基于WGCNA分析CPPU和TDZ对天工墨玉葡萄香气影响及关键基因挖掘

doi:10.3969/j.issn.1004-1524.20240346

浙江农业学报 ›› 2025, Vol. 37 ›› Issue (2): 311-320.DOI: 10.3969/j.issn.1004-1524.20240346

基于WGCNA分析CPPU和TDZ对天工墨玉葡萄香气影响及关键基因挖掘

郑婷(), 向江, 魏灵珠, 吴江, 程建徽()

浙江省农业科学院园艺研究所,浙江杭州 310021

收稿日期:2024-04-13 出版日期:2025-02-25 发布日期:2025-03-20
作者简介:*程建徽,E-mail: chengjianhui@zaas.ac.cn
郑婷(1989—),女,山东德州人,博士,助理研究员,主要从事葡萄品质形成机理研究。E-mail: zhengting12309@163.com
通讯作者: 程建徽
基金资助:
国家自然科学基金(32202414);浙江省农业(果品)新品种选育重大科技专项(2021C02066-6);国家现代农业产业技术体系(CARS-29-13);浙江省“三农九方”农业科技协作计划项目(2024SNJF026);浙江省“三农九方”农业科技协作计划项目(2023SNJF018)

Analysis on the effects of CPPU and TDZ on the aroma of Tiangong Moyu grape using WGCNA and the exploration of the key genes

ZHENG Ting(), XIANG Jiang, WEI Lingzhu, WU Jiang, CHENG Jianhui()

Institute of Horticulture, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China

Received:2024-04-13 Online:2025-02-25 Published:2025-03-20
Contact: CHENG Jianhui

摘要/Abstract

摘要： 为研究处理剂对无核葡萄香气的影响,以天工墨玉为试材,用氯吡脲(CPPU)和噻苯隆(TDZ)搭配赤霉素(GA₃)处理花序,检测成熟果实香气组分和萜类代谢基因表达水平,结合转录组通过加权基因共表达网络分析(WGCNA)探析萜类组分的变化,筛选相关基因。结果表明,在成熟果实中共有35种香气组分,其中,萜类13种。处理后,特征物质——橙花醇、香叶醇和芳樟醇的含量显著(P<0.05)下降。萜类合成甲羟戊酸途径(mevalonate pathway,MVA)关键基因HMGR在处理后表达降低;2-甲基赤藓糖醇-4-磷酸路径(methylerythritol phosphate pathway,MEP)关键基因DXR在CPPU处理后呈下降-上升趋势,TDZ相反。WGCNA结果显示,芳樟醇、橙花醇和香叶醇聚在一起,与turquoise模块高度相关,关联到MYB44基因在处理后大幅下调,尤其TDZ处理后,与物质含量变化一致。研究结果为葡萄生产中CPPU和TDZ的使用提供了理论依据。

关键词: 葡萄, 香气, 氯吡脲(CPPU), 噻苯隆(TDZ), 加权基因共表达网络分析(WGCNA)

Abstract:

In order to study the effect of treatment agents on the aroma of seedless grapes, the early-maturing seedless grape variety Tiangong Moyu was used as the experimental material, and the inflorescence of Tiangong Moyu was treated with forchlorfenuron (CPPU) and thidiazuron (TDZ) combined with gibberellin (GA₃). The aroma components in mature grape fruits were determined, and the expression levels of terpenoid metabolism-related genes in fruit development stage were detected. Combined with transcriptome data, the terpenoid components were analyzed by the weighted gene co-expression network analysis (WGCNA), and the genes related to grape aroma formation were screened. The results showed that a total of 35 components were detected in the mature Tiangong Moyu grape fruits, including 13 terpenoids. After CPPU and TDZ treatment, the content of characteristic aroma substances, namely, nerol, geraniol and linalool decreased significantly (P<0.05). The expression of HMGR(HMGR1 and HMGR3), the rate-limiting enzyme gene of terpenoid synthesis mevalonate pathway (MVA), decreased after treatment. The methylerythritol phosphate pathway (MEP) rate-limiting enzyme gene DXR showed a downward-upward trend after CPPU treatment, while it was the opposite in TDZ. WGCNA results showed that linalool, nerol and geraniol were clustered together, which were highly correlated with the turquoise module. The MYB44 gene was significantly down-regulated after treatment, especially after TDZ treatment, which was consistent with the content changes of the three substances. The results provide a theoretical basis for the application of CPPU and TDZ in grape fruit production.

