bHLH转录因子UNE10克隆及其在丁香罗勒挥发性化合物合成调控中的功能

doi:10.13560/j.cnki.biotech.bull.1985.2025-0218

生物技术通报 ›› 2025, Vol. 41 ›› Issue (9): 207-218.doi: 10.13560/j.cnki.biotech.bull.1985.2025-0218

• 研究报告 • 上一篇

bHLH转录因子UNE10克隆及其在丁香罗勒挥发性化合物合成调控中的功能

王斌¹^,²^,³(), 林冲², 袁晓²^,⁴, 蒋园园¹^,²^,³, 王玉昆¹^,²^,³, 肖艳辉¹^,²^,³()

^1.韶关学院广东省粤北食药资源利用与保护重点实验室，韶关 512005
^2.韶关学院生物与农业学院，韶关 512005
^3.粤北特色果蔬工程技术研究中心/韶关市芳香植物工程技术研究中心，韶关 512005
^4.华南农业大学园艺学院，广州 510642

收稿日期:2025-03-02 出版日期:2025-09-26 发布日期:2025-09-24
通讯作者: 肖艳辉，女，硕士，教授，研究方向：芳香植物栽培与利用；E-mail: yhxiao@sgu.edu.cn
作者简介:王斌，男，博士，副教授，研究方向：次生代谢产物合成调控与利用；E-mail: b_wang@sgu.edu.cn
基金资助:
广东省基础与应用基础研究基金(2024A1515010312);广东省教育厅重点科研平台项目(2024GCZX006)

Cloning of bHLH Transcription Factor UNE10 and Its Regulatory Roles in the Biosynthesis of Volatile Compounds in Clove Basil

WANG Bin¹^,²^,³(), LIN Chong², YUAN Xiao²^,⁴, JIANG Yuan-yuan¹^,²^,³, WANG Yu-kun¹^,²^,³, XIAO Yan-hui¹^,²^,³()

^1.Guangdong Provincial Key Laboratory of Utilization and Conservation of Food and Medicinal Resources in Northern Region, Shaoguan University, Shaoguan 512005
^2.College of Biology and Agriculture, Shaoguan University, Shaoguan 512005
^3.Guangdong Provincial Engineering and Technology Research Center of Special Fruit and Vegetables in Northern Region/Engineering Technology Research Center of Shaoguan Aromatic Plant, Shaoguan 512005
^4.College of Horticulture, South China Agricultural University, Guangzhou 510642

Received:2025-03-02 Published:2025-09-26 Online:2025-09-24

摘要/Abstract

摘要：

目的 bHLH转录因子是调控芳香植物次生代谢产物合成的关键因子，探究bHLH转录因子UNE10对丁香罗勒（Ocimum gratissimum）挥发性化合物合成的调控作用，为丁香罗勒精油组分精细调控提供关键调控因子。方法从丁香罗勒中挖掘在叶片显著高表达的bHLH家族基因OgUNE10，利用PCR法扩增OgUNE10的编码序列，通过农杆菌介导的超表达方法在丁香罗勒叶片中过表达OgUNE10，探究其对丁香罗勒叶片挥发性化合物合成的影响。结果 OgUNE10表达呈现明显的组织特异性，在叶片中显著高表达。从丁香罗勒叶片中克隆了OgUNE10基因，其编码161个氨基酸残基的蛋白质。OgUNE10含有保守的bHLH结构域，定位于细胞核，与模式植物bHLH氨基酸序列相似性较低。在叶片中过表达OgUNE10显著影响丁香罗勒精油主要组分的含量，提高丁香酚含量并降低草蒿脑和肉桂酸甲酯含量。过表达OgUNE10显著上调3个苯丙氨酸解氨酶（PAL）基因和2个甲酯酶（MES）基因的表达，下调1个甲酯转移酶（MTA）基因的表达，并影响6个萜烯合成相关基因的表达。结论 OgUNE10可能通过整合苯丙烷途径、MEP和MVA途径调控丁香罗勒挥发性次生代谢产物的生物合成。

关键词: 丁香罗勒, 挥发性化合物, 合成调控, bHLH转录因子, UNE亚家族

Abstract:

Objective bHLH transcription factors are key regulators in the biosynthesis of secondary metabolites in aromatic plants. The investigation into the regulatory roles of the bHLH transcription factor UNE10 in the biosynthesis of volatile compounds in clove basil (Ocimum gratissimum) provides critical regulatory factors for fine-tuning the essential oil components of this plant. Method The bHLH family gene OgUNE10, which has significantly high expression in the leaves of clove basil, was identified. The coding sequence of OgUNE10 was amplified using PCR, followed by its overexpression in the leaves of clove basil through Agrobacterium-mediated transformation to assess its impacts on the biosynthesis of volatile compounds. Result OgUNE10 expression in clove basil demonstrated notable tissue specificity, with particularly high levels in the leaves. The OgUNE10 gene was successfully cloned from the leaves and encoded a protein consisting of 161 amino acid residues, featuring a conserved bHLH domain. This protein was localized in the nucleus and presented low homology with bHLH amino acid sequences from model plants. The overexpression of OgUNE10 in the leaves significantly affected the contents of major components in the essential oil of clove basil, resulting in an increase in eugenol content while decreasing the contents of estragole and methyl cinnamate. Furthermore, OgUNE10 overexpression led to significant upregulations on three PAL (phenylalanine ammonia lyase) and two MES (methylesterase) genes, but downregulations on one MTA (methyltransferases) gene, while also affecting the expression of six terpene synthase-related genes. Conclusion OgUNE10 regulates the biosynthesis of volatile compounds by integrating the phenylpropanoid pathway, MEP and MVA pathways in clove basil.

Key words: clove basil, volatile compounds, biosynthesis regulation, bHLH transcription factor, UNE subfamily

王斌, 林冲, 袁晓, 蒋园园, 王玉昆, 肖艳辉. bHLH转录因子UNE10克隆及其在丁香罗勒挥发性化合物合成调控中的功能[J]. 生物技术通报, 2025, 41(9): 207-218.

WANG Bin, LIN Chong, YUAN Xiao, JIANG Yuan-yuan, WANG Yu-kun, XIAO Yan-hui. Cloning of bHLH Transcription Factor UNE10 and Its Regulatory Roles in the Biosynthesis of Volatile Compounds in Clove Basil[J]. Biotechnology Bulletin, 2025, 41(9): 207-218.

图/表 11

表1 引物序列

Table 1 Primer sequence

基因名称 Gene name	正向序列 Forward sequence（5′-3′）	反向序列 Reverse sequence（5′-3′）	实验用途 Experimental purpose
OgUNE10	ggactctagaggatccccgggATGAAGACATTGCAGAAGTTGGTC	ataagggactgaccacccgggGTTCTTG GAGGCAGAAGCCG	TA克隆、亚细胞定位和过表达 TA cloning， subcellular localization， and overexpression
OgACTIN7	GGAGCTCGTCTTTGCTGTCT	GAGCGGGAAATTGTGAGGGA	RT-qPCR
OgUNE10	TGTCTATGATGGGCATGGGC	GCCTGACAAGCAAAGAAGGC
OgPAL1	TTTTTCGCCCGTCAAGAAGC	CTGGAGAGTGGGAATCCAGC
OgPAL2	GGATCCCCTGCAGAAACCAA	ATCAATCAGCGGGTTGTCGT
OgPAL4	ATGCCGGAGAAACCTTGGAG	ATTCTCCCTCGCTCAGCAAC
OgMES1	GATAGGCCCTCGTGGAGTTG	TAACTGCATCAACGCCCAGT
OgMES2	CTCCATGGTCCGGGCTATTC	TGTCCCCTCAAATCCAAGGC
OgMAT1	TCTACACACGCACACTCGAC	GTACCAGCACCATGCTCCAT
OgEFS1	CCACACTTGTTGCTCCTTGC	GCTGGTAGTTTCCAGAGCGT
OgMD1	GGTCTTCAACAGCCCGATCA	ACGTCGGAACTTTTGACGGA
OgCyt P450	AATCCGATAATGGGTCCGCC	TTGCTGCAGGAGAAGTAGCC
OgG8H1	TGGGTCCGTCAAATCCTTGG	ACTGCCGGAAAAGTAGCCTC
OgiPR1	GCAAAGTGAAGAGGCCAACG	TTCTGCCGAATGAATGGGCT
OgGD2	ATCTTGGCGTGGACGAGTTT	TCAGCAACGGAACAATGGGA

