脱落酸对‘京香玉’葡萄果实单萜物质合成的影响

doi:10.16420/j.issn.0513-353x.2021-1117

摘要/Abstract

摘要：

为探究脱落酸（Abscisic acid，ABA）对葡萄果实中单萜物质合成的影响，以玫瑰香型鲜食葡萄品种‘京香玉’为试材，在果实始熟前1周和始熟期用300 mg · L^-1 ABA浸泡果实2次，测定处理后7、14、21、28和35 d果实中游离态单萜化合物含量以及ABA合成和信号转导通路、单萜合成通路中相关基因的表达量。在果实中共检测到28种单萜类化合物，并随果实的发育成熟不断积累。外源ABA处理的成熟果实中游离态单萜含量比对照提高45.20%。通过实时荧光定量PCR发现，ABA合成基因（VvNCED1、VvGT）在处理后7 d表达量明显高于对照，VvβG1相对表达量在处理后持续上升；ABA信号受体基因VvPYL1、信号转导相关基因VvPP2C2、VvSnRK2.1、VvABI5的转录水平分别在处理后21和35 d明显高于对照；单萜合成通路上游关键酶基因（VvDXS2、VvDXS3、VvDXR、VvGPPS）以及单萜合酶基因VvPNLinNer的表达水平在处理后7 d明显高于对照。根据以上结果，推测外源ABA处理可增强果实内部ABA信号的转导，进而引起单萜合酶底物水平的提升，加快向单萜类化合物的转变，改善玫瑰香型葡萄的品质。

关键词: 葡萄, 玫瑰香型, 脱落酸, 单萜, 基因表达

Abstract:

In order to explore the effects of abscisic acid（ABA）on monoterpene synthesis in grape berries，this study used table grape species‘Jingxiangyu'with muscat flavor as experimental material，and soaked the clusters with 300 mg · L^-1 ABA twice at one week before and during veraison. The contents of free monoterpenoids at 7，14，21，28 and 35 days after first treatment were analyzed qualitatively and quantitatively by GC-MS. The results showed that 28 kinds of monoterpenes were detected in berries，most of which accumulated during ripening. The total contents of free monoterpenoids in treated berries at maturity was 45.20% higher than that in the control，and the substance that changed most strongly was linalool. To investigate how ABA regulate monoterpenes synthesis，quantitative real-time PCR was used. The results indicated that the expression of ABA synthesis genes（VvNCED1 and VvGT）was much stronger than that of the control. The relative expression of VvβG1 increased after ABA treatment constantly. The transcriptional levels of ABA receptor VvPYL1 and signal transduction related genes VvPP2C2，VvSnRK2.1 and VvABI5 were distinctly higher than the control on 21 and 35 days after treatment respectively. The expression levels of upstream genes about monoterpene synthesis（VvDXS2，VvDXS3，VvDXR，VvGPPS）and monoterpene synthase gene VvPNLinNer was significantly higher. According to the above results，it’s suggested that exogenous ABA enhance ABA signal transduction in berries，which lead to the increase of monoterpene substrate level and facilitate monoterpene synthesis，finally improve the quality of rose flavor grapes.

Key words: grape, muscat flavor, abscisic acid, monoterpenoids, gene expression

中图分类号:

S663.1

王晓晨, 聂子页, 刘先菊, 段伟, 范培格, 梁振昌. 脱落酸对‘京香玉’葡萄果实单萜物质合成的影响[J]. 园艺学报, 2023, 50(2): 237-249.

WANG Xiaochen, NIE Ziye, LIU Xianju, DUAN Wei, FAN Peige, LIANG Zhenchang. Effects of Abscisic Acid on Monoterpene Synthesis in‘Jingxiangyu’Grape Berries[J]. Acta Horticulturae Sinica, 2023, 50(2): 237-249.

