基于代谢组与转录组联合分析解析秋石斛花香形成机制

doi:10.16420/j.issn.0513-353x.2025-1050

园艺学报 ›› 2026, Vol. 53 ›› Issue (2): 557-573.doi: 10.16420/j.issn.0513-353x.2025-1050

基于代谢组与转录组联合分析解析秋石斛花香形成机制

李崇晖¹^,^*(), 洪小雨¹^,³, 陆顺教¹, 廖易¹, 尹俊梅¹^,²^,^*()

¹ 中国热带农业科学院热带作物品种资源研究所，农业农村部华南作物基因资源与种质创制重点实验室，海南省热带作物资源遗传改良与创新重点实验室，海南省热带观赏植物种质创新利用工程技术研究中心，海口 571101
² 中国热带农业科学院三亚研究院，海南三亚 571101
³ 华中农业大学园艺林学学院，武汉 430070

收稿日期:2025-10-29 修回日期:2026-01-12 出版日期:2026-02-25 发布日期:2026-02-12
基金资助:
中央级公益性科研院所基本科研业务费专项(1630032022002); 海南省科技人才创新项目(KJRC2023C19); 现代农业产业技术体系建设专项资助(CARS-23-G60)

Elucidation of Floral Scent Formation in Phalaenopsis-Type Dendrobium Hybrids by Integrated Metabolome and Transcriptome Analysis

LI Chonghui¹(), HONG Xiaoyu¹^,³, LU Shunjiao¹, LIAO Yi¹, YIN Junmei¹^,²()

¹ Tropical Crop Germplasm Research Institute，Chinese Academy of Tropical Agricultural Sciences，Key Laboratory of Crop Gene Resources and Germplasm Enhancement in Southern China，Ministry of Agriculture and Rural Affairs，Key Laboratory of Tropical Crops Germplasm Resources Genetic Improvement and Innovation of Hainan Province，The Engineering Technology Research Center of Tropical Ornamental Plant Germplasm Innovation and Utilization，Haikou 571101，China
² Sanya Research Institute，Chinese Academy of Tropical Agricultural Sciences，Sanya，Hainan 571101，China
³ School of Horticulture and Forestry，Huazhong Agricultural University，Wuhan 430070，China

Received:2025-10-29 Revised:2026-01-12 Published:2026-02-25 Online:2026-02-12

摘要/Abstract

摘要：

秋石斛是重要的热带兰切花和盆花产品，但大多商业品种缺乏香气。以秋石斛芳香品种‘红珍珠’和非芳香品种‘迷你’为试验材料进行代谢组学和转录组研究，根据加权基因共表达网络分析和两组学联合分析筛选差异表达基因，并采用实时荧光定量PCR验证关键基因表达模式。结果表明，在鉴定的挥发性代谢物中，萜类化合物种类（159种）和含量最为丰富。相对气味活度值（rOAV）分析表明芳樟醇、3,6-壬二烯醛、2-戊基呋喃等化合物对‘红珍珠’花香贡献显著，这些化合物的香气被描述为花香、草香、果香。‘红珍珠’盛花期特异积累芳樟醇、茶螺烷等单萜类物质，而‘迷你’则以倍半萜类为主，因此，特定的单萜而非萜类总含量与‘红珍珠’的愉悦香气密切相关。秋石斛花香形成伴随着萜类生物合成通路基因的协同上调表达。筛选出11个花香核心候选基因，编码萜类合酶、S-芳樟醇合酶、倍半萜合酶等。这些基因在‘迷你’中表达量极低，在‘红珍珠’中的表达量随花发育显著上调，在盛花期最高，且与关键萜类代谢物积累显著正相关；RT-qPCR验证结果与转录组数据一致。关键萜类合成候选基因在花发育后期特异高表达是芳香秋石斛香气形成的重要分子机制。

关键词: 秋石斛, 花香, 萜类化合物, 芳樟醇, 茶螺烷

Abstract:

