei C, Liu H, Cao X, Zhang M, Li X, Chen K, Zhang B. 2021. Synthesis of flavor-related linalool is regulated by PpbHLH1 and associated with changes in DNA methylation during peach fruit ripening. Plant Biotechnology Journal, 19 (10):2082-2096.
[124]	Wei C C, 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.
[125]	Weidenhamer J D, Macias F A, Fischer N H, Williamson G B. 1993. Just how insoluble are monoterpenes? Journal of Chemical Ecology, 19 (8):1799-1807.
[126]	Wu S Q, Schalk M, Clark A, Miles R B, Coates R, Chappell J. 2006. Redirection of cytosolic or plastidic isoprenoid precursors elevates terpene production in plants. Nature Biotechnology, 24 (11):1441-1447.
[127]	Xiao Y T, Wang Q, Erb M, Turlings T C J, Ge L Q, Hu L F, Li J C, Han X, Zhang T F, Lu J, Zhang G R, Lou Y G. 2012. Specific herbivore-induced volatiles defend plants and determine insect community composition in the field. Ecology Letters, 15 (10):1130-1139.
[14]	Borg-Karlson A K, Unelius C R, Valterová I, Anders Nilsson L. 1996. Floral fragrance chemistry in the early flowering shrub Daphne mezereum. Phytochemistry, 41 (6):1477-1483.
[15]	Cao Y, Hu S L, Dai Q L, Liu Y S. 2014. Tomato terpene synthases TPS5 and TPS 39 account for a monoterpene linalool production in tomato fruits. Biotechnology Letters, 36 (8):1717-1725.
[16]	Cao Y, Wang J, Germinara G S, Wang L J, Yang H, Gao Y L, Li C. 2020. Behavioral responses of Thrips hawaiiensis(Thysanoptera:Thripidae)to volatile compounds identified from Gardenia jasminoides ellis(Gentianales:Rubiaceae). Insects, 11 (7):408.
[17]	Carroll M J, Schmelz E A, Meagher R L, Teal P E A. 2006. Attraction of Spodoptera frugiperda larvae to volatiles from herbivore-damaged maize seedlings. Journal of Chemical Ecology, 32 (9):1911-1924.
[18]	Chappell J, Wolf F, Proulx J, Cuellar R, Saunders C. 1995. Is the reaction catalyzed by 3-hydroxy-3-methylglutaryl coenzyme a reductase a rate-limiting step for isoprenoid biosynthesis in plants? Plant Physiology, 109 (4):1337-1343.
[19]	Chen X, Yauk Y K, Nieuwenhuizen N J, Matich A J, Wang M Y, Perez R L, Atkinson R G, Beuning L L. 2010. Characterisation of an(S)-linalool synthase from kiwifruit(Actinidia arguta)that catalyses the first committed step in the production of floral lilac compounds. Functional Plant Biology,37:232-243.
[20]	Chen Zumin, Xiao Nuoya, Zhang Yanxia, Shi Xiaomin, Guo Shuaiqi, Gao Hu, Wang Zhenping. 2021. Effects of water stress on the volatile compounds and related biosynthetic genes expression in‘Muscat Hamburg’grape berries. Acta Horticulturae Sinica, 48 (5):883-896. (in Chinese)
	陈祖民, 校诺娅, 张艳霞, 史晓敏, 郭帅奇, 高虎, 王振平. 2021. 水分胁迫对‘玫瑰香’葡萄果实挥发性化合物及相关基因表达的影响. 园艺学报, 48 (5):883-896.
[128]	Xiao Y Y, Tan H B, Huang H T, Yu J Z, Zeng L T, Liao Y Y, Wu P, Yang Z Y. 2022. Light synergistically promotes the tea green leafhopper infestation-induced accumulation of linalool oxides and their glucosides in tea(Camellia sinensis). Food Chemistry,394:133460.
[129]	Yactayo-Chang J P, Broadhead G T, Housler R J, Resende M F R,Jr., Verma K, Louis J, Basset G J, Beck J J, Block A K. 2024. Maize terpene synthase 1 impacts insect behavior via the production of monoterpene volatiles β-myrcene and linalool. Phytochemistry,218:113957.
[130]	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.
[131]	Yang Z R, Zhan T, Xie C Z, Huang S, Zheng X S. 2023. Genome-wide analyzation and functional characterization on the TPS family provide insight into the biosynthesis of mono-terpenes in the camphor tree. Plant Physiology and Biochemistry,196:55-64.
[132]	Yang Z Z, Li Y Q, Gao F Z, Jin W, Li S Y, Kimani S, Yang S, Bao T T, Gao X, Wang 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.
[133]	Yao C C, Du L X, Liu Q S, Hu X Y, Ye W F, Turlings T C J, Li Y H. 2023a. Stemborer-induced rice plant volatiles boost direct and indirect resistance in neighboring plants. New Phytologist, 237 (6):2375-2387.
[21]	Ciriello M, Cirillo V, Formisano L, De Pascale S, Romano R, Fusco G M, Nicastro R, Carillo P, Kyriacou M C, Soteriou G A, Rouphael Y. 2023. Salt-induced stress impacts the phytochemical composition and aromatic profile of three types of basil in a genotype-dependent mode. Plants, 12 (11):2167.
[22]	Danner H, Boeckler G A, Irmisch S, Yuan J S, Chen F, Gershenzon J, Unsicker S B, Köllner T G. 2011. Four terpene synthases produce major compounds of the gypsy moth feeding-induced volatile blend of Populus trichocarpa. Phytochemistry, 72 (9):897-908.
[23]	Dicke M, Van Beek T A, Posthumus M A, Ben Dom N, Van Bokhoven H, De Groot A. 1990. Isolation and identification of volatile kairomone that affects acarine predatorprey interactions involvement of host plant in its production. Journal of Chemical Ecology, 16 (2):381-396.
[24]	Dong F, Yang Z, Baldermann S, Sato Y, Asai T, Watanabe N. 2011. Herbivore-induced volatiles from tea(Camellia sinensis)plants and their involvement in intraplant communication and changes in endogenous nonvolatile metabolites. Journal of Agricultural and Food Chemistry, 59 (24):13131-13135.
[25]	Dötterl S, Jürgens A, Seifert K, Laube T, Weissbecker B, Schütz S. 2006. Nursery pollination by a moth in Silene latifolia:the role of odours in eliciting antennal and behavioural responses. New Phytologist, 169 (4):707-718.
[26]	Dudareva N, Cseke L, Blanc V M, Pichersky E. 1996. Evolution of floral scent in Clarkia:novel patterns of S-linalool synthase gene expression in the C. breweri flower. Plant Cell, 8 (7):1137-1148.
[134]	Yao Y, Fu W H, Yu Y, Wan S Y, Zhang W P, Ming R. 2023b. The synthesis of papaya fruit flavor-related linalool was regulated by CpTPS18 and CpNAC56. Plant Reproduction, 37 (3):295-308.
[135]	Yu Z M, Zhao C H, Zhang G H,Teixeira da Silva J A,Duan J. 2020. Genome-wide identification and expression profile of TPS gene family in Dendrobium officinale and the role of DoTPS10 in linalool biosynthesis. International Journal of Molecular Sciences, 21 (15):5419.
[136]	Yu Z M, Zhang G H,Teixeira da Silva J A,Zhao C H,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.
[137]	Yuan J S, Köllner T G, Wiggins G, Grant J, Degenhardt J, Chen F. 2008. Molecular and genomic basis of volatile-mediated indirect defense against insects in rice. Plant Journal, 55 (3):491-503.