图/表 5

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基因 Gene	基因ID Gene ID	正向引物（5′-3′） Forward primer	反向引物（5′-3′） Reverse primer
VvACTIN7	VIT_04s0044g00580	CTTGCATCCCTCAGCACCTT	TCCTGTGGACAATGGATGGA
VvNCED1	VIT_19s0093g00550	CCACACTCCCAAAAGAGAAGGA	CGGAGCAAAATTACCGGCG
VvGT	VIT_03s0063g00050	CGTCAACTGCTCATCCAACG	GGCAGAACAAAGACCTCCGA
VvβG1	VIT_01s0011g00760	AGGTGCTCCATCCCCCTATT	AGTGGCTTCAATGTCCTCTGT
VvPYL1	VIT_02s0012g01270	CTCAGCCGGAGTTCCAAGAG	GAGTAGGGAGGAGCATTGGC
VvPP2C2	VIT_04s0008g01420	CGTAGGAGATGGGAAGTGGC	TTGTTTGAGTCCCTCGGCAG
VvSnRK2.1	VIT_02s0236g00130	GGCAGCTCCTTTCCCGTATT	ATCCTTCCCCTGGTGTCCTT
VvSnRK2.6	VIT_03s0063g01080	CAGTGAGGATGAGGCACGTT	GGAGCAATGTATGCAGGGGT
VvABI5	VIT_08s0007g03420	GCAGGTCAGTTCGGTTTGGA	CTGCCAGAGCCTGTTGAAGA
VvABF2	VIT_18s0001g10450	CGGGATGGGAATGGTTGGTT	ACCTTCTCTACAGCCCCACT
VvDXS2	VIT_11s0052g01780	CTGCTGCCCAGGACAACAAT	CAGCCAACGTCTCAAGCTCC
VvDXS3	VIT_04s0008g04970	TTGAAAGGGAAACGGGAAC	TGGGTGTAAAGAATGACGACTG
VvDXR	VIT_17s0000g08390	TGCTGGGGGTCCTTTTGTCCTTCC	TCAACGGGCCAATCCCTGAATGC
VvHDR	VIT_03s0063g02030	CGTTATGTTAGTAGTTGGTGGGTG	CTTATTCTGTTTCCTGGACCTATTC
VvGPPS	VIT_15s0024g00850	ATGGTGGTTGCTGAGGTCC	CAACAATAGGACTGTGGGACG
VvPNLinNer1	VIT_00s0385g00010	AGATGGGATTTGTCTGCTTTCA	CTTATGCTCCTTGTGGACCTTG

基因 Gene	基因ID Gene ID	正向引物（5′-3′） Forward primer	反向引物（5′-3′） Reverse primer
VvACTIN7	VIT_04s0044g00580	CTTGCATCCCTCAGCACCTT	TCCTGTGGACAATGGATGGA
VvNCED1	VIT_19s0093g00550	CCACACTCCCAAAAGAGAAGGA	CGGAGCAAAATTACCGGCG
VvGT	VIT_03s0063g00050	CGTCAACTGCTCATCCAACG	GGCAGAACAAAGACCTCCGA
VvβG1	VIT_01s0011g00760	AGGTGCTCCATCCCCCTATT	AGTGGCTTCAATGTCCTCTGT
VvPYL1	VIT_02s0012g01270	CTCAGCCGGAGTTCCAAGAG	GAGTAGGGAGGAGCATTGGC
VvPP2C2	VIT_04s0008g01420	CGTAGGAGATGGGAAGTGGC	TTGTTTGAGTCCCTCGGCAG
VvSnRK2.1	VIT_02s0236g00130	GGCAGCTCCTTTCCCGTATT	ATCCTTCCCCTGGTGTCCTT
VvSnRK2.6	VIT_03s0063g01080	CAGTGAGGATGAGGCACGTT	GGAGCAATGTATGCAGGGGT
VvABI5	VIT_08s0007g03420	GCAGGTCAGTTCGGTTTGGA	CTGCCAGAGCCTGTTGAAGA
VvABF2	VIT_18s0001g10450	CGGGATGGGAATGGTTGGTT	ACCTTCTCTACAGCCCCACT
VvDXS2	VIT_11s0052g01780	CTGCTGCCCAGGACAACAAT	CAGCCAACGTCTCAAGCTCC
VvDXS3	VIT_04s0008g04970	TTGAAAGGGAAACGGGAAC	TGGGTGTAAAGAATGACGACTG
VvDXR	VIT_17s0000g08390	TGCTGGGGGTCCTTTTGTCCTTCC	TCAACGGGCCAATCCCTGAATGC
VvHDR	VIT_03s0063g02030	CGTTATGTTAGTAGTTGGTGGGTG	CTTATTCTGTTTCCTGGACCTATTC
VvGPPS	VIT_15s0024g00850	ATGGTGGTTGCTGAGGTCC	CAACAATAGGACTGTGGGACG
VvPNLinNer1	VIT_00s0385g00010	AGATGGGATTTGTCTGCTTTCA	CTTATGCTCCTTGTGGACCTTG