Phalaenopsis-Type Dendrobium hybrids（Den-Phals）have high ornamental value，but most cultivars lack floral fragrance. Using the aromatic cultivar D.‘Udomsri Beauty’（HZZ）and the non-aromatic cultivar D. Yaya Victoria（MN）as experimental materials，metabolomics analysis and transcriptome sequencing were conducted. Weighted Gene Co-expression Network Analysis and integrated analysis of both omics datasets were performed to screen differentially expressed genes, and real-time quantitative PCR（RT-qPCR）was used to validate the expression patterns of key genes. The results showed that among the identified volatile metabolites，terpenoids were the most abundant in both variety（159 kinds）and content. Relative odor activity value（rOAV）analysis identified compounds such as linalool，3,6-nonadienal，and 2-pentylfuran，were described as having floral，green，and fruity scent characteristics-as significant contributors to the floral fragrance of HZZ. The fragrant HZZ specifically accumulated monoterpenes like linalool and theaspirane during full-bloom stage, whereas the non-fragrant MN primarily accumulated sesquiterpenes. Specific monoterpenes，rather than total terpene content，are closely associated with the pleasant fragrance of Den-Phals. The formation of fragrance in Den-Phals was accompanied by coordinated upregulation of genes in the terpenoid biosynthesis pathway. Further analysis identified 11 core candidate genes（e.g.，genes encoding terpene synthase，S-linalool synthase，sesquiterpene synthase，etc.）with very low expression in MN，while their expression in HZZ significantly increased during flower development，peaking at full-bloom stage，and showed a significant positive correlation with the accumulation of key terpenoid metabolites. RT-qPCR validation confirmed consistency with the transcriptome data. The specific high expression of key terpenoid synthesis candidate genes during the late developmental stages of the fragrant cultivar is an important molecular mechanism underlying fragrance formation in Den-Phals.

Key words: Phalaenopsis-Type Dendrobium hybrids, floral scent, terpenoids, linalool, theaspirane

李崇晖, 洪小雨, 陆顺教, 廖易, 尹俊梅. 基于代谢组与转录组联合分析解析秋石斛花香形成机制[J]. 园艺学报, 2026, 53(2): 557-573.

LI Chonghui, HONG Xiaoyu, LU Shunjiao, LIAO Yi, YIN Junmei. Elucidation of Floral Scent Formation in Phalaenopsis-Type Dendrobium Hybrids by Integrated Metabolome and Transcriptome Analysis[J]. Acta Horticulturae Sinica, 2026, 53(2): 557-573.

图/表 13

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基因ID号/基因名 Gene ID/Gene name	引物正向序列（5′-3′） Forward primer sequence	引物反向序列（5′-3′） Reverse primer sequence
CL7532.Contig2_All	CTTGAGCGATTCACCGATGT	AATGTCCAAGCCCTTCTCCT
Unigene66_All	TTGTTGATCACTTGGAGAGG	AGCCATGCATCCTCAATAAT
CL105.Contig16_All	GAAAGAGGGTGGTGACGAA	TCAATGGCGAGTTGGAGGAG
Unigene78_All	TCAAGTTTCTCCAACGAGGTTT	CATTCCAGCCTTCTCAAGTTCA
CL55.Contig2_All	CTTCATAGACCACACGCCAAT	ATCGCTGTTCAGAACTTGACAT
Unigene1_All	TTGAAGCGGCATATAAGTGATG	GAAGTGACGACGGTGACAA
Unigene6_All	GAAGTGACGACGGTGACAA	TTGAAGCGGCATATAAGTGATG
β-Actin	GTCAGGGACATCAAGGAGAAG	TGGGCACCTAAATCTCTCAGC
TUA	CAAAGAAGATGCAGCCAACAAC	AAGACCAGTGCAGTTGTCAGCTA