[138]	Yue R, Li Y L, Qi Y J, Liang X Y, Zheng Z Q, Ye Z L, Tong W, Si X Y, Zhang Y R, Xia E H, Li P H. 2025. Divergent MYB paralogs determine spatial distribution of linalool mediated by JA and DNA demethylation participating in aroma formation and cold tolerance of tea plants. Plant Biotechnology Journal, 23 (5):1455-1475.
[139]	Yue Y C, Yu R C, Fan Y P. 2014. Characterization of two monoterpene synthases involved in floral scent formation in Hedychium coronarium. Planta, 240 (4):745-762.
[140]	Yue Y C, Yu R C, Fan Y P. 2015. Transcriptome profiling provides new insights into the formation of floral scent in Hedychium coronarium. BMC Genomics,16:470.
[141]	Zeng X L, Liu C, Zheng R R, Cai X, Luo J, Zou J J, Wang C Y. 2016. Emission and accumulation of monoterpene and the key terpene synthase(TPS)associated with monoterpene biosynthesis in Osmanthus fragrans Lour. Frontiers in Plant Science,6:1232.
[27]	Falara V, Akhtar T A, Nguyen T T H, Spyropoulou E A, Bleeker P M, Schauvinhold I, Matsuba Y, Bonini M E, Schilmiller A L, Last R L, Schuurink R C, Pichersky E. 2011. The tomato terpene synthase gene family. Plant Physiology, 157 (2):770-789.
[28]	Fan Y Z, Tian C, Tong S Y, Liu Q, Xu F, Shi B B, Ai H L, Liu J K. 2023. The antifungal properties of terpenoids from the endophytic fungus Bipolaris eleusines. Natural Products and Bioprospecting, 13 (1):43.
[29]	Galata M, Sarker L S, Mahmoud S S. 2014. Transcriptome profiling,and cloning and characterization of the main monoterpene synthases of Coriandrum sativum L. Phytochemistry,102:64-73.
[30]	Gao F Z, Liu B F, Li M, Gao X Y, Fang Q, Liu C, Ding H, Wang L, Gao X. 2018. Identification and characterization of terpene synthase genes accounting for volatile terpene emissions in flowers of Freesia x hybrida. Journal of Experimental Botany, 69 (18):4249-4265.
[31]	Gao J, Chen Y C, Gao M, Wu L W, Zhao Y X, Wang Y D. 2023. LcWRKY17,a WRKY transcription factor from Litsea cubeba,effectively promotes monoterpene synthesis. International Journal of Molecular Sciences, 24 (8):7210.
[32]	Gasmi L, Martínez-Solís M, Frattini A, Ye M, Collado María C, Turlings T C J, Erb M, Herrero S. 2018. Can herbivore-induced volatiles protect plants by increasing the herbivores’susceptibility to natural pathogens? Applied and Environmental Microbiology, 85 (1):e01468-01418.
[142]	Zhang T X, Guo Y H, Shi X J, Yang Y J, Chen J T, Zhang Q X, Sun M. 2020. Overexpression of LiTPS2 from a cultivar of lily(Lilium‘Siberia’)enhances the monoterpenoids content in tobacco flowers. Plant Physiology and Biochemistry,151:391-399.
[143]	Zhang X H, Zhang X H, Teixeira da Silva J A, Niu M Y, Zhang T, Liu H F, Zheng F, Yuan Y F, Li Y, Fang L, Zeng S J, Ma G H. 2021. Functional characterization of an Indian sandalwood(Santalum album L.)dual-localized bifunctional nerolidol/linalool synthase gene involved in stress response. Phytochemistry,183:112610.
[144]	Zhang Xianying, Huo Zhiguo, You Changyan, Hu Fei. 2014. Effects of volatiles in twenty non-host plants on the repellented and attractive behaviors of brown planthopper. Nilaparvata lugens., Journal of South China Agricultural University, 35 (3):63-68. (in Chinese)
	张献英, 霍治国, 犹昌艳, 胡飞. 2014. 20种非寄主植物挥发物对褐飞虱拒避与引诱行为的影响. 华南农业大学学报, 35 (3):63-68.
[145]	Zhang Y, Zhang T, Wang X H, Bian Z P, Zhang X F, Yang G Q, Lu Y H. 2024. Volatiles from essential oils of three Lamiaceae plants repel the winged cotton aphid,disturb its feeding behavior and reduce its fecundity. Pest Management Science, 80 (9):4253-4263.
[146]	Zhang Z Z, Li Y B, Qi L, Wan X C. 2006. Antifungal activities of major tea leaf volatile constituents toward Colletorichum camelliae Massea. Journal of Agricultural and Food Chemistry, 54 (11):3936-3940.
[147]	Zhao H Y, Li M, Zhao Y Y, Lin X J, Liang H L, Wei J S, Wei W K, Ma D, Zhou Z Y, Yang J F. 2021. A comparison of two monoterpenoid synthases reveals molecular mechanisms associated with the difference of bioactive monoterpenoids between Amomum villosum and Amomum longiligulare. Frontiers in Plant Science,12:695551.
[33]	Gharbi K, Tay J W. 2022. Fumigant toxicity of essential oils against Frankliniella occidentalis and F. insularis(Thysanoptera:Thripidae)as affected by polymer release and adjuvants. Insects, 13 (6):496.
[34]	Ginglinger J F, Boachon B, Höfer R, Paetz C, Köllner T G, Miesch L, Lugan R, Baltenweck R, Mutterer J, Ullmann P, Beran F, Claudel P, Verstappen F, Fischer M J C, Karst F, Bouwmeester H, Miesch M, Schneider B, Gershenzon J, Ehlting J, Werck-Reichhart D. 2013. Gene coexpression analysis reveals complex metabolism of the monoterpene alcohol linalool in Arabidopsis Flowers. Plant Cell, 25 (11):4640-4657.
[35]	Gong Lintao, Su Xiujuan, Liao Yan, Keremuhan·Wusiman, Zhou Di, Yin Songsong. 2021. Cloning,expression and enzyme activity detection of linalool synthase gene in Lavender. Crops,(6):78-87. (in Chinese)
	龚林涛, 苏秀娟, 廖燕, 克热木汗·吾斯曼, 周迪, 尹松松. 2021. 薰衣草芳樟醇合酶基因的克隆、表达及酶活性检测. 作物杂志,(6):78-87.
[36]	Green S A, Chen X, Nieuwenhuizen N J, Matich A J, Wang M Y, Bunn B J, Yauk Y K, Atkinson R G. 2012. Identification,functional characterization,and regulation of the enzyme responsible for floral(E)-nerolidol biosynthesis in kiwifruit(Actinidia chinensis). Journal of Experimental Botany, 63 (5):1951-1967.
[37]	Günther C S, Chervin C, Marsh K B, Newcomb R D, Souleyre E J F. 2011. Characterisation of two alcohol acyltransferases from kiwifruit(Actinidia spp.) reveals distinct substrate preferences. Phytochemistry, 72 (8):700-710.
[38]	Guo Y H, Guo Z Y, Zhong J, Liang Y L, Feng Y, Zhang P, Zhang Q, Sun M X. 2023. Positive regulatory role of R2R3 MYBs in terpene biosynthesis in Lilium‘Siberia’. Horticultural Plant Journal, 9 (5):1024-1038.
[39]	Han Y J, Lu M M, Yue S M, Li K, Dong M F, Liu L X, Wang H Y, Shang F D. 2022. Comparative methylomics and chromatin accessibility analysis in Osmanthus fragrans uncovers regulation of genic transcription and mechanisms of key floral scent production. Horticulture Research,9:uhac096.