游离态单萜化合物 Free monoter-penes	处理后7 d 7 days after treatment		处理后14 d 14 days after treatment		处理后21 d 21 days after treatment
游离态单萜化合物 Free monoter-penes	对照 Control	ABA	对照 Control	ABA	对照 Control	ABA
游离态单萜物质总量 Total content of free monoterpenes	20.93 ± 3.75	7.96 ± 2.25^*	56.99 ± 1.31	26.26 ± 2.86^*	69.92 ± 2.02	80.56 ± 2.22^*
里那醇 Linalool	3.28 ± 0.75	1.37 ± 0.51^*	36.79 ± 2.22	14.40 ± 4.55^*	41.21 ± 3.07	52.43 ± 3.23^*
π 蒎烯 π Pinene	0.49 ± 0.28	0.15 ± 0.13	2.99 ± 0.11	1.28 ± 0.29^*	3.58 ± 0.34	5.47 ± 0.57^*
脱氢里那醇 Hotrienol	1.72 ± 0.51	0.57 ± 0.58	2.46 ± 0.55	1.54 ± 0.65	3.62 ± 0.41	4.64 ± 0.52^*
顺式里那醇氧化物 cis-Linalool Oxide	4.81 ± 2.17	0.79 ± 0.94^*	3.25 ± 0.07	2.07 ± 0.10^*	5.53 ± 0.14	3.04 ± 0.27^*
顺式罗勒烯 cis-Ocimene	0.25 ± 0.11	0.38 ± 0.05	0.76 ± 0.09	0.31 ± 0.08^*	1.07 ± 0.13	1.30 ± 0.11^*
右旋柠檬烯d-Limonene	0.71 ± 0.49	1.06 ± 0.10	1.23 ± 0.08	0.45 ± 0.14^*	1.33 ± 0.06	1.82 ± 0.18^*
别罗勒烯Allocimene B	0.06 ± 0.05	0 ± 0	0.40 ± 0.03	0.17 ± 0.03^*	0.53 ± 0.03	0.73 ± 0.08^*
右旋π蒎烯1R-πPinene	0.09 ± 0.05	0.18 ± 0.03	0.32 ± 0.04	0.13 ± 0.03^*	0.43 ± 0.06	0.57 ± 0.10
反式罗勒烯 trans-Ocimene	0.10 ± 0.06	0.17 ± 0.03	0.29 ± 0.06	0.13 ± 0.03^*	0.41 ± 0.05	0.59 ± 0.11^*
反式香叶醇trans-Geraniol	2.07 ± 1.05	0.60 ± 0.29	1.33 ± 0.39	1.34 ± 0.45	1.94 ± 0.44	1.23 ± 0.15^*
π水芹烯 π Phellandrene	Nd	Nd	0.35 ± 0.03	0.09 ± 0.07^*	0.44 ± 0.03	0.57 ± 0.04^*
顺式松油醇π Terpieol	0.08 ± 0.01	0.09 ± 0.02	0.25 ± 0.04	0.10 ± 0.07^*	0.25 ± 0.04	0.51 ± 0.07^*
3-蒈烯 3-Carene	Nd	Nd	0.28 ± 0.03	0.13 ± 0.04^*	0.39 ± 0.03	0.53 ± 0.12