基因ID号/基因名 Gene ID/Gene name	引物正向序列（5′-3′） Forward primer sequence	引物反向序列（5′-3′） Reverse primer sequence
CL7532.Contig2_All	CTTGAGCGATTCACCGATGT	AATGTCCAAGCCCTTCTCCT
Unigene66_All	TTGTTGATCACTTGGAGAGG	AGCCATGCATCCTCAATAAT
CL105.Contig16_All	GAAAGAGGGTGGTGACGAA	TCAATGGCGAGTTGGAGGAG
Unigene78_All	TCAAGTTTCTCCAACGAGGTTT	CATTCCAGCCTTCTCAAGTTCA
CL55.Contig2_All	CTTCATAGACCACACGCCAAT	ATCGCTGTTCAGAACTTGACAT
Unigene1_All	TTGAAGCGGCATATAAGTGATG	GAAGTGACGACGGTGACAA
Unigene6_All	GAAGTGACGACGGTGACAA	TTGAAGCGGCATATAAGTGATG
β-Actin	GTCAGGGACATCAAGGAGAAG	TGGGCACCTAAATCTCTCAGC
TUA	CAAAGAAGATGCAGCCAACAAC	AAGACCAGTGCAGTTGTCAGCTA

分类 Class	阈值/ （μg · g^-1） Threshold	相对含量/（μg · g^-1FW） Relative content		rOAV		香味描述 Aroma description
分类 Class	阈值/ （μg · g^-1） Threshold	红珍珠 D.‘Udomsri Bwauty’	迷你 D.‘Yaya Victoria’	红珍珠 D. ‘Udomsri Bwauty’	迷你 D.‘Yaya Victoria’	香味描述 Aroma description
醛Aldehyde
3,6-壬二烯醛 (Z,Z)-3,6-Nonadienal	5.00E-05	0.577 ± 0.006	0.197 ± 0.019	11 531.56 ± 110.58	3 937.34 ± 378.01	油腻、肥皂、黄瓜 Fatty，soapy，cucumber
(E,E)-2,4-十一烯醛 (E,E)-2,4-Undecadienal	1.00E-03	0.093 ± 0.060	1.471 ± 0.118	92.72 ± 5.57	1 470.58 ± 118.01	油腻、焦糖味、辛辣、柑橘、奶油、烘焙 Oily，caramel，spicy，citrus，buttery，baked
醛Aldehyde
反,顺-2,6-壬二烯醛 (E,Z)-2,6-Nonadienal	1.00E-05	0.032 ± 0.032	0.096 ± 0.014	3 186.86 ± 2 614.20	9 577.90 ± 960.95	黄瓜、青草 Cucumber，green
杂环化合物Heterocyclic compound
2-乙酰基噻吩 1-(2-thienyl)-Ethanone	1.00E-03	0.153 ± 0.002	0.037 ± 0.003	153.05 ± 1.53	36.64 ± 2.69	硫磺、坚果、榛子、核桃 Sulfury，nutty，hazelnut，walnut
2-乙基-3,5-二甲基吡嗪 2-ethyl-3,5-dimethyl-Pyrazine	4.00E-05	0.011 ± 0.000	0.001 ± 0.000	273.62 ± 5.10	20.44 ± 2.16	灼烧、杏仁、烤、坚果、咖啡 Burnt，almond，roasted，nutty，coffee
2-戊基呋喃 2-pentyl-Furan	6.00E-03	0.699 ± 0.023	0.258 ± 0.100	116.56 ± 3.81	43.01 ± 16.72	果香、青草、泥土、豆类、蔬菜、金属 Fruity，green，earthy，Beany，vegetable，metallic
萜类Terpenoids
(+)-柏木脑Cedrol	5.00E-04	0.013 ± 0.001	0.237 ± 0.027	26.92 ± 2.27	474.35 ± 53.95	雪松木、木香、干燥、甜 Cedarwood，woody，dry，sweet
芳樟醇Linalool	6.00E-03	0.777 ± 0.018	0.184 ± 0.016	129.57 ± 3.02	30.69 ± 2.74	花香、青草Floral，green
酯Ester
苯甲酸甲酯 Methyl ester benzoic acid	5.20E-04	0.202 ± 0.043	0.039 ± 0.004	389.31 ± 82.70	74.49 ± 7.64	苯酚、冬青油、杏仁、花香 Phenol，wintergreen，almond，floral，
含硫化合物Sulfur compounds
苯甲硫醇 Benzenemethanethiol	3.50E-06	0.374 ± 0.003	0.051 ± 0.004	106 787.09 ± 938.44	14 608.38 ± 1 056.60	尖锐、葱蒜、硫磺、辣根、薄荷、咖啡Sharp，alliaceous，onion，sulfury，garlic，horseradish，minty，coffee
芳烃Aromatics
二甲萘烷醇 Octahydro-4,8a-dimethyl-4a (2H)-Naphthalenol	2.10E-04	0.003 ± 0.001	0.026 ± 0.003	16.39 ± 2.92	125.25 ± 16.22	新鲜、霉味、土、土壤 Fresh，musty，earthy，soil