[40]	Hany A F, Wagner de S T, José C Z. 2021. Toxicity and repellent activity of monoterpene enantiomers to rice weevils(Sitophilus oryzae). Pest Management Science, 77 (7):3500-3507.
[41]	He J, Fandino R A, Halitschke R, Luck K, Köllner T G, Murdock M H, Ray R, Gase K, Knaden M, Baldwin I T, Schuman M C. 2019. An unbiased approach elucidates variation in(S)-(+)-linalool,a context-specific mediator of a tri-trophic interaction in wild tobacco. Proceedings of the National Academy of Sciences, 116 (29):14651-14660.
[42]	He J, Halitschke R, Baldwin I T, Schuman M C. 2021. Natural variation in linalool metabolites:one genetic locus,many functions? Journal of Integrative Plant Biology, 63 (8):1416-1421.
[43]	Hemmerlin A, Harwood J L, Bach T J. 2012. A raison d’être for two distinct pathways in the early steps of plant isoprenoid biosynthesis? Progress in Lipid Research, 51 (2):95-148.
[44]	Höfer R, Boachon B, Renault H, Gavira C, Miesch L, Iglesias J, Ginglinger J F, Allouche L, Miesch M, Grec S, Larbat R, Werck-Reichhart D. 2014. Dual function of the cytochrome P 450 CYP76 family from Arabidopsis thaliana in the metabolism of monoterpenols and phenylurea herbicides. Plant Physiology, 166 (3):1149-1161.
[148]	Zhao M Y, Wang L, Wang J M, Jin J Y, Zhang N, Lei L, Gao T, Jing T T, Zhang S R, Wu Y, Wu B, Hu Y Q, Wan X C, Schwab W, Song C K. 2020. Induction of priming by cold stress via inducible volatile cues in neighboring tea plants. Journal of Integrative Plant Biology, 62 (10):1461-1468.
[149]	Zhou Lirong, Zhang Lingling, Xiong Shijie, Ma Jiawei, Zhu Yongxing, Sun Chong, Zhu Xuedong, Liu Yiqing. 2024. Inhibitory effect of linalool against the ginger Fusarium wilt pathogen. Food Science, 45 (6):72-79. (in Chinese)
	周丽荣, 张玲玲, 熊诗洁, 马佳伟, 朱永兴, 孙冲, 朱学栋, 刘奕清. 2024. 芳樟醇对生姜枯萎病菌的抑制作用. 食品科学, 45 (6):72-79.
[150]	Zhou Y, Deng R F, Xu X L, Yang Z Y. 2020. Enzyme catalytic efficiencies and relative gene expression levels of(R)-linalool synthase and(S)-linalool synthase determine the proportion of linalool enantiomers in camellia sinensis var. sinensis. Journal of Agricultural and Food Chemistry, 68 (37):10109-10117.
[151]	Zhou Y, Dong F, Kunimasa A, Zhang Y Q, Cheng S H, Lu J M, Zhang L, Murata A, Mayer F, Fleischmann P, Watanabe N, Yang Z Y. 2014. Occurrence of glycosidically conjugated 1-phenylethanol and its hydrolase β-primeverosidase in tea(Camellia sinensis)flowers. Journal of Agricultural and Food Chemistry, 62 (32):8042-8050.
[152]	Zhou Y, Zeng L T, Liu X Y, Gui J D, Mei X, Fu X M, Dong F, Tang J C, Zhang L Y, Yang Z Y. 2017. Formation of(E)-nerolidol in tea(Camellia sinensis)leaves exposed to multiple stresses during tea manufacturing. Food Chemistry,231:78-86.
[153]	Zhu B Q, Cai J, Wang Z Q, Xu X Q, Duan C Q, Pan Q H. 2014. Identification of a plastid-localized bifunctional nerolidol/linalool synthase in relation to linalool biosynthesis in young grape berries. International Journal of Molecular Sciences, 15 (12):21992-22010.
[45]	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. Frontiers in Plant Science,12:700958.
[46]	Huang X Z, Xiao Y T, Köllner T G, Jing W X, Kou J F, Chen J Y, Liu D F, Gu S H, Wu J X, Zhang Y J, Guo Y Y. 2017. The terpene synthase gene family in Gossypium hirsutum harbors a linalool synthase GhTPS 12 implicated in direct defence responses against herbivores. Plant Cell and Environment, 41 (1):261-274.
[47]	Ibdah M, Hino S, Nawade B, Yahyaa M, Bosamia T C, Shaltiel-Harpaz L. 2022. Identification and characterization of three nearly identical linalool/nerolidol synthase from Acorus calamus. Phytochemistry,202:113318.
[48]	Iijima Y, Gang D R, Fridman E, Lewinsohn E, Pichersky E. 2004. Characterization of geraniol synthase from the peltate glands of sweet basil. Plant Physiology, 134 (1):370-379.
[49]	Jia J W, Crock J, Lu S, Croteau R, Chen X Y. 1999.( 3R)-Linalool synthase from Artemisia annua L.: cDNA isolation,characterization,and wound induction. Archives of Biochemistry and Biophysics, 372 (1):143-149.
[50]	Jiao C Y, Gong J Q, Guo Z J, Li S W, Zuo Y X, Shen Y B. 2022. Linalool activates oxidative and calcium burst and CAM3-ACA 8 participates in calcium recovery in Arabidopsis leaves. International Journal of Molecular Sciences, 23 (10):5357.
[51]	Jiang Y, Pan X Y, Li Y M, Yang Y Z, Jia Y N, Lei B, Feng J T, Ma Z Q, Liu X L, Yan H. 2023. Linalool induces resistance against tobacco mosaic virus in tobacco plants. Plant Diseases, 107 (7):2144-2152.
[52]	Jiang Y F, Qian R J, Zhang W B, Wei G, Ma X H, Zheng J, Köllner T G, Chen F. 2020. Composition and biosynthesis of scent compounds from sterile flowers of an ornamental plant Clematis florida cv.‘Kaiser’. Molecules, 25 (7):1711.
[53]	Ke Y G, Abbas F, Zhou Y W, Yu R C, Fan Y P. 2021. Auxin-responsive R2R3-MYB transcription factors HcMYB1 and HcMYB 2 activate volatile biosynthesis in Hedychium coronarium flowers. Frontiers in Plant Science,12:710826.
[54]	Kessler A, Baldwin I T. 2001. Defensive function of herbivore-induced plant volatile emissions in nature. Science, 291 (5511):2141-2144.
[55]	Kheloul L, Anton S, Gadenne C, Kellouche A. 2020. Fumigant toxicity of Lavandula spica essential oil and linalool on different life stages of Tribolium confusum(Coleoptera:Tenebrionidae). Journal of Asia-Pacific Entomology, 23 (2):320-326.
[56]	Knudsen J T, Tollsten L. 1993. Trends in floral scent chemistry in pollination syndromes:floral scent composition in moth-pollinated taxa. Botanical Journal of the Linnean Society, 113 (3):263-284.
[57]	Knudsen J T, Tollsten L, Bergström L G. 1993. Floral scents-a checklist of volatile compounds isolated by head-space techniques. Phytochemistry, 33 (2):253-280.
[58]	Krug C, Cordeiro G D, Schäffler I, Silva C I, Oliveira R, Schlindwein C, Dötterl S, Alves-Dos-Santos I. 2018. Nocturnal bee pollinators are attracted to guarana flowers by their scents. Frontiers in Plant Science,9:.
[59]	Landmann C, Fink B, Festner M, Dregus M, Engel K-H, Schwab W. 2007. Cloning and functional characterization of three terpene synthases from lavender(Lavandula angustifolia). Archives of Biochemistry and Biophysics, 465 (2):417-429.
