基于WGCNA发掘茶树糖苷类香气前体含量性状相关的候选基因

doi:10.3724/SP.J.1006.2026.54082

作物学报 ›› 2026, Vol. 52 ›› Issue (2): 494-513.doi: 10.3724/SP.J.1006.2026.54082

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

基于WGCNA发掘茶树糖苷类香气前体含量性状相关的候选基因

张力岚(), 杨军, 王让剑()

福建省农业科学院茶叶研究所 / 国家茶树改良中心福建分中心, 福建福州 350013

收稿日期:2025-07-02 接受日期:2025-09-10 出版日期:2026-02-12 网络出版日期:2025-09-23
通讯作者: *王让剑 E-mail:wangrj@faas.cn
作者简介:E-mail: lilanzhang0114@foxmail.com
基金资助:
福建省属公益类科研院所基本科研专项-竞争性项目(2024R1075);福建省农业科学院科技专项(ZYTS202408);福建省茶产业技术体系项目(闽农科教(2023) 16号)

Identification of candidate genes related to glycoside aroma precursor content in tea plant using WGCNA

Zhang Li-Lan(), Yang Jun, Wang Rang-Jian()

Tea Research Institute, Fujian Academy of Agricultural Sciences / Fujian Branch, National Center for Tea Improvement, Fuzhou 350013, Fujian, China

Received:2025-07-02 Accepted:2025-09-10 Published:2026-02-12 Published online:2025-09-23
Contact: *Wang Rang-Jian E-mail:wangrj@faas.cn
Supported by:
Basic Research Projects of Public Welfare Research Institutes in Fujian Province-Competitive Projects(2024R1075);Science and Technology Specific Project of Fujian Academy of Agricultural Sciences(ZYTS202408);the Fujian Tea Industry Technology System (Fujian Agricultural Science and Education(2023)No.16

摘要/Abstract

摘要：

通过分析秋季茶树不同品种新梢的转录组信息, 可以挖掘调控糖苷类香气前体(glycoside aroma precursor, GAP)合成的关键基因, 为研究茶树GAP的合成机制和指导高GAP含量品种选育提供理论参考。本研究以茶树14个品种的新梢为研究材料, 测定8种不同类型的GAP含量, 利用加权基因共表达网络分析(weighted gene co-expression network analysis, WGCNA)转录组数据与新梢GAP含量的表型数据, 鉴定出与GAP合成积累相关的共表达模块与候选基因。8种GAP的含量在茶树新梢中的分布是不均匀的, 其中苯丙烷GAP的含量要远高于萜烯类GAP含量。在低GAP含量品种与高GAP含量品种之间进行比较, 共鉴定出4277个差异表达基因(differentially expressed genes, DEGs)。DEGs的实时荧光定量PCR (quantitative real-time polymerase chain reaction, qRT-PCR)变化趋势与转录组基本一致, 利用该转录组数据获得的分析结果可信。利用WGCNA对过滤后的高表达基因进行划分, 共获得26个共表达模块, 确定了关键模块MEorange、MEyellow、MEdarkturquoise和MElightcyan与8个主要的GAP化合物显著相关(P < 0.01)。对模块中的基因进行GO与KEGG富集分析, 根据基因的连接度以及功能注释, 筛选出16个关键基因, 包括13个结构基因(4个GST、3个GT、3个TPS、1个CYP450、1个CHI和1个DFR)与3个转录因子基因(2个NAC和1个WRKY)。萜烯类GAP与苯丙烷GAP在茶树秋季新梢中的积累具有显著的差异性, 联合分析筛选到与GAP化合物合成积累密切相关的13个结构基因和3个转录因子基因, 这些基因可能在调控茶树秋季新梢GAP的合成积累中起关键作用。

关键词: 茶树, 糖苷类香气前体, 转录组分析, 共表达分析, 调控基因

Abstract:

By analyzing transcriptome data from tender shoots of different tea cultivars in autumn, key genes involved in the regulation of glycoside aroma precursor (GAP) biosynthesis were identified. This study provides a theoretical foundation for elucidating the biosynthetic mechanisms of GAPs and for guiding the breeding of tea cultivars with high GAP content. Tender shoots from 14 tea cultivars were used to quantify eight types of GAPs. Weighted gene co-expression network analysis (WGCNA) was applied to integrate transcriptomic data with GAP content, enabling the identification of co-expression modules and candidate genes related to GAP biosynthesis and accumulation. The distribution of the eight GAP types in tender shoots was uneven, with phenylpropanoid-type GAPs showing significantly higher accumulation than terpene-type GAPs. A total of 4277 differentially expressed genes (DEGs) were identified by comparing high-GAP and low-GAP cultivars. Quantitative real-time polymerase chain reaction (qRT-PCR) validation of selected DEGs showed results consistent with transcriptomic data, confirming the reliability of the analysis. WGCNA identified 26 co-expression modules, among which four modules—MEorange, MEyellow, MEdarkturquoise, and MElightcyan—were significantly correlated with the main GAP types (P < 0.01). GO and KEGG enrichment analyses were performed on genes within these key modules. Based on gene connectivity and functional annotation, 16 key genes were identified, including 13 structural genes (four GSTs, three GTs, three TPSs, one CYP450, one CHI, and one DFR) and three transcription factor genes (two NACs and one WRKY). The seasonal variation in terpene- and phenylpropanoid-type GAP accumulation in autumn was notable, and the identified genes are likely to play central roles in regulating GAP biosynthesis and accumulation in tender tea shoots.

Key words: Camellia sinensis, glycoside aroma precursor, transcriptome analysis, co-expression analysis, regulatory genes

张力岚, 杨军, 王让剑. 基于WGCNA发掘茶树糖苷类香气前体含量性状相关的候选基因[J]. 作物学报, 2026, 52(2): 494-513.

Zhang Li-Lan, Yang Jun, Wang Rang-Jian. Identification of candidate genes related to glycoside aroma precursor content in tea plant using WGCNA[J]. Acta Agronomica Sinica, 2026, 52(2): 494-513.

图/表 13

表1

附表1

图1

表2

表3

图2

图3

图4

图5

表4

每个模块内的基因主要富集到的GO条目"

模块 Module	性状 Trait	GO ID	显著富集的项目 Significantly enriched term	条目类型 Term type	基因数量 Gene number	P值 P-value
MEorange	PG, PP	GO:0010876	脂质定位 Lipid localization	BP	2	0.0030
		GO:0006401	RNA分解代谢过程 RNA catabolic process	BP	2	0.0097
		GO:0000159	蛋白磷酸酶2A型复合物 Protein phosphatase type 2A complex	CC	2	0.0056
		GO:0009378	四向连接解旋酶活性 Four-way junction helicase activity	MF	2	0.0003
		GO:0019888	蛋白磷酸酶活性调节 Protein phosphatase regulator activity	MF	2	0.0086
MEyellow	PG	GO:0006384	RNA聚合酶III启动子的转录启动 Transcription initiation from RNA polymerase III promoter	BP	3	0.0002
		GO:0008284	细胞增殖的正向调控 Positive regulation of cell proliferation	BP	3	0.0013
		GO:0009231	核黄素生物合成过程 Riboflavin biosynthetic process	BP	3	0.0047