分类 Class	阈值/ （μg · g^-1） Threshold	相对含量/（μg · g^-1FW） Relative content		rOAV		香味描述 Aroma description
分类 Class	阈值/ （μg · g^-1） Threshold	红珍珠 D.‘Udomsri Bwauty’	迷你 D.‘Yaya Victoria’	红珍珠 D. ‘Udomsri Bwauty’	迷你 D.‘Yaya Victoria’	香味描述 Aroma description
醛Aldehyde
3,6-壬二烯醛 (Z,Z)-3,6-Nonadienal	5.00E-05	0.577 ± 0.006	0.197 ± 0.019	11 531.56 ± 110.58	3 937.34 ± 378.01	油腻、肥皂、黄瓜 Fatty，soapy，cucumber
(E,E)-2,4-十一烯醛 (E,E)-2,4-Undecadienal	1.00E-03	0.093 ± 0.060	1.471 ± 0.118	92.72 ± 5.57	1 470.58 ± 118.01	油腻、焦糖味、辛辣、柑橘、奶油、烘焙 Oily，caramel，spicy，citrus，buttery，baked
醛Aldehyde
反,顺-2,6-壬二烯醛 (E,Z)-2,6-Nonadienal	1.00E-05	0.032 ± 0.032	0.096 ± 0.014	3 186.86 ± 2 614.20	9 577.90 ± 960.95	黄瓜、青草 Cucumber，green
杂环化合物Heterocyclic compound
2-乙酰基噻吩 1-(2-thienyl)-Ethanone	1.00E-03	0.153 ± 0.002	0.037 ± 0.003	153.05 ± 1.53	36.64 ± 2.69	硫磺、坚果、榛子、核桃 Sulfury，nutty，hazelnut，walnut
2-乙基-3,5-二甲基吡嗪 2-ethyl-3,5-dimethyl-Pyrazine	4.00E-05	0.011 ± 0.000	0.001 ± 0.000	273.62 ± 5.10	20.44 ± 2.16	灼烧、杏仁、烤、坚果、咖啡 Burnt，almond，roasted，nutty，coffee
2-戊基呋喃 2-pentyl-Furan	6.00E-03	0.699 ± 0.023	0.258 ± 0.100	116.56 ± 3.81	43.01 ± 16.72	果香、青草、泥土、豆类、蔬菜、金属 Fruity，green，earthy，Beany，vegetable，metallic
萜类Terpenoids
(+)-柏木脑Cedrol	5.00E-04	0.013 ± 0.001	0.237 ± 0.027	26.92 ± 2.27	474.35 ± 53.95	雪松木、木香、干燥、甜 Cedarwood，woody，dry，sweet
芳樟醇Linalool	6.00E-03	0.777 ± 0.018	0.184 ± 0.016	129.57 ± 3.02	30.69 ± 2.74	花香、青草Floral，green
酯Ester
苯甲酸甲酯 Methyl ester benzoic acid	5.20E-04	0.202 ± 0.043	0.039 ± 0.004	389.31 ± 82.70	74.49 ± 7.64	苯酚、冬青油、杏仁、花香 Phenol，wintergreen，almond，floral，
含硫化合物Sulfur compounds
苯甲硫醇 Benzenemethanethiol	3.50E-06	0.374 ± 0.003	0.051 ± 0.004	106 787.09 ± 938.44	14 608.38 ± 1 056.60	尖锐、葱蒜、硫磺、辣根、薄荷、咖啡Sharp，alliaceous，onion，sulfury，garlic，horseradish，minty，coffee
芳烃Aromatics
二甲萘烷醇 Octahydro-4,8a-dimethyl-4a (2H)-Naphthalenol	2.10E-04	0.003 ± 0.001	0.026 ± 0.003	16.39 ± 2.92	125.25 ± 16.22	新鲜、霉味、土、土壤 Fresh，musty，earthy，soil