Key words: grape, aroma, forchlorfenuron (CPPU), thidiazuron (TDZ), weighted gene co-expression network analysis (WGCNA)

中图分类号:

S663.1

郑婷, 向江, 魏灵珠, 吴江, 程建徽. 基于WGCNA分析CPPU和TDZ对天工墨玉葡萄香气影响及关键基因挖掘[J]. 浙江农业学报, 2025, 37(2): 311-320.

ZHENG Ting, XIANG Jiang, WEI Lingzhu, WU Jiang, CHENG Jianhui. Analysis on the effects of CPPU and TDZ on the aroma of Tiangong Moyu grape using WGCNA and the exploration of the key genes[J]. Acta Agriculturae Zhejiangensis, 2025, 37(2): 311-320.

图/表 6

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基因 Gene	上游引物序列 Forward primer sequence(5'-3')	下游引物序列 Reverse primer sequence(5'-3')
VvHMGR1	CGACTTCGTTCAGGGGTTTA	GACTCCAGCGAGTACGAAGG
VvHMGR2	AAGGCCATTTTCACTTGTGG	TTGCAAATCTGCTTGACCTG
VvHMGR3	TAGGGCAGTGCTGTGAGATG	TCAATTCTGCAGCCCTCTTT
VvAACT1	GGATCTCGATTAGGGCATGA	GTGCAGAAATACCACGCTCA
VvAACT2	GTGATGGTGCTGCTGCTTTA	CTGATTTGCAAGAGCCACAA
VvHMGS	ATGGGATTGACCCAAAACAA	CCACCCAGTTGACACAGTTG
VvMVK	CTTGGGTTCATCTGCCTCAT	TCAATTCCAGATGGCTTTCC
VvpMVK	CAACATCAGTGGTTGCTGCT	CCAACTTTCCCTTGTGCAAT
VvMDC	CCCGAGCTAGCAAAATTGAG	ACAAGCACTGCCTGAACCTT
VvFPS	CAGGCGAAAATTTGGACAAT	GCTGGATCTGCTTTCCCATA
VvDXS1	CCAGGTGGTGCATGATGTAG	AAACAACTGGGCCTGTCATC
VvDXS2	AGCAAGCAAGAAGGAATCCA	GCCCACCAGTTTTTGACACT
VvDXS3	CTGCCCATTTCCAAGAAAAA	CATAGGGCTTTCGACCATGT
VvDXS4	AGCAAGGAGGAATCCAAGGT	GCCCACCAGTTTTTGACACT
VvDXS5	CCCCCGAAGACAAGATCATA	CCCAATCACTGCAACAACAC
VvDXS6	GGACAGGCATATGAGGCAAT	GCCTCTTTAGGGCTTTGCTT
VvDXR	GGGAAAAAGATCACGGTTGA	AGGATTGGCAAACGCATATC
VvMCT	GGGAGAAGTGGATCATCGAA	TTCCTTCACTGCACTCATGC
VvCMK	CAATTCTTCTGCGATCACCA	CTACCAGCAGAAGCCCTGAC
VvHDS	GGCTCTTGGAAGTGAGCATC	CAATGCAACAGAAGGAGCAA
VvHDR	ATTCCGATTTTGATGCGAAG	TCCATAGGCTTCTGCCAGTT
VvIDI	TGCTGCACAGAGCTTTCAGT	TCAACAGGTGCATCTTCAGC
VvUBI	GGTGGTATTATTGAGCCATCCTT	AACCTCCAATCCAGTCATCTACT