[60]	Laule O, Fürholz A, Chang H-S, Zhu T, Wang X, Heifetz P B, Gruissem W, Lange M. 2003. Crosstalk between cytosolic and plastidial pathways of isoprenoid biosynthesis in Arabidopsis thaliana. Proceedings of the National Academy of Sciences of the United States of Americai, 100 (11):6866-6871.
[61]	Li S S, Zhang L, Sun M, Lv M W, Yang Y, Xu W Z, Wang L S. 2023. Biogenesis of flavor-related linalool is diverged and genetically conserved in tree peony(Paeonia × suffruticosa). Horticulture Research, 10 (2):uhac253.
[62]	Lin Y L, Lee Y R, Huang W K, Chang S T, Chu F H. 2014. Characterization of S-(+)-linalool synthase from several provenances of Cinnamomum osmophloeum. Tree Genetics and Genomes, 10 (1):75-86.
[63]	Liu Cai, Xi Wan, Yuan Jinmei, Zhu Linlin, Chen Hongguo, Zou Jingjing, Zheng Riru, Wang Caiyun. 2020. Molecular loning and functional characterization of linalool synthase gene OfTPS5 in Osmanthus fragrans‘Lianzi Dangui’flowers. Acta Horticulturae Sinica, 47 (2):310-320. (in Chinese)
	刘偲, 席婉, 袁金梅, 朱琳琳, 陈洪国, 邹晶晶, 郑日如, 王彩云. 2020. 桂花‘莲籽丹桂’芳樟醇合酶基因OfTPS5的克隆及功能鉴定. 园艺学报, 47 (2):310-320.
[64]	Liu G H, Wang Q, Chen H, Wang Y X, Zhou X G, Bao D M, Wang N, Sun J, Huang F Y, Yang M, Zhang H, Yan P, Li X, Shi J, Fu J Y. 2024. Plant-derived monoterpene S-linalool and β-ocimene generated by CsLIS and CsOCS-SCZ are key chemical cues for attracting parasitoid wasps for suppressing Ectropis obliqua infestation in Camellia sinensis L. Plant Cell and Environment, 47 (3):913-927.
[65]	Liu G H, Yang M, Yang X M, Ma X Y, Fu J Y. 2021. Five TPSs are responsible for volatile terpenoid biosynthesis in Albizia julibrissin. Journal of Plant Physiology,258-259:153358.
[66]	Liu G F, Liu J J, He Z R, Wang F M, Yang H, Yan Y F, Gao M J, Gruber M Y, Wan X C, Wei S. 2017a. Implementation of CsLIS/NES in linalool biosynthesis involves transcript splicing regulation in Camellia sinensis. Plant Cell and Environment, 41 (1):176-186.
[67]	Liu H R, Cao X M, Liu X H, Xin R, Wang J J, Gao J, Wu B P, Gao L X, Xu C J, Zhang B, Grierson D, Chen K S. 2017b. UV‐B irradiation differentially regulates terpene synthases and terpene content of peach. Plant Cell and Environment, 40 (10):2261-2275.
[68]	Liu P J, Zhang X F, Wang R Y, Chen S L, Zhang T. 2023. Monoterpene alcohols induced by sweet potato weevil larvae deter conspecific adults from feeding and oviposition. Pest Management Science, 79 (10):3504-3510.
[69]	Lu Kai, Li Xin, Zhou Jialiang, Jie Xiaojun, Qi Shu, Zhou Qiang. 2010. Influence of the herbivore-induced rice volatiles on fungal disease. Chinese Science Bulletin, 55 (30):2927-2932. (in Chinese)
	卢凯, 李欣, 周嘉良, 解晓军, 戚舒, 周强. 2010. 虫害诱导的水稻挥发物抑制水稻病原菌的生长. 科学通报, 55 (30):2927-2932.
[70]	Lukas K, Harig T, Schulz S, Hadersdorfer J, Dötterl S. 2019. Flowers of European pear release common and uncommon volatiles that can be detected by honey bee pollinators. Chemoecology, 29 (5):211-223.
[71]	Luo C W, Huang Z Y, Li K, Chen X M, Chen Y, Sun Y Y. 2013. EAG responses of Apis cerana to floral compounds of a biodiesel plant,Jatropha curcas(Euphorbiaceae). Journal of Economic Entomology, 106 (4):1653-1658.
[72]	Luo Yujie, Wang Yang, Zhou Qiong, He Jie, Li Xing. 2023. Olfactory and behavioral responses of Papilio polytes(Lepidoptera:Papilioidae)adults to volatiles from the branches and leaves of citrus. Acta Entomologica Sinica, 66 (12):1612-1625. (in Chinese)
	罗宇婕, 汪洋, 周琼, 何杰, 李幸. 2023. 玉带凤蝶成虫对柑橘枝叶挥发物的嗅觉和行为反应. 昆虫学报, 66 (12):1612-1625.
[73]	Ma Qing, Ma Rui, Su Ping, Shen Ye, Chen Meilan, Jin Baolong, Ouyang Shaolin, Guo Juan, Cui Guanghong, Huang Luqi. 2023. Systematic identification of chemical forms of key terpene synthase in Cinnamomum camphora. China Journal of Chinese Materia Medica, 48 (9):2307-2315. (in Chinese)
	马青, 马蕊, 苏平, 申业, 陈美兰, 靳保龙, 欧阳少林, 郭娟, 崔光红, 黄璐琦. 2023. 樟树化学型形成关键萜类合酶的系统鉴定. 中国中药杂志, 48 (9):2307-2315.
[74]	Magnard J L, Bony A R, Bettini F, Campanaro A, Blerot B, Baudino S, Jullien F. 2018. Linalool and linalool nerolidol synthases in roses,several genes for little scent. Plant Physiology and Biochemistry,127:74-87.
[75]	Maimone T J, Baran P S. 2007. Modern synthetic efforts toward biologically active terpenes. Nature Chemical Biology, 3 (7):396-407.
[76]	Majetic C J, Wiggam S, Ferguson C J, Raguso R A. 2015. Timing is everything:temporal variation in floral scent,and its connections to pollinator behavior and female reproductive success in Phlox divaricata. American Midland Naturalist, 173 (2):191-207.
[77]	Majroomi S B, Abdollahi M B. 2018. The impact of cold stress on genes expression pattern of mono-and sesquiterpene biosynthesis and their contents in Ocimum basilicum L. Phytochemistry,156:250-256.
[78]	Mant J, Brändli C, Vereecken N J, Schulz C M, Francke W, Schiestl F P. 2005. Cuticular hydrocarbons as sex pheromone of the bee Colletes cunicularius and the key to its mimicry by the sexually deceptive orchidOphrys exaltata. Journal of Chemical Ecology, 31 (8):1765-1787.
[79]	Martin D M, Aubourg S, Schouwey M B, Daviet L, Schalk M, Toub O, Lund S T, Bohlmann J. 2010. Functional annotation,genome organization and phylogeny of the grapevine(Vitis vinifera)terpene synthase gene family based on genome assembly,FLcDNA cloning,and enzyme assays. BMC Plant Biology, 10 (1):226.
[80]	Martin D M, Fäldt J, Bohlmann J R. 2004. Functional characterization of nine Norway Spruce TPS genes and evolution of gymnosperm terpene synthases of the TPS-d subfamily. Plant Physiology, 135 (4):1908-1927.
[81]	Martin D M, Gershenzon J, Bohlmann J. 2003. Induction of volatile terpene biosynthesis and diurnal emission by methyl jasmonate in foliage of norway spruce. Plant Physiology, 132 (3):1586-1599.