组别 Group	差异代谢物总数 Total	上调 Up	下调 Down	组别 Group	差异代谢物总数 Total	上调 Up	下调 Down
HZZ-S2 vs. HZZ-S1	15	12	3	HZZ-S4 vs. HZZ-S3	35	16	19
HZZ-S3 vs. HZZ-S1	167	43	124	MN-S4 vs. MN-S2	174	100	74
HZZ-S4 vs. HZZ-S1	226	45	181	HZZ-S2 vs. MN-S2	63	40	23
HZZ-S3 vs. HZZ-S2	98	34	64	HZZ-S4 vs. MN-S4	187	63	124
HZZ-S4 vs. HZZ-S2	190	42	148

基于代谢组与转录组联合分析解析秋石斛花香形成机制

Elucidation of Floral Scent Formation in Phalaenopsis-Type Dendrobium Hybrids by Integrated Metabolome and Transcriptome Analysis

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组别 Group	差异表达基因数 Number of DEGs	KEGG途径 KEGG pathway
组别 Group	差异表达基因数 Number of DEGs	萜类骨架生物合成 Terpenoid backbone biosynthesis ko00900	单萜生物合成 Monoterpenoid biosynthesis ko00902	二萜生物合成 Diterpenoid biosynthesis ko00904	倍半萜和三萜生物合成 Sesquiterpenoid and triterpenoid biosynthesis ko00909	柠檬烯和蒎烯降解 Limonene and pinene degradation ko00903
HZZ-S2 vs. HZZ-S1	上调Up	28	3	11	2	8
	下调 Down	1	1	3	3	0
HZZ-S3 vs. HZZ-S1	上调Up	94	12	74	37	27
	下调Down	14	2	11	14	0
HZZ-S4 vs. HZZ-S1	上调Up	81	17	88	48	26
	下调Down	18	2	7	15	3
HZZ-S2 vs. MN-S2	上调Up	118	11	65	28	25
	下调Down	71	8	42	25	10
HZZ-S4 vs. MN-S4	上调Up	131	24	109	64	27
	下调Down	62	12	33	37	10