侧柏烯2-Thujene	Nd	Nd	0.28 ± 0.03	0.13 ± 0.04^*	0.39 ± 0.03	0.53 ± 0.12
萜品油烯Terpinolene	Nd	Nd	0.20 ± 0.01	0.02 ± 0.05^*	0.25 ± 0.02	0.37 ± 0.02^*
顺式香叶醇cis-Geraniol	0.30 ± 0.14	0.10 ± 0.06	0.19 ± 0.04	0.17 ± 0.05	0.30 ± 0.05	0.26 ± 0.02
玫瑰醚 Rose oxide	1.18 ± 0.17	0.76 ± 0.23	0.58 ± 0.02	0.66 ± 0.19	0.69 ± 0.03	0.50 ± 0.02^*
顺式萜品醇 π Terpineol	0.14 ± 0.02	0.12 ± 0.04	0.24 ± 0.02	0.12 ± 0.02	0.22 ± 0.02	0.25 ± 0.01^*
环氧芳樟醇Epoxylinalol	0.17 ± 0.03	0.09 ± 0.03	0.40 ± 0.09	0.19 ± 0.01^*	0.25 ± 0.01	0.25 ± 0.02
莰酮 Camphor	0.02 ± 0.03	Nd	0.02 ± 0.03	0.05 ± 0.01	0.07 ± 0.01	0.07 ± 0.01
香茅醇Citronellol	0.17 ± 0.00	0.10 ± 0.03	0.12 ± 0.04	0.11 ± 0.03	0.15 ± 0.07	0.16 ± 0.10
香茅醛 Citral	0.10 ± 0.01	0.06 ± 0.01^*	0.07 ± 0.01	0.06 ± 0.01	0.08 ± 0.01	0.08 ± 0.01
左旋薄荷醇l-(-)-Menthol	0.11 ± 0.04	0.12 ± 0.04	0.12 ± 0.02	0.12 ± 0.02	0.15 ± 0.03	0.17 ± 0.02
左旋4-萜品醇 l-4-terpineol	Nd	Nd	Nd	Nd	Nd	Nd
薄荷醇 Menthol	0.04 ± 0.03	0.04 ± 0.04	0.05 ± 0	0.05 ± 0.01	0.06 ± 0.01	0.07 ± 0.01
异香叶醇Isogeraniol	Nd	0.03 ± 0.05	Nd	0.05 ± 0.04	Nd	Nd
游离态单萜化合物 Free monoter-penes	处理后28 d 28 days after treatment		处理后35 d 35 days after treatment
游离态单萜化合物 Free monoter-penes	对照 Control	ABA	对照 Control	ABA
游离态单萜物质总量 Total contents of free monoterpenes	153.64 ± 4.23	194.23 ± 3.74^**	214.71 ± 8.01	311.74 ± 12.72^**
里那醇 Linalool	98.29 ± 4.56	122.89 ± 4.39^*	149.48 ± 11.88	206.58 ± 10.64^*
π 蒎烯 π Pinene	9.69 ± 1.12	12.83 ± 0.71^*	14.48 ± 1.05	24.36 ± 2.66^*
脱氢里那醇 Hotrienol	7.11 ± 0.84	13.82 ± 0.97^*	6.59 ± 1.47	12.96 ± 2.19^*