[82]	Masumoto N, Korin M, Ito M. 2010. Geraniol and linalool synthases from wild species of perilla. Phytochemistry, 71 (10):1068-1075.
[83]	Maurya A K, Patel R C, Frost C J. 2020. Acute toxicity of the plant volatile indole depends on herbivore specialization. Journal of Pest Science, 93 (3):1107-1117.
[84]	Mei X, Liu X Y, Zhou Y, Wang X Q, Zeng L T, Fu X M, Li J L, Tang J C, Dong F, Yang Z Y. 2017. Formation and emission of linalool in tea(Camellia sinensis) leaves infested by tea green leafhopper(Empoasca(Matsumurasca)onukii Matsuda). Food Chemistry,237:356-363.
[85]	Mitra S, Karmakar A, Mukherjee A, Barik A. 2017. The role of leaf volatiles of Ludwigia octovalvis(Jacq.)raven in the attraction of Altica cyanea(Weber)(Coleoptera:Chrysomelidae). Journal of Chemical Ecology, 43 (7):679-692.
[86]	Miyake T, Yamaoka R, Yahara T. 1998. Floral scents of hawkmoth-pollinated flowers in Japan. Journal of Plant Research, 111 (2):199-205.
[87]	Mizutani M, Nakanishi H, Ema J I, Ma S J, Noguchi E, Inohara-Ochiai M, Fukuchi-Mizutani M, Nakao M, Sakata K. 2002. Cloning of β-primeverosidase from tea leaves,a key enzyme in tea aroma formation. Plant Physiology, 130 (4):2164-2176.
[88]	Nagegowda D A, Gutensohn M, Wilkerson C G, Dudareva N. 2008. Two nearly identical terpene synthases catalyze the formation of nerolidol and linalool in snapdragon flowers. Plant Journal, 55 (2):224-239.
[89]	Naskar S, Roy C, Ghosh S, Mukhopadhyay A, Hazarika L K, Chaudhuri R K, Roy S, Chakraborti D. 2021. Elicitation of biomolecules as host defense arsenals during insect attacks on tea plants[Camellia sinensis(L.)Kuntze]. Applied Microbiology and Biotechnologyl, 105 (19):7187-7199.
[90]	Nawade B, Shaltiel-Harpaz L, Yahyaa M, Kabaha A, Kedoshim R, Bosamia T C, Ibdah M. 2020. Characterization of terpene synthase genes potentially involved in black fig fly(Silba adipata)interactions with Ficus carica. Plant Science,298:110549.
[91]	Nieuwenhuizen N J, Green S A, Chen X, Bailleul E J D, Matich A J, Wang M Y, Atkinson R G. 2013. Functional genomics reveals that a compact terpene synthase gene family can account for terpene volatile production in apple. Plant Physiology, 161 (2):787-804.
[92]	Ochieng S, Park K, Baker T. 2002. Host plant volatiles synergize responses of sex pheromone-specific olfactory receptor neurons in male Helicoverpa zea. Journal of Comparative Physiology A, 188 (4):325-333.
[93]	Ohgami S, Ono E, Horikawa M, Murata J, Totsuka K, Toyonaga H, Ohba Y, Dohra H, Asai T, Matsui K, Mizutani M, Watanabe N, Ohnishi T. 2015. Volatile glycosylation in tea plants:sequential glycosylations for the biosynthesis of aroma β-primeverosides are catalyzed by two Camellia sinensis glycosyltransferases. Plant Physiology, 168 (2):464-477.
[94]	Özek T, Tabanca N, Demirci F, Wedge D E, Başer K H C. 2010. Enantiomeric distribution of some linalool containing essential oils and their biological activities. RNA Biology, 4 (1):22-30.
[95]	Paddon C J, Keasling J D. 2014. Semi-synthetic artemisinin:a model for the use of synthetic biology in pharmaceutical development. Nature Reviews Microbiology, 12 (5):355-367.
[96]	Pandey V, Tiwari D C, Dhyani V, Bhatt I D, Rawal R S, Nandi S K. 2021. Physiological and metabolic changes in two Himalayan medicinal herbs under drought,heat and combined stresses. Physiology and Molecular Biology of Plants, 27 (7):1523-1538.
[97]	Pei H R, Xie G, Yao X, Wang S H, Yan J, Dai L Y, Wang Y S. 2024. Exploring the binding affinity and characteristics of DcitOBP9 in citrus psyllids. Gene 9:923:148551.
[98]	Pichersky E, Raguso R A, Lewinsohn E, Croteau R. 1994. Floral scent production in Clarkia(Onagraceae)(I. Localization and Developmental Modulation of Monoterpene Emission and Linalool Synthase Activity). Plant Physiology, 106 (4):1533-1540.
[99]	Pragadheesh V S, Chanotiya C S, Rastogi S, Shasany A K. 2017. Scent from Jasminum grandiflorum flowers:investigation of the change in linalool enantiomers at various developmental stages using chemical and molecular methods. Phytochemistry,140:83-94.
[100]	Reddy A S N, Marquez Y, Kalyna M, Barta A. 2013. Complexity of the alternative splicing landscape in plants. Plant Cell, 25 (10):3657-3683.
[101]	Reis M, Pansarim E, Silva U, Amaral M d C, Amaral C, Marsaioli A. 2004. Pollinator attraction devices(floral fragrances)of some Brazilian orchids. Arkivoc,6:103-111.
[102]	Reisenman C E, Riffell J A, Bernays E A, Hildebrand J G. 2010. Antagonistic effects of floral scent in an insect-plant interaction. Proceedings. Biological Sciences, 277 (1692):2371-2379.
[103]	Rohmer M. 1999. The discovery of a mevalonate-independent pathway for isoprenoid biosynthesis in bacteria,algae and higher plants. Natural Product Reports, 16 (5):565-574.
[104]	Sarker L S, Adal A M, Mahmoud S S. 2019. Diverse transcription factors control monoterpene synthase expression in lavender(Lavandula). Planta, 251 (1):5.
[105]	Schiestl F P, Glaser F. 2012. Specific ant-pollination in an alpine orchid and the role of floral scent in attracting pollinating ants. Alpine Botany, 122 (1):1-9.
[106]	Shang J Z, Tian J P, Cheng H H, Yan Q M, Li L, Jamal A, Xu Z P, Xiang L, Saski C A, Jin S X, Zhao K G, Liu X Q, Chen L Q. 2020. The chromosome-level wintersweet(Chimonanthus praecox)genome provides insights into floral scent biosynthesis and flowering in winter. Genome Biology, 21 (1):200.
[107]	Shimada T, Endo T, Fujii H, Rodríguez A, Peña L, Omura M. 2014. Characterization of three linalool synthase genes from Citrus unshiu Marc. and analysis of linalool-mediated resistance against Xanthomonas citri subsp. citri and Penicilium italicum in citrus leaves and fruits. Plant Science,229:154-166.
[108]	Shimada T, Endo T, Rodríguez A, Fujii H, Goto S, Matsuura T, Hojo Y, Ikeda Y, Mori I C, Fujikawa T, Peña L, Omura M. 2017. Ectopic accumulation of linalool confers resistance to Xanthomonas citri subsp. citri in transgenic sweet orange plants. Tree Physiology, 37 (5):654-664.
[109]	Siani A C, Tappin M R R, Ramos M F S, Mazzei J L, Ramos M C K V, de Aquino Neto F R, Frighetto N. 2002. Linalool from Lippia alba:study of the reproducibility of the essential oil profile and the enantiomeric purity. Journal of Agricultural and Food Chemistry, 50 (12):3518-3521.