代谢通路注释 Pathway annotation	Unigene ID	NR数据库基因注释 NR database gene annotation	关联代谢物 Conjoint metabolite
单萜生物合成 Monoterpenoid biosynthesis（ko00902）	CL7532.Contig5_All	萜烯合酶样-10 terpene synthase 10-like	α-氧化蒎烯，萜品油烯，(2S-顺)-四氢化-4-甲基-2-(2-甲基-1-丙烯基)-2H-吡喃 [4.1.1.0(2,4)]octane, 2,7,7-trimethyl-3-oxatricyclo，1-Methyl-4-(1-methylethylidene)-cyclohexene，(2S,4R)-4-methyl-2-(2-methylprop-1-en-1-yl)tetrahydro-2H-pyran
二萜生物合成 Diterpenoid biosythesis （ko00904）	CL105.Contig2_All	S-芳樟醇合酶样蛋白S-linalool synthase-like	α-氧化蒎烯，萜品油烯 [4.1.1.0(2,4)]octane, 2,7,7-trimethyl-3-oxatricyclo，1-Methyl-4-(1-methylethylidene)-cyclohexene
	CL105.Contig16_All	S-芳樟醇合酶样蛋白S-linalool synthase-like	茶螺烷 [4.5]dec-6-ene, 2,6,10,10-tetramethyl-1-oxaspiro
	CL105.Contig21_All	S-芳樟醇合酶样蛋白S-linalool synthase-like	α-氧化蒎烯，萜品油烯，(2S-顺)-四氢化-4-甲基-2-(2-甲基-1-丙烯基)-2H-吡喃[4.1.1.0(2,4)]octane, 2,7,7-trimethyl-3-oxatricyclo，1-Methyl-4-(1-methylethylidene)-cyclohexene、 (2S,4R)-4-Methyl-2-(2-methylprop-1-en-1-yl)tetrahydro-2H-pyran
	CL105.Contig39_All	S-芳樟醇合酶样蛋白S-linalool synthase-like	茶螺烷 [4.5]dec-6-ene, 2,6,10,10-tetramethyl-1-oxaspiro
	Unigene66_All	S-芳樟醇合酶样蛋白S-linalool synthase-like	茶螺烷，α-氧化蒎烯，萜品油烯，(2S-顺)-四氢化-4-甲基-2-(2-甲基-1-丙烯基)-2H-吡喃 [4.5]dec-6-ene, 2,6,10,10-tetramethyl-1-oxaspiro， [4.1.1.0(2,4)]octane, 2,7,7-trimethyl-3-oxatricyclo，1-Methyl-4-(1-methylethylidene)-cyclohexene，(2S,4R)-4-Methyl-2-(2-methylprop-1-en-1-yl)tetrahydro-2H-pyran
	Unigene78_All	S-芳樟醇合酶样蛋白S-linalool synthase-like	α-氧化蒎烯，萜品油烯，(2S-顺)-四氢化-4-甲基-2-(2-甲基-1-丙烯基)-2H-吡喃 [4.1.1.0(2,4)]octane, 2,7,7-trimethyl-3-oxatricyclo，1-Methyl-4-(1-methylethylidene)-cyclohexene，(2S,4R)-4-Methyl-2-(2-methylprop-1-en-1-yl)tetrahydro-2H-pyran
倍半萜和三萜生物合成 Sesquiterpenoid and triterpenoid biosynthesis（ko00909）	CL55.Contig2_All	倍半萜合酶 Sesquiterpene synthase	α-氧化蒎烯，萜品油烯，(2S-顺)-四氢化-4-甲基-2-(2-甲基-1-丙烯基)-2H-吡喃 [4.1.1.0(2,4)]octane, 2,7,7-trimethyl-3-oxatricyclo，1-Methyl-4-(1-methylethylidene)-cyclohexene，(2S,4R)-4-Methyl-2-(2-methylprop-1-en-1-yl)tetrahydro-2H-pyran
	CL55.Contig47_All	倍半萜合酶 Sesquiterpene synthase	萜品油烯，(2S-顺)-四氢化-4-甲基-2-(2-甲基-1-丙烯基)-2H-吡喃 1-Methyl-4-(1-methylethylidene)-cyclohexene，(2S,4R)-4-Methyl-2-(2-methylprop-1-en-1-yl)tetrahydro-2H-pyran
	Unigene1_All	(-)-大根香叶烯D合酶 (-)-germacrene D synthase	萜品油烯，(2S-顺)-四氢化-4-甲基-2-(2-甲基-1-丙烯基)-2H-吡喃1-Methyl-4-(1-methylethylidene)-cyclohexene，(2S,4R)-4-Methyl-2-(2-methylprop-1-en-1-yl)tetrahydro-2H-pyran
	Unigene6_All	(-)-大根香叶烯D合酶 (-)-germacrene D synthase	萜品油烯1-Methyl-4-(1-methylethylidene)-cyclohexene

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基于代谢组与转录组联合分析解析秋石斛花香形成机制

Elucidation of Floral Scent Formation in Phalaenopsis-Type Dendrobium Hybrids by Integrated Metabolome and Transcriptome Analysis

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