顺式里那醇氧化物 cis-Linalool Oxide	6.92 ± 0.66	7.43 ± 0.24	5.49 ± 0.55	8.38 ± 0.54^*
顺式罗勒烯 cis-Ocimene	3.14 ± 0.45	4.02 ± 0.22^*	4.62 ± 0.23	8.37 ± 1.10^*
右旋柠檬烯d-Limonene	2.96 ± 0.26	3.67 ± 0.16^*	4.41 ± 0.30	6.99 ± 0.92^*
别罗勒烯Allocimene B	1.32 ± 0.21	1.75 ± 0.08^*	2.08 ± 0.21	3.55 ± 0.52^*
右旋π蒎烯1R-π Pinene	1.11 ± 0.10	1.52 ± 0.13^*	2.07 ± 0.10	3.51 ± 0.58^*
反式罗勒烯trans-Ocimene	1.15 ± 0.10	1.53 ± 0.16^*	2.06 ± 0.08	3.50 ± 0.58^*
反式香叶醇trans-Geraniol	4.83 ± 0.39	3.70 ± 0.70^*	3.88 ± 1.05	2.40 ± 2.03
π水芹烯 π Phellandrene	0.90 ± 0.30	1.23 ± 0.05^*	1.46 ± 0.13	2.33 ± 0.29^*
顺式松油醇 π Terpieol	0.67 ± 0.11	1.34 ± 0.18^*	0.95 ± 0.28	2.13 ± 0.42^*
3-蒈烯3-Carene	0.87 ± 0.11	1.03 ± 0.04^*	1.40 ± 0.08	2.03 ± 0.17^*
侧柏烯2-Thujene	0.86 ± 0.10	1.01 ± 0.04^*	1.32 ± 0.13	1.92 ± 0.17^*
萜品油烯Terpinolene	0.55 ± 0.10	0.86 ± 0.06^*	0.86 ± 0.13	1.52 ± 0.22^*
顺式香叶醇cis-Geraniol	0.90 ± 0.10	0.97 ± 0.18	0.82 ± 0.21	1.20 ± 0.26
玫瑰醚Rose oxide	1.01 ± 0.03	1.11 ± 0.03^*	0.72 ± 0.01	1.04 ± 0.07^*
顺式萜品醇 π Terpineol	0.43 ± 0.03	0.54 ± 0.01^*	0.60 ± 0.05	0.92 ± 0.16^*
环氧芳樟醇Epoxylinalol	0.43 ± 0.07	0.49 ± 0.17	0.48 ± 0.07	0.66 ± 0.13
莰酮 Camphor	0.10 ± 0.03	0.25 ± 0.02^*	0.10 ± 0.03	0.25 ± 0.08^*
香茅醇Citronellol	0.24 ± 0.02	0.44 ± 0.21	0.35 ± 0.25	0.22 ± 0.14
香茅醛Citral	0.16 ± 0.03	0.14 ± 0.02	0.12 ± 0.03	0.15 ± 0.02
左旋薄荷醇l-(-)-Menthol	0.13 ± 0.01	0.15 ± 0.01^*	0.13 ± 0.01	0.13 ± 0.02
左旋4-萜品醇 l-4-terpineol	Nd	Nd	0.02 ± 0.03	0.09 ± 0.02^*
薄荷醇Menthol	0.04 ± 0.03	0.05 ± 0.03	0.06 ± 0.00	0.04 ± 0.03
异香叶醇Isogeraniol	Nd	Nd	Nd	Nd