[110]	Soliman M M, Elsaba Y M, Soliman M S A, Ahmed E Z. 2024. Composition and antimicrobial activity of Rosmarinus officinalis L. and Artemisia monosperma L. leaf essential oils and methanolic extracts from plants grown in normal and saline habitats in Egypt. Scientific Reports, 14 (1):7342.
[111]	Steenhuisen S L, Jürgens A, Johnson S D. 2013. Effects of volatile compounds emitted by Protea species(Proteaceae)on antennal electrophysiological responses and attraction of cetoniine beetles. Journal of Chemical Ecology, 39 (3):438-446.
[112]	Sugiura M, Ito S, Saito Y, Niwa Y, Koltunow A M, Sugimoto O, Sakai H. 2014. Molecular cloning and characterization of a linalool synthase from lemon myrtle. Bioscience,Biotechnology,and Biochemistry, 75 (7):1245-1248.

科Family	物种Species	基因Gene	合成产物Synthetic product	参考文献Reference
杨柳科 Salicaceae	毛果杨 Populus trichocarpa	PtTPS3	（S）-（+）-芳樟醇和（S）-橙花叔醇（S）-（+）-linalool and （S）-nerolidol	Danner et al.,2011
杨柳科 Salicaceae	毛果杨 Populus trichocarpa	PtTPS4	主要产物：（R）-（−）-芳樟醇和（R）-橙花叔醇；次要产物：（S）-（+）-芳樟醇和（S）-橙花叔醇 Major products：（R）-（-）-linalool and（R）-nerolidol； minor products：（S）-（+）-linalool and（S）-nerolidol	Danner et al.,2011
禾本科 Poaceae	水稻 Oryza sativa	OsTPS3	（S）-（+）-芳樟醇（S）-（+）-linalool	Yuan et al.,2008
禾本科 Poaceae	玉米 Zea mays	ZmTPS1	芳樟醇和β-月桂烯 Linalool and β-myrcene	Yactayo-Chang et al.， 2024
毛茛科 Ranunculaceae	铁线莲 Clematis florida	CfTPS1、CfTPS2	芳樟醇和橙花叔醇 Linalool and nerolidol	Jiang et al.,2020
蔷薇科 Rosaceae	苹果 Malus domestica	MdLIS-RG1	（S）-（+）-芳樟醇（S）-（+）-linalool	Nieuwenhuizen et al.， 2013
	桃 Prunus persica	PpTPS1、PpTPS3	芳樟醇 Linalool	Liu et al.,2017b； Wei et al.,2021
	草莓 Fragaria × ananassa	FaNES1	（S）-（+）-芳樟醇和（E）-橙花叔醇（S）-（+）-linalool and（E）-nerolidol	Aharoni et al.,2004
	月季花 Rosa chinensis	RcLINS	（R）-（-）-芳樟醇（R）-（−）-linalool	Magnard et al.,2018； Ibdah et al.,2022
豆科 Fabaceae	合欢 Albizia julibrissin	AjTPS10	芳樟醇 Linalool	Liu et al.,2021
芸香科 Rutaceae	柑橘 Citrus unshiu	CuSTS3-1、CuSTS3-2	芳樟醇 Linalool	Shimada et al.,2014
芸香科 Rutaceae	柑橘 Citrus unshiu	CuSTS4	芳樟醇和橙花叔醇 Linalool and nerolidol	Shimada et al.,2014
唇形科 Lamiaceae	罗勒 Ocimum basilicum	ObLIS	（R）-（-）-芳樟醇（R）-（-）-linalool	Iijima et al.,2004
	柠檬薄荷 Mentha citrata	McLIS	（R）-（-）-芳樟醇（R）-（-）-linalool	Sugiura et al.,2014
	薰衣草 Lavandula augustifolia	LaLINS	（R）-（-）-芳樟醇（R）-（-）-linalool	Landmann et al.,2007
		LIS1、LIS2	芳樟醇 Linalool	龚林涛等,2021
	紫苏 Perilla hirtella Perilla setoyensis	PhTps-5042L、 PhTps-5073L、 PsTps-5031L	芳樟醇 Linalool	Masumoto et al.,2010
菊科 Asteraceae	黄花蒿 Artemisia annua	AaQH1、AaQH5	（R）-（-）-芳樟醇（R）-（-）-linalool	Jia et al.,1999
茄科 Solanaceae	番茄 Solanum lycopersicum	TPS5、TPS39	芳樟醇 Linalool	Cao et al.,2014
		TPS37、TPS39	芳樟醇和橙花叔醇 Linalool and nerolidol	Falara et al.,2011
		LeMTS1	（R）-（-）-芳樟醇（R）-（-）-linalool	van Schie et al.,2007
	烟草 Nicotiana attenuata	NaLIS	（S）-（+）-芳樟醇（S）-（+）-linalool	He et al.,2019
伞形科 Apiaceae	芫荽 Coriandrum sativum	CsLINS	（S）-（+）-芳樟醇（S）-（+）-linalool	Galata et al.,2014
葡萄科 Vitaceae	葡萄 Vitis vinifera	VvPNRLin	（R）-（-）-芳樟醇（R）-（-）-linalool	Martin et al.,2010
葡萄科 Vitaceae	葡萄 Vitis vinifera	VvPNLinNer1、 VvPNLinNer2、 VvCSLinNer	（S）-（+）-芳樟醇和（E）-橙花叔醇（S）-（+）-linalool and（E）-nerolidol	Martin et al.,2010
山茶科 Theaceae	茶 Camellia sinensis	CsLIS1、CsLIS2	（S）-（+）-芳樟醇（S）-（+）-linalool	Mei et al.,2017
		CsRLIS	（R）-（-）-芳樟醇（R）-（-）-linalool	Zhou et al.,2020
		CsLIS/NES-1	芳樟醇 Linalool	Liu et al.,2017a
锦葵科 Malvaceae	陆地棉 Gossypium hirsutum	GhTPS12、GhTPS14	（S）-（+）-芳樟醇（S）-（+）-linalool	Huang et al.,2017
		GhTPS4	（R）-（-）-芳樟醇（R）-（-）-linalool
		GhTPS10	芳樟醇 Linalool
十字花科 Brassicaceae	拟南芥 Arabidopsis thaliana	AtTPS14	（S）-（+）-芳樟醇（S）-（+）-linalool	Ginglinger et al.,2013
十字花科 Brassicaceae	拟南芥 Arabidopsis thaliana	AtTPS10	（R）-（-）-芳樟醇（R）-（-）-linalool	Ginglinger et al.,2013
木樨科 Oleaceae	木樨 Osmanthus fragrans	OfTPS1、 OfTPS2	（S）-（+）-芳樟醇（S）-（+）-linalool	Zeng et al.,2016
	木樨 Osmanthus fragrans	OfTPS5	芳樟醇 Linalool	刘偲等,2020
	素馨 Jasminum grandiflorum	LIS1	（R）-（-）-芳樟醇（R）-（-）-linalool	Pragadheesh et al.,2017
樟科 Lauraceae	土肉桂 Cinnamomum osmophloeum	CoLIS-D4、CoLIS-HS、 CoLIS-LL、CoLIS-T1	（S）-（+）-芳樟醇（S）-（+）-linalool	Lin et al.,2014