基因 Gene	上游引物序列 Forward primer sequence(5'-3')	下游引物序列 Reverse primer sequence(5'-3')
VvHMGR1	CGACTTCGTTCAGGGGTTTA	GACTCCAGCGAGTACGAAGG
VvHMGR2	AAGGCCATTTTCACTTGTGG	TTGCAAATCTGCTTGACCTG
VvHMGR3	TAGGGCAGTGCTGTGAGATG	TCAATTCTGCAGCCCTCTTT
VvAACT1	GGATCTCGATTAGGGCATGA	GTGCAGAAATACCACGCTCA
VvAACT2	GTGATGGTGCTGCTGCTTTA	CTGATTTGCAAGAGCCACAA
VvHMGS	ATGGGATTGACCCAAAACAA	CCACCCAGTTGACACAGTTG
VvMVK	CTTGGGTTCATCTGCCTCAT	TCAATTCCAGATGGCTTTCC
VvpMVK	CAACATCAGTGGTTGCTGCT	CCAACTTTCCCTTGTGCAAT
VvMDC	CCCGAGCTAGCAAAATTGAG	ACAAGCACTGCCTGAACCTT
VvFPS	CAGGCGAAAATTTGGACAAT	GCTGGATCTGCTTTCCCATA
VvDXS1	CCAGGTGGTGCATGATGTAG	AAACAACTGGGCCTGTCATC
VvDXS2	AGCAAGCAAGAAGGAATCCA	GCCCACCAGTTTTTGACACT
VvDXS3	CTGCCCATTTCCAAGAAAAA	CATAGGGCTTTCGACCATGT
VvDXS4	AGCAAGGAGGAATCCAAGGT	GCCCACCAGTTTTTGACACT
VvDXS5	CCCCCGAAGACAAGATCATA	CCCAATCACTGCAACAACAC
VvDXS6	GGACAGGCATATGAGGCAAT	GCCTCTTTAGGGCTTTGCTT
VvDXR	GGGAAAAAGATCACGGTTGA	AGGATTGGCAAACGCATATC
VvMCT	GGGAGAAGTGGATCATCGAA	TTCCTTCACTGCACTCATGC
VvCMK	CAATTCTTCTGCGATCACCA	CTACCAGCAGAAGCCCTGAC
VvHDS	GGCTCTTGGAAGTGAGCATC	CAATGCAACAGAAGGAGCAA
VvHDR	ATTCCGATTTTGATGCGAAG	TCCATAGGCTTCTGCCAGTT
VvIDI	TGCTGCACAGAGCTTTCAGT	TCAACAGGTGCATCTTCAGC
VvUBI	GGTGGTATTATTGAGCCATCCTT	AACCTCCAATCCAGTCATCTACT