游离态单萜化合物 Free monoter-penes	处理后7 d 7 days after treatment		处理后14 d 14 days after treatment		处理后21 d 21 days after treatment
游离态单萜化合物 Free monoter-penes	对照 Control	ABA	对照 Control	ABA	对照 Control	ABA
游离态单萜物质总量 Total content of free monoterpenes	20.93 ± 3.75	7.96 ± 2.25^*	56.99 ± 1.31	26.26 ± 2.86^*	69.92 ± 2.02	80.56 ± 2.22^*
里那醇 Linalool	3.28 ± 0.75	1.37 ± 0.51^*	36.79 ± 2.22	14.40 ± 4.55^*	41.21 ± 3.07	52.43 ± 3.23^*
π 蒎烯 π Pinene	0.49 ± 0.28	0.15 ± 0.13	2.99 ± 0.11	1.28 ± 0.29^*	3.58 ± 0.34	5.47 ± 0.57^*
脱氢里那醇 Hotrienol	1.72 ± 0.51	0.57 ± 0.58	2.46 ± 0.55	1.54 ± 0.65	3.62 ± 0.41	4.64 ± 0.52^*
顺式里那醇氧化物 cis-Linalool Oxide	4.81 ± 2.17	0.79 ± 0.94^*	3.25 ± 0.07	2.07 ± 0.10^*	5.53 ± 0.14	3.04 ± 0.27^*
顺式罗勒烯 cis-Ocimene	0.25 ± 0.11	0.38 ± 0.05	0.76 ± 0.09	0.31 ± 0.08^*	1.07 ± 0.13	1.30 ± 0.11^*
右旋柠檬烯d-Limonene	0.71 ± 0.49	1.06 ± 0.10	1.23 ± 0.08	0.45 ± 0.14^*	1.33 ± 0.06	1.82 ± 0.18^*
别罗勒烯Allocimene B	0.06 ± 0.05	0 ± 0	0.40 ± 0.03	0.17 ± 0.03^*	0.53 ± 0.03	0.73 ± 0.08^*
右旋π蒎烯1R-πPinene	0.09 ± 0.05	0.18 ± 0.03	0.32 ± 0.04	0.13 ± 0.03^*	0.43 ± 0.06	0.57 ± 0.10
反式罗勒烯 trans-Ocimene	0.10 ± 0.06	0.17 ± 0.03	0.29 ± 0.06	0.13 ± 0.03^*	0.41 ± 0.05	0.59 ± 0.11^*
反式香叶醇trans-Geraniol	2.07 ± 1.05	0.60 ± 0.29	1.33 ± 0.39	1.34 ± 0.45	1.94 ± 0.44	1.23 ± 0.15^*
π水芹烯 π Phellandrene	Nd	Nd	0.35 ± 0.03	0.09 ± 0.07^*	0.44 ± 0.03	0.57 ± 0.04^*
顺式松油醇π Terpieol	0.08 ± 0.01	0.09 ± 0.02	0.25 ± 0.04	0.10 ± 0.07^*	0.25 ± 0.04	0.51 ± 0.07^*
3-蒈烯 3-Carene	Nd	Nd	0.28 ± 0.03	0.13 ± 0.04^*	0.39 ± 0.03	0.53 ± 0.12
侧柏烯2-Thujene	Nd	Nd	0.28 ± 0.03	0.13 ± 0.04^*	0.39 ± 0.03	0.53 ± 0.12
萜品油烯Terpinolene	Nd	Nd	0.20 ± 0.01	0.02 ± 0.05^*	0.25 ± 0.02	0.37 ± 0.02^*
顺式香叶醇cis-Geraniol	0.30 ± 0.14	0.10 ± 0.06	0.19 ± 0.04	0.17 ± 0.05	0.30 ± 0.05	0.26 ± 0.02
玫瑰醚 Rose oxide	1.18 ± 0.17	0.76 ± 0.23	0.58 ± 0.02	0.66 ± 0.19	0.69 ± 0.03	0.50 ± 0.02^*
顺式萜品醇 π Terpineol	0.14 ± 0.02	0.12 ± 0.04	0.24 ± 0.02	0.12 ± 0.02	0.22 ± 0.02	0.25 ± 0.01^*
环氧芳樟醇Epoxylinalol	0.17 ± 0.03	0.09 ± 0.03	0.40 ± 0.09	0.19 ± 0.01^*	0.25 ± 0.01	0.25 ± 0.02
莰酮 Camphor	0.02 ± 0.03	Nd	0.02 ± 0.03	0.05 ± 0.01	0.07 ± 0.01	0.07 ± 0.01
香茅醇Citronellol	0.17 ± 0.00	0.10 ± 0.03	0.12 ± 0.04	0.11 ± 0.03	0.15 ± 0.07	0.16 ± 0.10
香茅醛 Citral	0.10 ± 0.01	0.06 ± 0.01^*	0.07 ± 0.01	0.06 ± 0.01	0.08 ± 0.01	0.08 ± 0.01