	香樟 Cinnamomum camphora	CcTPS54	芳樟醇 Linalool	Yang et al.,2023
		CcTPS3	芳樟醇 Linalool	马青等,2023
		CcTPS6	芳樟醇和橙花叔醇 Linalool and nerolidol	马青等,2023
鸢尾科 Iridaceae	香雪兰 Fressia × hybrida	FhTPS1	芳樟醇 Linalool	Gao et al.,2018
百合科 Liliaceae	百合 Lilium brownii	LiTPS2	芳樟醇和其他单萜、橙花叔醇和其他倍半萜 Linalool and other monoterpenes，nerolidol and other sesquiterpenes	Zhang et al.,2020
		LoTPS1	芳樟醇和（Z）-β-罗勒烯 Linalool and （Z）-β-ocimene	Abbas et al.,2018
		LoTPS3	芳樟醇和橙花叔醇 Linalool and nerolidol	Abbas et al.,2018
姜科 Zingiberaceae	姜花 Hedychium coronarium	HcTPS5	芳樟醇 Linalool	Yue et al.,2015； Ke et al.,2021
	姜花 Hedychium coronarium	HcTPS8	芳樟醇和倍半萜 Linalool and sesquiterpenes	Yue et al.,2014
	海南砂仁 Amomum longiligulare	AlTPS2	芳樟醇 Linalool	Zhao et al.,2021
	阳春砂仁 Amomum villosum	AvTPS2	芳樟醇 Linalool	Zhao et al.,2021
桑科 Moraceae	无花果 Ficus carica	FcTPS7 FcTPS8	芳樟醇 Linalool	Nawade et al.,2020
车前科 Plantaginaceae	金鱼草 Antirrhinum majus	AmNES/LIS-2	（S）-（+）-芳樟醇（S）-（+）-linalool	Nagegowda et al.,2008
柳叶菜科 Onagraceae	仙女扇 Clarkia breweri	LIS	（S）-（+）-芳樟醇（S）-（+）-linalool	Dudareva et al.,1996
腊梅科 Calycanthaceae	腊梅 Chimonanthus praecox	CpTPS4	芳樟醇 Linalool	Shang et al.,2020
猕猴桃科 Actinidiaceae	猕猴桃 Actinidia chinensis Actinidia arguta Actinidia polygama	AcLIS/NES	主要产物芳樟醇，次要产物橙花叔醇 Major products：linalool，minor products：nerolidol	Wang et al.,2023
猕猴桃科 Actinidiaceae		AaLS1、ApLS1	（S）-（+）-芳樟醇（S）-（+）-linalool	Chen et al.,2010； Günther et al.,2011
桃金娘科 Myrtaceae	柠檬香桃 Backhousia citriodora	BcLS	芳樟醇 Linalool	Sugiura et al.,2014
番木瓜科 Caricaceae	番木瓜 Carica papaya	CpTPS18	芳樟醇 Linalool	Yao et al.,2023b
檀香科 Santalaceae	檀香 Santalum album	SaNES/LIS	芳樟醇和橙花叔醇 Linalool and nerolidol	Zhang et al.,2021
菖蒲科 Acoraceae	菖蒲 Acorus calamus	AcTPS3、AcTPS4、 AcTPS5	芳樟醇和橙花叔醇 Linalool and nerolidol	Ibdah et al.,2022
兰科 Orchidaceae	铁皮石斛 Dendrobium officinale	DoTPS10	芳樟醇 Linalool	Yu et al.,2020
	蝴蝶兰 Phalaenopsis bellina	PbTPS4、PbTPS10	芳樟醇 Linalool	Huang et al.,2021
		PbTPS3	芳樟醇、β-罗勒烯 Linalool and β-ocimene
		PbTPS5	芳樟醇、香叶醇 Linalool and geraniol
芍药科 Paeoniaceae	牡丹 Paeonia delavayi Paeonia suffruticosa	PdTPS1、PdTPS4	芳樟醇 Linalool	Li et al.,2023
芍药科 Paeoniaceae	牡丹 Paeonia delavayi Paeonia suffruticosa	PsTPS14	芳樟醇 Linalool	王佩云等,2024
松科 Pinaceae	欧洲云杉 Picea abies	PaLIN	主要产物（R）-（-）-芳樟醇；次要产物（S）-（+）-芳樟醇 Major products：（R）-（-）-linalool； minor products：（S）-（+）-linalool	Martin et al.,2004
橄榄科 Burseraceae	乳香 Boswellia serrata	BsTPS2	（S）-（+）-芳樟醇（S）-（+）-linalool	Bhargav et al.,2023

科Family	物种Species	基因Gene	合成产物Synthetic product	参考文献Reference
杨柳科 Salicaceae	毛果杨 Populus trichocarpa	PtTPS3	（S）-（+）-芳樟醇和（S）-橙花叔醇（S）-（+）-linalool and （S）-nerolidol	Danner et al.,2011
杨柳科 Salicaceae	毛果杨 Populus trichocarpa	PtTPS4	主要产物：（R）-（−）-芳樟醇和（R）-橙花叔醇；次要产物：（S）-（+）-芳樟醇和（S）-橙花叔醇 Major products：（R）-（-）-linalool and（R）-nerolidol； minor products：（S）-（+）-linalool and（S）-nerolidol	Danner et al.,2011
禾本科 Poaceae	水稻 Oryza sativa	OsTPS3	（S）-（+）-芳樟醇（S）-（+）-linalool	Yuan et al.,2008
禾本科 Poaceae	玉米 Zea mays	ZmTPS1	芳樟醇和β-月桂烯 Linalool and β-myrcene	Yactayo-Chang et al.， 2024
毛茛科 Ranunculaceae	铁线莲 Clematis florida	CfTPS1、CfTPS2	芳樟醇和橙花叔醇 Linalool and nerolidol	Jiang et al.,2020
蔷薇科 Rosaceae	苹果 Malus domestica	MdLIS-RG1	（S）-（+）-芳樟醇（S）-（+）-linalool	Nieuwenhuizen et al.， 2013
	桃 Prunus persica	PpTPS1、PpTPS3	芳樟醇 Linalool	Liu et al.,2017b； Wei et al.,2021
	草莓 Fragaria × ananassa	FaNES1	（S）-（+）-芳樟醇和（E）-橙花叔醇（S）-（+）-linalool and（E）-nerolidol	Aharoni et al.,2004
	月季花 Rosa chinensis	RcLINS	（R）-（-）-芳樟醇（R）-（−）-linalool	Magnard et al.,2018； Ibdah et al.,2022
豆科 Fabaceae	合欢 Albizia julibrissin	AjTPS10	芳樟醇 Linalool	Liu et al.,2021
芸香科 Rutaceae	柑橘 Citrus unshiu	CuSTS3-1、CuSTS3-2	芳樟醇 Linalool	Shimada et al.,2014
芸香科 Rutaceae	柑橘 Citrus unshiu	CuSTS4	芳樟醇和橙花叔醇 Linalool and nerolidol	Shimada et al.,2014
唇形科 Lamiaceae	罗勒 Ocimum basilicum	ObLIS	（R）-（-）-芳樟醇（R）-（-）-linalool	Iijima et al.,2004
	柠檬薄荷 Mentha citrata	McLIS	（R）-（-）-芳樟醇（R）-（-）-linalool	Sugiura et al.,2014
	薰衣草 Lavandula augustifolia	LaLINS	（R）-（-）-芳樟醇（R）-（-）-linalool	Landmann et al.,2007
		LIS1、LIS2	芳樟醇 Linalool	龚林涛等,2021
	紫苏 Perilla hirtella Perilla setoyensis	PhTps-5042L、 PhTps-5073L、 PsTps-5031L	芳樟醇 Linalool	Masumoto et al.,2010