类别 Types	组分 Compound	T1	T2	T3	T4	T5
醛类 Aldehydes	己醛 Hexanal	1 098.71 ±93.94 a	1 190.80 ±159.92 a	1 273.72 ±315.18 a	1 230.44 ±249.88 a	854.03 ±88.46 a
	3-己烯醛 3-Hexenal	75.56 ±2.89 a	88.95 ±10.74 a	102.72 ±11.32 a	98.23 ±14.49 a	98.85 ±4.56 a
	庚醛 Heptaldehyde	29.90 ±0.69 ab	23.75 ±2.68 b	4.14 ±1.01 c	35.06 ±3.98 a	34.55 ±5.69 a
	2-己烯醛 2-Hexenal	4 873.81 ±130.35 a	5 426.69 ±394.66 a	5 959.74 ±665.93 a	6 269.82 ±708.49 a	5 651.58 ±270.12 a
	壬醛 Nonanal	43.97 ±4.08 b	44.99 ±3.84 b	96.08 ±2.99 a	46.03 ±9.94 b	59.61 ±7.51 b
	(E,E)-2,4-己二烯醛 2,4-Hexadienal,(E,E)-	143.41 ±1.07 b	163.60 ±19.08 ab	170.19 ±18.03 ab	183.49 ±6.20 a	161.47 ±8.48 ab
	反式-2,4-庚二烯醛 2,4-Heptadienal,(E,E)-	20.18 ±3.63 a	25.42 ±3.75 a	20.07 ±5.24 a	33.81 ±5.45 a	29.80 ±4.17 a
	苯甲醛 Benzaldehyde	124.89 ±22.46 a	120.34 ±29.71 a	145.50 ±10.01 a	138.06 ±25.73 a	118.35 ±28.39 a
	反式-2-壬烯醛 2-nonenal, (E)-	15.76 ±0.26 b	13.51 ±0.84 b	15.95 ±1.10 b	39.41 ±0.13 a	28.12 ±3.45 ab
醇类 Alcohols	己醇 Hexyl alcohol	743.16 ±105.07 ab	580.92 ±92.46 b	744.75 ±24.25 ab	866.29 ±9.02 a	606.89 ±44.81 b
	1-辛烯-3-醇 1-Octen-3-ol	44.15 ±4.09 a	42.13 ±12.83 a	33.33 ±10.35 a	47.57 ±10.52 a	44.37 ±11.25 a
	庚醇 1-Heptanol	87.94 ±7.37 a	80.21 ±9.85 a	94.23 ±10.29 a	97.20 ±11.85 a	88.32 ±13.18 a
	4-萜烯醇 Terpinen-4-ol	20.91 ±0.04 b	29.03 ±1.80 b	28.70 ±0.92 b	74.57 ±0.63 a	50.86 ±14.71 ab
	苄醇 Benzyl alcohol	88.18 ±5.24 a	41.42 ±6.25 a	62.86 ±4.36 a	114.47 ±14.03 a	56.58 ±8.18 a
	苯乙醇 Phenylethyl alcohol	368.96 ±10.39 a	334.68 ±43.21 a	361.86 ±33.24 a	419.78 ±52.44 a	286.58 ±58.09 a
萜类 Terpenes	β-蒎烯 β-Pinene	20.10 ±1.83 a	23.38 ±4.19 a	25.98 ±3.34 a	27.25 ±5.82 a	24.52 ±2.11 a
	D-柠檬烯 D-Limonene	36.99 ±0.16 a	30.85 ±8.85 a	44.78 ±10.78 a	39.65 ±4.58 a	31.31 ±3.27 a
	萜品烯 Terpinene	42.66 ±3.44 b	80.90 ±9.56 ab	113.02 ±26.72 a	122.98 ±21.20 a	55.30 ±10.64 b
	萜品油烯 Terpinolene	9.51 ±0.27 a	10.82 ±3.18 a	11.15 ±1.92 a	10.11 ±1.19 a	10.59 ±0.24 a
	玫瑰醚 Rose oxide	25.84 ±1.26 a	31.74 ±6.73 a	34.13 ±1.28 a	34.36 ±1.56 a	32.86 ±2.93 a
	芳樟醇 Linalol	92.18 ±10.35 a	57.27 ±0.99 b	43.39 ±6.04 b	77.92 ±2.09 ab	68.58 ±9.97 ab
	α-松油醇 α-terpineol	71.21 ±7.33 ab	70.06 ±0.15 ab	58.59 ±12.17 b	122.34 ±0.86 a	101.97 ±36.37 ab
	柠檬醛 Citral	57.29 ±1.11 a	37.62 ±3.64 a	65.46 ±29.94 a	88.31 ±31.68 a	56.96 ±29.44 a
	β-香茅醇 β-Cephrol	727.66 ±52.77 b	539.71 ±51.99 c	983.80 ±7.94 a	692.33 ±52.82 b	740.30 ±71.14 b
	橙花醇 Nerol	782.80 ±34.43 a	488.29 ±43.43 bc	481.45 ±26.90 bc	572.95 ±49.98 b	371.46 ±60.45 c
	香叶醇 Lemonol	1 154.37 ±84.05 a	613.16 ±117.24 b	618.15 ±39.42 b	623.57 ±47.30 b	463.94 ±82.80 b
	β-紫罗兰酮 β-Ionone	4.15 ±0.06 a	4.56 ±0.62 a	4.91 ±0.41 a	53.44 ±42.55 a	35.10 ±53.35 a
	香叶酸 Geranic acid	62.44 ±1.28 a	47.20 ±5.52 a	48.46 ±2.67 a	251.47 ±170.87 a	129.66 ±146.83 a
酮类 Ketones	2-辛酮 2-Octanone	329.65 ±32.30 a	380.26 ±28.15 a	384.34 ±21.82 a	326.14 ±45.51 a	385.86 ±13.43 a
酯类 Esters	乙酸乙酯 Ethyl acetate	389.73 ±21.81 bc	316.85 ±48.11 c	617.26 ±21.50 a	600.96 ±33.92 a	528.44 ±65.96 ab
	庚酸乙酯 Ethyl heptanoate	44.94 ±1.41 c	49.80 ±2.77 bc	53.35 ±1.60 bc	66.27 ±3.13 a	57.47 ±4.49 b
	3-羟基扁桃酸乙酯 Ethyl 3-hydroxymandelate	24.49 ±0.22 b	25.72 ±0.03 b	31.98 ±0.95 ab	57.94 ±2.18 a	31.46 ±7.94 ab
	苯甲酸乙酯 Ethyl benzoate	45.92 ±9.97 a	19.36 ±3.49 b	15.65 ±0.27 b	42.86 ±0.85 a	19.36 ±5.60 b
	苯乙酸乙酯 Ethyl phenylacetate	8.33 ±0.09 a	7.90 ±0.88 a	9.10 ±0.06 a	49.14 ±4.06 a	18.81 ±3.72 a
	丁酸丁酯 Butyl butyrate	33.30 ±9.18 a	4.70 ±0.79 a	6.45 ±1.19 a	37.74 ±0.07 a	16.33 ±20.71 a