左旋薄荷醇l-(-)-Menthol	0.11 ± 0.04	0.12 ± 0.04	0.12 ± 0.02	0.12 ± 0.02	0.15 ± 0.03	0.17 ± 0.02
左旋4-萜品醇 l-4-terpineol	Nd	Nd	Nd	Nd	Nd	Nd
薄荷醇 Menthol	0.04 ± 0.03	0.04 ± 0.04	0.05 ± 0	0.05 ± 0.01	0.06 ± 0.01	0.07 ± 0.01
异香叶醇Isogeraniol	Nd	0.03 ± 0.05	Nd	0.05 ± 0.04	Nd	Nd
游离态单萜化合物 Free monoter-penes	处理后28 d 28 days after treatment		处理后35 d 35 days after treatment
游离态单萜化合物 Free monoter-penes	对照 Control	ABA	对照 Control	ABA
游离态单萜物质总量 Total contents of free monoterpenes	153.64 ± 4.23	194.23 ± 3.74^**	214.71 ± 8.01	311.74 ± 12.72^**
里那醇 Linalool	98.29 ± 4.56	122.89 ± 4.39^*	149.48 ± 11.88	206.58 ± 10.64^*
π 蒎烯 π Pinene	9.69 ± 1.12	12.83 ± 0.71^*	14.48 ± 1.05	24.36 ± 2.66^*
脱氢里那醇 Hotrienol	7.11 ± 0.84	13.82 ± 0.97^*	6.59 ± 1.47	12.96 ± 2.19^*
顺式里那醇氧化物 cis-Linalool Oxide	6.92 ± 0.66	7.43 ± 0.24	5.49 ± 0.55	8.38 ± 0.54^*
顺式罗勒烯 cis-Ocimene	3.14 ± 0.45	4.02 ± 0.22^*	4.62 ± 0.23	8.37 ± 1.10^*
右旋柠檬烯d-Limonene	2.96 ± 0.26	3.67 ± 0.16^*	4.41 ± 0.30	6.99 ± 0.92^*
别罗勒烯Allocimene B	1.32 ± 0.21	1.75 ± 0.08^*	2.08 ± 0.21	3.55 ± 0.52^*
右旋π蒎烯1R-π Pinene	1.11 ± 0.10	1.52 ± 0.13^*	2.07 ± 0.10	3.51 ± 0.58^*
反式罗勒烯trans-Ocimene	1.15 ± 0.10	1.53 ± 0.16^*	2.06 ± 0.08	3.50 ± 0.58^*
反式香叶醇trans-Geraniol	4.83 ± 0.39	3.70 ± 0.70^*	3.88 ± 1.05	2.40 ± 2.03
π水芹烯 π Phellandrene	0.90 ± 0.30	1.23 ± 0.05^*	1.46 ± 0.13	2.33 ± 0.29^*
顺式松油醇 π Terpieol	0.67 ± 0.11	1.34 ± 0.18^*	0.95 ± 0.28	2.13 ± 0.42^*
3-蒈烯3-Carene	0.87 ± 0.11	1.03 ± 0.04^*	1.40 ± 0.08	2.03 ± 0.17^*
侧柏烯2-Thujene	0.86 ± 0.10	1.01 ± 0.04^*	1.32 ± 0.13	1.92 ± 0.17^*
萜品油烯Terpinolene	0.55 ± 0.10	0.86 ± 0.06^*	0.86 ± 0.13	1.52 ± 0.22^*
顺式香叶醇cis-Geraniol	0.90 ± 0.10	0.97 ± 0.18	0.82 ± 0.21	1.20 ± 0.26
玫瑰醚Rose oxide	1.01 ± 0.03	1.11 ± 0.03^*	0.72 ± 0.01	1.04 ± 0.07^*
顺式萜品醇 π Terpineol	0.43 ± 0.03	0.54 ± 0.01^*	0.60 ± 0.05	0.92 ± 0.16^*
环氧芳樟醇Epoxylinalol	0.43 ± 0.07	0.49 ± 0.17	0.48 ± 0.07	0.66 ± 0.13
莰酮 Camphor	0.10 ± 0.03	0.25 ± 0.02^*	0.10 ± 0.03	0.25 ± 0.08^*
香茅醇Citronellol	0.24 ± 0.02	0.44 ± 0.21	0.35 ± 0.25	0.22 ± 0.14
香茅醛Citral	0.16 ± 0.03	0.14 ± 0.02	0.12 ± 0.03	0.15 ± 0.02
左旋薄荷醇l-(-)-Menthol	0.13 ± 0.01	0.15 ± 0.01^*	0.13 ± 0.01	0.13 ± 0.02
左旋4-萜品醇 l-4-terpineol	Nd	Nd	0.02 ± 0.03	0.09 ± 0.02^*
薄荷醇Menthol	0.04 ± 0.03	0.05 ± 0.03	0.06 ± 0.00	0.04 ± 0.03
异香叶醇Isogeraniol	Nd	Nd	Nd	Nd