菊科 Asteraceae	黄花蒿 Artemisia annua	AaQH1、AaQH5	（R）-（-）-芳樟醇（R）-（-）-linalool	Jia et al.,1999
茄科 Solanaceae	番茄 Solanum lycopersicum	TPS5、TPS39	芳樟醇 Linalool	Cao et al.,2014
		TPS37、TPS39	芳樟醇和橙花叔醇 Linalool and nerolidol	Falara et al.,2011
		LeMTS1	（R）-（-）-芳樟醇（R）-（-）-linalool	van Schie et al.,2007
	烟草 Nicotiana attenuata	NaLIS	（S）-（+）-芳樟醇（S）-（+）-linalool	He et al.,2019
伞形科 Apiaceae	芫荽 Coriandrum sativum	CsLINS	（S）-（+）-芳樟醇（S）-（+）-linalool	Galata et al.,2014
葡萄科 Vitaceae	葡萄 Vitis vinifera	VvPNRLin	（R）-（-）-芳樟醇（R）-（-）-linalool	Martin et al.,2010
葡萄科 Vitaceae	葡萄 Vitis vinifera	VvPNLinNer1、 VvPNLinNer2、 VvCSLinNer	（S）-（+）-芳樟醇和（E）-橙花叔醇（S）-（+）-linalool and（E）-nerolidol	Martin et al.,2010
山茶科 Theaceae	茶 Camellia sinensis	CsLIS1、CsLIS2	（S）-（+）-芳樟醇（S）-（+）-linalool	Mei et al.,2017
		CsRLIS	（R）-（-）-芳樟醇（R）-（-）-linalool	Zhou et al.,2020
		CsLIS/NES-1	芳樟醇 Linalool	Liu et al.,2017a
锦葵科 Malvaceae	陆地棉 Gossypium hirsutum	GhTPS12、GhTPS14	（S）-（+）-芳樟醇（S）-（+）-linalool	Huang et al.,2017
		GhTPS4	（R）-（-）-芳樟醇（R）-（-）-linalool
		GhTPS10	芳樟醇 Linalool
十字花科 Brassicaceae	拟南芥 Arabidopsis thaliana	AtTPS14	（S）-（+）-芳樟醇（S）-（+）-linalool	Ginglinger et al.,2013
十字花科 Brassicaceae	拟南芥 Arabidopsis thaliana	AtTPS10	（R）-（-）-芳樟醇（R）-（-）-linalool	Ginglinger et al.,2013
木樨科 Oleaceae	木樨 Osmanthus fragrans	OfTPS1、 OfTPS2	（S）-（+）-芳樟醇（S）-（+）-linalool	Zeng et al.,2016
	木樨 Osmanthus fragrans	OfTPS5	芳樟醇 Linalool	刘偲等,2020
	素馨 Jasminum grandiflorum	LIS1	（R）-（-）-芳樟醇（R）-（-）-linalool	Pragadheesh et al.,2017
樟科 Lauraceae	土肉桂 Cinnamomum osmophloeum	CoLIS-D4、CoLIS-HS、 CoLIS-LL、CoLIS-T1	（S）-（+）-芳樟醇（S）-（+）-linalool	Lin et al.,2014
	香樟 Cinnamomum camphora	CcTPS54	芳樟醇 Linalool	Yang et al.,2023
		CcTPS3	芳樟醇 Linalool	马青等,2023
		CcTPS6	芳樟醇和橙花叔醇 Linalool and nerolidol	马青等,2023
鸢尾科 Iridaceae	香雪兰 Fressia × hybrida	FhTPS1	芳樟醇 Linalool	Gao et al.,2018
百合科 Liliaceae	百合 Lilium brownii	LiTPS2	芳樟醇和其他单萜、橙花叔醇和其他倍半萜 Linalool and other monoterpenes，nerolidol and other sesquiterpenes	Zhang et al.,2020
		LoTPS1	芳樟醇和（Z）-β-罗勒烯 Linalool and （Z）-β-ocimene	Abbas et al.,2018
		LoTPS3	芳樟醇和橙花叔醇 Linalool and nerolidol	Abbas et al.,2018
姜科 Zingiberaceae	姜花 Hedychium coronarium	HcTPS5	芳樟醇 Linalool	Yue et al.,2015； Ke et al.,2021
	姜花 Hedychium coronarium	HcTPS8	芳樟醇和倍半萜 Linalool and sesquiterpenes	Yue et al.,2014
	海南砂仁 Amomum longiligulare	AlTPS2	芳樟醇 Linalool	Zhao et al.,2021
	阳春砂仁 Amomum villosum	AvTPS2	芳樟醇 Linalool	Zhao et al.,2021
桑科 Moraceae	无花果 Ficus carica	FcTPS7 FcTPS8	芳樟醇 Linalool	Nawade et al.,2020
车前科 Plantaginaceae	金鱼草 Antirrhinum majus	AmNES/LIS-2	（S）-（+）-芳樟醇（S）-（+）-linalool	Nagegowda et al.,2008
柳叶菜科 Onagraceae	仙女扇 Clarkia breweri	LIS	（S）-（+）-芳樟醇（S）-（+）-linalool	Dudareva et al.,1996
腊梅科 Calycanthaceae	腊梅 Chimonanthus praecox	CpTPS4	芳樟醇 Linalool	Shang et al.,2020
猕猴桃科 Actinidiaceae	猕猴桃 Actinidia chinensis Actinidia arguta Actinidia polygama	AcLIS/NES	主要产物芳樟醇，次要产物橙花叔醇 Major products：linalool，minor products：nerolidol	Wang et al.,2023
猕猴桃科 Actinidiaceae		AaLS1、ApLS1	（S）-（+）-芳樟醇（S）-（+）-linalool	Chen et al.,2010； Günther et al.,2011
桃金娘科 Myrtaceae	柠檬香桃 Backhousia citriodora	BcLS	芳樟醇 Linalool	Sugiura et al.,2014
番木瓜科 Caricaceae	番木瓜 Carica papaya	CpTPS18	芳樟醇 Linalool	Yao et al.,2023b
檀香科 Santalaceae	檀香 Santalum album	SaNES/LIS	芳樟醇和橙花叔醇 Linalool and nerolidol	Zhang et al.,2021
菖蒲科 Acoraceae	菖蒲 Acorus calamus	AcTPS3、AcTPS4、 AcTPS5	芳樟醇和橙花叔醇 Linalool and nerolidol	Ibdah et al.,2022
兰科 Orchidaceae	铁皮石斛 Dendrobium officinale	DoTPS10	芳樟醇 Linalool	Yu et al.,2020
	蝴蝶兰 Phalaenopsis bellina	PbTPS4、PbTPS10	芳樟醇 Linalool	Huang et al.,2021
		PbTPS3	芳樟醇、β-罗勒烯 Linalool and β-ocimene
		PbTPS5	芳樟醇、香叶醇 Linalool and geraniol
芍药科 Paeoniaceae	牡丹 Paeonia delavayi Paeonia suffruticosa	PdTPS1、PdTPS4	芳樟醇 Linalool	Li et al.,2023
芍药科 Paeoniaceae	牡丹 Paeonia delavayi Paeonia suffruticosa	PsTPS14	芳樟醇 Linalool	王佩云等,2024
松科 Pinaceae	欧洲云杉 Picea abies	PaLIN	主要产物（R）-（-）-芳樟醇；次要产物（S）-（+）-芳樟醇 Major products：（R）-（-）-linalool； minor products：（S）-（+）-linalool	Martin et al.,2004
橄榄科 Burseraceae	乳香 Boswellia serrata	BsTPS2	（S）-（+）-芳樟醇（S）-（+）-linalool	Bhargav et al.,2023