		GO:0009991	响应细胞外部刺激 Response to extracellular stimulus	BP	4	0.0062
		GO:0016050	囊泡组织 Vesicle organization	BP	2	0.0065
		GO:0006643	膜脂代谢过程 Membrane lipid metabolic process	BP	2	0.0089
		GO:0000127	转录因子TFIIIC复合物 Transcription factor TFIIIC complex	CC	2	0.0031
		GO:0004527	外切酶活性 Exonuclease activity	MF	3	0.0059
		GO:0016298	脂肪酶活性 Lipase activity	MF	2	0.0065
		GO:0005199	细胞壁的结构组成 Structural constituent of cell wall	MF	2	0.0076
		GO:0045300	酰基-[酰基载体蛋白]脱饱和酶活性 Acyl-[acyl-carrier-protein] desaturase activity	MF	2	0.0088
		GO:0004045	氨基酰基tRNA水解酶活性 Aminoacyl-tRNA hydrolase activity	MF	3	0.0091
MEdarkturquoise	GG	GO:0022900	电子传递链 Electron transport chain	BP	3	0.0019
		GO:0001678	细胞葡萄糖稳态 Cellular glucose homeostasis	BP	2	0.0043
		GO:0009205	嘌呤核糖核苷三磷酸代谢过程 Purine ribonucleoside triphosphate metabolic process	BP	2	0.0050
		GO:0009167	嘌呤核糖核苷一磷酸代谢过程 Purine ribonucleoside monophosphate metabolic process	BP	2	0.0050
		GO:0009126	嘌呤核苷一磷酸代谢过程 Purine nucleoside monophosphate metabolic process	BP	2	0.0050
MEdarkturquoise	GG	GO:0009199	三磷酸核糖核苷代谢过程 Ribonucleoside triphosphate metabolic process	BP	2	0.0050
		GO:0009144	嘌呤核苷三磷酸代谢过程 Purine nucleoside triphosphate metabolic process	BP	2	0.0050
		GO:0006091	前体代谢物和能量的产生 Generation of precursor metabolites and energy	BP	3	0.0064
		GO:0009141	核苷三磷酸代谢过程 Nucleoside triphosphate metabolic process	BP	2	0.0066
		GO:0051560	线粒体钙离子稳态 Mitochondrial calcium ion homeostasis	BP	2	0.0074
		GO:0009161	核糖核苷一磷酸代谢过程 Ribonucleoside monophosphate metabolic process	BP	2	0.0083
		GO:0005758	线粒体膜间隙 Mitochondrial intermembrane space	CC	2	0.0057
		GO:0005666	DNA导向的RNA聚合酶III复合物 DNA-directed RNA polymerase III complex	CC	2	0.0094
		GO:0008146	磺基转移酶活性 Sulfotransferase activity	MF	4	0.0002
		GO:0015018	半乳糖基半乳糖基木糖基蛋白3-β-葡糖醛酸基转移酶活性 Galactosylgalactosylxylosylprotein 3-beta-glucuronosyltransferase activity	MF	2	0.0026
		GO:0005536	葡萄糖结合 Glucose binding	MF	2	0.0032
		GO:0004396	己糖激酶活性 Hexokinase activity	MF	2	0.0045
		GO:0005199	细胞壁的结构组成 Structural constituent of cell wall	MF	2	0.0045
		GO:0015292	转运体活性 Uniporter activity	MF	2	0.0077
		GO:0030246	碳水化合物结合 Carbohydrate binding	MF	7	0.0092
MElightcyan	GG, GP	GO:0006891	高尔基体内囊泡介导的转运 Intra-Golgi vesicle-mediated transport	BP	3	0.0010
		GO:0006013	甘露糖代谢过程 Mannose metabolic process	BP	2	0.0023
		GO:0009094	L-苯丙氨酸生物合成过程 L-phenylalanine biosynthetic process	BP	2	0.0050
		GO:0016592	中间复合体 Mediator complex	CC	4	0.0007
		GO:0046658	质膜固定组件 Anchored component of plasma membrane	CC	3	0.0013
		GO:0098588	细胞器界膜 Bounding membrane of organelle	CC	5	0.0030
		GO:0044444	细胞质部分 Cytoplasmic part	CC	21	0.0033
MElightcyan	GG, GP	GO:0031226	质膜的固有成分 Intrinsic component of plasma membrane	CC	3	0.0057
		GO:0005773	液泡 Vacuole	CC	5	0.0096
		GO:0019213	脱乙酰酶活性 Deacetylase activity	MF	2	0.0010
		GO:0005388	钙转运ATP酶活性 Calcium-transporting ATPase activity	MF	3	0.0017
		GO:0004559	α-甘露糖苷酶活性 Alpha-mannosidase activity	MF	2	0.0025
		GO:0004664	预苯酸脱水酶活性 Prephenate dehydratase activity	MF	2	0.0047
		GO:0010333	萜烯合酶活性 Terpene synthase activity	MF	3	0.0080