类别 Types	组分 Compound	T1	T2	T3	T4	T5
醛类 Aldehydes	己醛 Hexanal	1 098.71 ±93.94 a	1 190.80 ±159.92 a	1 273.72 ±315.18 a	1 230.44 ±249.88 a	854.03 ±88.46 a
	3-己烯醛 3-Hexenal	75.56 ±2.89 a	88.95 ±10.74 a	102.72 ±11.32 a	98.23 ±14.49 a	98.85 ±4.56 a
	庚醛 Heptaldehyde	29.90 ±0.69 ab	23.75 ±2.68 b	4.14 ±1.01 c	35.06 ±3.98 a	34.55 ±5.69 a
	2-己烯醛 2-Hexenal	4 873.81 ±130.35 a	5 426.69 ±394.66 a	5 959.74 ±665.93 a	6 269.82 ±708.49 a	5 651.58 ±270.12 a
	壬醛 Nonanal	43.97 ±4.08 b	44.99 ±3.84 b	96.08 ±2.99 a	46.03 ±9.94 b	59.61 ±7.51 b
	(E,E)-2,4-己二烯醛 2,4-Hexadienal,(E,E)-	143.41 ±1.07 b	163.60 ±19.08 ab	170.19 ±18.03 ab	183.49 ±6.20 a	161.47 ±8.48 ab
	反式-2,4-庚二烯醛 2,4-Heptadienal,(E,E)-	20.18 ±3.63 a	25.42 ±3.75 a	20.07 ±5.24 a	33.81 ±5.45 a	29.80 ±4.17 a
	苯甲醛 Benzaldehyde	124.89 ±22.46 a	120.34 ±29.71 a	145.50 ±10.01 a	138.06 ±25.73 a	118.35 ±28.39 a
	反式-2-壬烯醛 2-nonenal, (E)-	15.76 ±0.26 b	13.51 ±0.84 b	15.95 ±1.10 b	39.41 ±0.13 a	28.12 ±3.45 ab
醇类 Alcohols	己醇 Hexyl alcohol	743.16 ±105.07 ab	580.92 ±92.46 b	744.75 ±24.25 ab	866.29 ±9.02 a	606.89 ±44.81 b
	1-辛烯-3-醇 1-Octen-3-ol	44.15 ±4.09 a	42.13 ±12.83 a	33.33 ±10.35 a	47.57 ±10.52 a	44.37 ±11.25 a
	庚醇 1-Heptanol	87.94 ±7.37 a	80.21 ±9.85 a	94.23 ±10.29 a	97.20 ±11.85 a	88.32 ±13.18 a
	4-萜烯醇 Terpinen-4-ol	20.91 ±0.04 b	29.03 ±1.80 b	28.70 ±0.92 b	74.57 ±0.63 a	50.86 ±14.71 ab
	苄醇 Benzyl alcohol	88.18 ±5.24 a	41.42 ±6.25 a	62.86 ±4.36 a	114.47 ±14.03 a	56.58 ±8.18 a
	苯乙醇 Phenylethyl alcohol	368.96 ±10.39 a	334.68 ±43.21 a	361.86 ±33.24 a	419.78 ±52.44 a	286.58 ±58.09 a
萜类 Terpenes	β-蒎烯 β-Pinene	20.10 ±1.83 a	23.38 ±4.19 a	25.98 ±3.34 a	27.25 ±5.82 a	24.52 ±2.11 a
	D-柠檬烯 D-Limonene	36.99 ±0.16 a	30.85 ±8.85 a	44.78 ±10.78 a	39.65 ±4.58 a	31.31 ±3.27 a
	萜品烯 Terpinene	42.66 ±3.44 b	80.90 ±9.56 ab	113.02 ±26.72 a	122.98 ±21.20 a	55.30 ±10.64 b