表4

表5

图6

表6

不同模块中核心基因的功能注释"

模块 Module	基因ID Gene ID	基因功能描述 Gene description
MEorange	CSS0025104	RAB11结合蛋白RELCH同源物 RAB11-binding protein RELCH homolog (relch)
	CSS0027015	核糖核酸酶H2亚基C Ribonuclease H2 subunit C (RNASEH2C)
	CSS0018839	木葡聚糖半乳糖基转移酶 Xyloglucan galactosyltransferase (XLT)
	CSS0006239	三肽基肽酶2 Tripeptidyl-peptidase 2 (TPP2)
	CSS0036120	线粒体蛋白C2orf69 Mitochondrial protein C2orf69 (C2orf69)
	CSS0011607	微管蛋白β6 Tubulin beta-6 chain (TUBB6)
	CSS0047454	26S蛋白酶体调节亚基4 26S proteasome regulatory subunit 4 (PSMD4)
	CSS0011688	2Fe-2S铁氧还蛋白 2Fe-2S ferredoxin (Fd)
MEyellow	CSS0011428	尿素活性转运蛋白 Urea active transporter (DUR3)
	CSS0035382	UDP糖基转移酶73B UDP-glycosyltransferase 73B (UGT73B)
	CSS0036512	尿苷核苷酶1 Uridine nucleosidase 1 (URH1)
	CSS0044880	IAA丙氨酸抗性蛋白1 IAA-alanine resistance protein 1 (IAR1)
	CSS0026521	线粒体二羧酸/三羧酸转运蛋白 Mitochondrial dicarboxylate/tricarboxylate transporter (DTC)
	CSS0035736	转录因子NAC29 NAC transcription factor 29 (NAC29)
	CSS0034202	查尔酮异构酶 Chalcone isomerase (CHI)
	CSS0002552	萜烯合酶3 Terpene synthase 3 (TPS3)
MEdarkturquoise	CSS0017124	脂肪酸酰胺水解酶 Fatty acid amide hydrolase (FAAH)
	CSS0001960	脂肪酸酰胺水解酶 Fatty acid amide hydrolase (FAAH)
	CSS0025505	TONNEAU 1蛋白 Protein TONNEAU 1 (TON1)
Medarkturquoise	CSS0032881	泛素羧基末端水解酶12 Ubiquitin carboxyl-terminal hydrolase 12 (UBP12)
	CSS0037168	肌动蛋白细胞骨架调控复合蛋白PAN1 Actin cytoskeleton-regulatory complex protein (PAN1)
	CSS0036268	蛋白酶抑制剂 Proteinase inhibitor (PI)
MElightcyan	CSS0023530	含锚蛋白重复结构域超家族 Ankyrin repeat-containing domain superfamily (ANKRD)
	CSS0041188	L-2-羟基戊二酸脱氢酶 L-2-hydroxyglutarate dehydrogenase (L2HGDH)
	CSS0032938	E3泛素蛋白连接酶 E3 ubiquitin-protein ligase (UBE3)
	CSS0026531	重金属相关异丙烯基化植物蛋白 Heavy metal-associated isoprenylated plant protein (HIPP)
	CSS0014110	Dof锌指蛋白 Dof zinc finger protein (DOF)
	CSS0024431	富含半胱氨酸的重复分泌蛋白 Cysteine-rich repeat secretory protein (CRRSP)
	CSS0038267	单铜氧化酶类蛋白 Monocopper oxidase-like protein (MOP)
	CSS0011278	富含半胱氨酸的受体类蛋白激酶 Cysteine-rich receptor-like protein kinase (CRK)

表6

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基因名称 Gene name	正向引物 Forward primer (5'-3')	反向引物 Reverse primer (5'-3')
CsACTIN	GCCATCTTTGATTGGAATGG	GGTGCCACAACCTTGATCTT
CsUGT84A13	AGAACCCGAAAGCGCCTAACTC	CACCCACAAAAACGAAACTCCA
CsAOS1	CGGATCGAACCACCCATATCAT	TCTCTCCCTCCTCACCCACAAA
CsCBP3	CGTAGCTATGGGCCCCTAACTTTC	GTGTGCCTTGTGTCCACCTCTTGA
CsBGLU11	TTAGAGAATTTGGCGATAGGGTTT	CAATGTATGGCTCAGAGGATGAGT
CsCSLE1	AATTGATGGAAGGAGGGAGTGTAG	GAAGTTGTGGTGATATTGGGGTCT
CsIRX9H	GCTATATTGGAGGGTCCGATTTGTAA	CAAGCTGCCTAATAGGCTCAAGAGTG
CsALDH2C4	GACTATAGACCCAAGAACAGGAGAAG	TAATTAGGATACAACCAGGTAAACGA
CsRVE6	TGACCGTGATTGGAAAAAGATTG	TTTGGACGAGGGGGAGGTAGATG
CsUBI3	ATCTTCGTGAAAACCCTAACTGGCA	ACTCCTTCTGGATGTTGTAGTCGGC

品种名称Cultivar name	苯甲醇葡萄糖苷BG	苯甲醇樱草糖苷BP	苯乙醇葡萄糖苷PG	苯乙醇樱草糖苷PP	芳樟醇樱草糖苷LP	香叶醇葡萄糖苷GG	香叶醇樱草糖苷GP	橙花叔醇樱草糖苷NP
BX	1.4737	9.9579	0	16.8421	0.7200	0.1632	3.5474	0.0001
LC	3.9899	3.7980	0.8949	9.9899	2.8081	0.1646	2.5455	0
DG	29.8020	30.2970	30.0990	233.6634	6.3960	0.2366	9.0000	0.0566
JLP	4.5496	0.6991	25.5856	13.6036	1.9189	1.3063	55.7658	1.1802
JB1	21.5385	36.4103	3.8718	89.7436	1.7607	0.1513	7.0427	1.8291
JB2	27.5824	7.1758	147.2527	219.7802	1.7363	0.2681	15.6044	0.0374
FY6	27.9808	33.3654	5.2500	52.6923	0.1962	0.1981	6.0673	0
JMD	34.1026	27.0086	10.9402	72.3932	1.4530	0.3607	23.5897	2.3846
ZCH	0.8018	0.1355	0.4536	0.4718	0.4918	0.0255	1.4273	0.0512
QQ	15.8772	7.1228	22.5439	70.7018	0.7035	0.0462	1.3772	0.3325
BYQL	8.1456	16.5049	1.7282	30.8738	0.8447	0.0413	9.7087	0.2243
JGY	16.1539	1.1795	31.0256	5.4274	1.5299	0.8103	7.8718	0
FDDBC	23.4211	13.5965	41.0526	174.5614	15.0000	0.3544	14.9123	0.4184
YMX	24.5775	15.9155	31.7606	78.8732	10.4930	0.2979	39.1549	1.7183

性状 Traits	最小值 Minimum (mg kg^-1)	最大值 Maximum (mg kg^-1)	平均值±标准差 Mean ± SD (mg kg^-1)	变异系数 CV (%)
苯甲醇葡萄糖苷 BG	0.8018	34.1026	17.1426±11.4892	67.02
苯甲醇樱草糖苷 BP	0.1355	36.4103	14.5119±12.6073	86.88
苯乙醇葡萄糖苷 PG	0	147.2527	25.1756±37.9287	150.66
苯乙醇樱草糖苷 PP	0.4718	233.6634	76.4013±78.6891	102.99
芳樟醇樱草糖苷 LP	0.1962	15.0000	3.2894±4.3736	132.96
香叶醇葡萄糖苷 GG	0.0255	1.3063	0.3160±0.3461	109.52
香叶醇樱草糖苷 GP	1.3772	55.7658	14.1154±15.7444	111.54
橙花叔醇樱草糖苷 NP	0	2.3846	0.5880±0.8265	140.54

分类 Classification	最大值 Maximum	最小值 Minimum	平均值 Average
N50长度 N50 length (bp)	1840	1329	1642
最长Reads长度 Maximum reads length (bp)	832,982	25,203	158,708
过滤核糖体RNA后Clean reads序列条数 Number of clean reads (except rRNA)	5,175,126	3,152,178	4,016,925
全长序列条数 Number of full-length reads	4,667,084	2,789,577	3,563,667
全长序列比例 Full-length percentage (%)	91.03	82.33	88.66

模块 Module	性状 Trait	KEGG 通路号 KEGG pathway ID	显著富集的项目 Significantly enriched term	基因数量 Gene number	P值 P-value
MEorange	PG, PP	ko00730	硫胺素代谢Thiamine metabolism	2	0.0171
MEorange	PG, PP	ko03018	RNA降解RNA degradation	4	0.0230
MEyellow	PG	ko01212	脂肪酸代谢Fatty acid metabolism	7	0.0045
		ko00561	甘油酯代谢Glycerolipid metabolism	7	0.0085
		ko01040	不饱和脂肪酸的生物合成 Biosynthesis of unsaturated fatty acids	3	0.0177
		ko00480	谷胱甘肽代谢Glutathione metabolism	7	0.0195
		ko00941	类黄酮生物合成Flavonoid biosynthesis	7	0.0267
		ko00500	淀粉和蔗糖代谢Starch and sucrose metabolism	11	0.0289
		ko00061	脂肪酸生物合成Fatty acid biosynthesis	4	0.0468
MEdarkturquoise	GG	ko03440	同源重组Homologous recombination	7	0.0077
		ko00130	泛醌和其他萜类醌生物合成 Ubiquinone and other terpenoid-quinone biosynthesis	5	0.0082
		ko00780	生物素代谢Biotin metabolism	3	0.0130
		ko00860	卟啉与叶绿素代谢 Porphyrin and chlorophyll metabolism	4	0.0140
		ko00908	玉米素的生物合成Zeatin biosynthesis	4	0.0335
		ko00061	脂肪酸生物合成Fatty acid biosynthesis	4	0.0367
		ko01212	脂肪酸代谢Fatty acid metabolism	5	0.0408
		ko00196	光合作用-触角蛋白 Photosynthesis-antenna proteins	2	0.0472
MElightcyan	GG, GP	ko04712	昼夜节律-植物Circadian rhythm-plant	4	0.0121
		ko00053	抗坏血酸和醛缩酯代谢 Ascorbate and aldarate metabolism	4	0.0210
		ko00270	半胱氨酸和蛋氨酸代谢 Cysteine and methionine metabolism	5	0.0267
		ko00909	倍半萜和三萜生物合成 Sesquiterpenoid and triterpenoid biosynthesis	3	0.0357

基于WGCNA发掘茶树糖苷类香气前体含量性状相关的候选基因

Identification of candidate genes related to glycoside aroma precursor content in tea plant using WGCNA

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