	萜品油烯 Terpinolene	9.51 ±0.27 a	10.82 ±3.18 a	11.15 ±1.92 a	10.11 ±1.19 a	10.59 ±0.24 a
	玫瑰醚 Rose oxide	25.84 ±1.26 a	31.74 ±6.73 a	34.13 ±1.28 a	34.36 ±1.56 a	32.86 ±2.93 a
	芳樟醇 Linalol	92.18 ±10.35 a	57.27 ±0.99 b	43.39 ±6.04 b	77.92 ±2.09 ab	68.58 ±9.97 ab
	α-松油醇 α-terpineol	71.21 ±7.33 ab	70.06 ±0.15 ab	58.59 ±12.17 b	122.34 ±0.86 a	101.97 ±36.37 ab
	柠檬醛 Citral	57.29 ±1.11 a	37.62 ±3.64 a	65.46 ±29.94 a	88.31 ±31.68 a	56.96 ±29.44 a
	β-香茅醇 β-Cephrol	727.66 ±52.77 b	539.71 ±51.99 c	983.80 ±7.94 a	692.33 ±52.82 b	740.30 ±71.14 b
	橙花醇 Nerol	782.80 ±34.43 a	488.29 ±43.43 bc	481.45 ±26.90 bc	572.95 ±49.98 b	371.46 ±60.45 c
	香叶醇 Lemonol	1 154.37 ±84.05 a	613.16 ±117.24 b	618.15 ±39.42 b	623.57 ±47.30 b	463.94 ±82.80 b
	β-紫罗兰酮 β-Ionone	4.15 ±0.06 a	4.56 ±0.62 a	4.91 ±0.41 a	53.44 ±42.55 a	35.10 ±53.35 a
	香叶酸 Geranic acid	62.44 ±1.28 a	47.20 ±5.52 a	48.46 ±2.67 a	251.47 ±170.87 a	129.66 ±146.83 a
酮类 Ketones	2-辛酮 2-Octanone	329.65 ±32.30 a	380.26 ±28.15 a	384.34 ±21.82 a	326.14 ±45.51 a	385.86 ±13.43 a
酯类 Esters	乙酸乙酯 Ethyl acetate	389.73 ±21.81 bc	316.85 ±48.11 c	617.26 ±21.50 a	600.96 ±33.92 a	528.44 ±65.96 ab
	庚酸乙酯 Ethyl heptanoate	44.94 ±1.41 c	49.80 ±2.77 bc	53.35 ±1.60 bc	66.27 ±3.13 a	57.47 ±4.49 b
	3-羟基扁桃酸乙酯 Ethyl 3-hydroxymandelate	24.49 ±0.22 b	25.72 ±0.03 b	31.98 ±0.95 ab	57.94 ±2.18 a	31.46 ±7.94 ab
	苯甲酸乙酯 Ethyl benzoate	45.92 ±9.97 a	19.36 ±3.49 b	15.65 ±0.27 b	42.86 ±0.85 a	19.36 ±5.60 b
	苯乙酸乙酯 Ethyl phenylacetate	8.33 ±0.09 a	7.90 ±0.88 a	9.10 ±0.06 a	49.14 ±4.06 a	18.81 ±3.72 a
	丁酸丁酯 Butyl butyrate	33.30 ±9.18 a	4.70 ±0.79 a	6.45 ±1.19 a	37.74 ±0.07 a	16.33 ±20.71 a

基于WGCNA分析CPPU和TDZ对天工墨玉葡萄香气影响及关键基因挖掘

Analysis on the effects of CPPU and TDZ on the aroma of Tiangong Moyu grape using WGCNA and the exploration of the key genes

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