图1 OgUNE10在丁香罗勒不同组织中的表达FPKM表示每百万reads中来自比对到某一基因每千碱基长度的reads数目，误差线为标准误。下同

Fig. 1 Expressions of OgUNE10 in different tissues of clove basil (Ocimum gratissimum)FPKM refers to the number of reads per kilobase length from alignment to a gene per million reads, error bars indicate standard errors. The same below

图2 OgUNE10 PCR产物电泳与序列A：以cDNA为模板扩增的PCR产物电泳图；B：以连接OgUNE10全长的重组质粒为模板扩增的PCR产物电泳图；C：转录组测序组装的OgUNE10全长和克隆序列。泳道M：DNA marker；泳道1-6：OgUNE10全长PCR扩增产物；R：RNA-Seq组装的序列；S：Sanger测序序列；红色字体为差异碱基

Fig. 2 Electrophoretic image of OgUNE10 PCR products and sequencesA: Electrophoretic image of PCR product amplified using cDNA as template. B: Electrophoretic image of PCR products amplified using the recombinant plasmid linking the full length of OgUNE10 as templates. C: Full length of OgUNE10 assembled by RNA-Seq and obtained from cloning sequence. Lane M: DNA marker. Lanes 1-6: Full length OgUNE10 PCR amplified products. R: RNA-Seq assembled sequence. S: Sanger sequencing sequence; red letters are different bases

图3 OgUNE10多序列比对（A）和保守性分析（B）红色方框内为bHLH保守结构域，蛋白登录号见图4-B

Fig. 3 Multiple sequence alignment (A) and conservation analysis (B) of OgUNE10Sequences in red box is the bHLH conserved domain, protein accession No. are indicated in Fig. 4-B below

表2 丁香罗勒OgUNE10与4种模式植物UNE10蛋白序列相似性比较

Table 2 Similarity comparison of the sequences between OgUNE10 of clove basil and UNE10 proteins from four model plants/%

植物类型Plant species	丁香罗勒Ocimum gratissimum	水稻Oryza sativa	拟南芥Arabidopsis thaliana	烟草Nicotiana tabacum	番茄Solanum lycopersicum
丁香罗勒 Ocimum gratissimum	100
水稻 Oryza sativa	23.55	100
拟南芥 Arabidopsis thaliana	29.51	52.57	100
烟草 Nicotiana tabacum	24.31	47.90	52.89	100
番茄 Solanum lycopersicum	22.87	47.98	53.05	79.45	100

图4 OgUNE10亚细胞定位（A）和系统发育分析（B）LaMYC4和MhbHLH96氨基酸序列分别见文献［13， 26］

Fig. 4 Subcellular localization (A) and phylogenetic analysis (B) of OgUNE10 proteinThe amino acid sequences of LaMYC4 and MhbHLH96 can be found in the references [13, 26], respectively

图5 丁香罗勒叶片中挥发性化合物的色谱图A：转化pBI121空载的叶片；B：过表达OgUNE10的叶片

Fig. 5 Chromatogram of volatile compounds in clove basil leavesA: Leaves transforming pBI 121 empty plasmid. B: Leaves overexpressing OgUNE10

表3 过表达OgUNE10对丁香罗勒叶片挥发性化合物种类和相对含量的影响

Table 3 Effects of OgUNE10 overexpressionon the types and relative content of volatile compounds in clove basil leaves

序号 No.	化合物 Compound	保留时间 Retention time （min）	相对含量 Relative content （%）
序号 No.	化合物 Compound	保留时间 Retention time （min）	35S：：GFP	35S：：OgUNE10-GFP
1	3-己烯醛 3-Hexenal	4.05	0.27±0.01	0.38±0.08*
2	α-蒎烯 α-Pinene	6.19	0.23±0.04	0.25±0.04
3	莰烯 Camphene	6.45	-	0.10±0.01
4	香桧烯 Sabinene	6.84	0.13±0.03	-
5	β-蒎烯 β-Pinene	6.90	0.24±0.04	0.27±0.02
6	β-月桂烯 β-Myrcene	7.08	0.14±0.01	0.19±0.03
7	D-枞油烯 D-sylvestrene	7.73	0.10±0.01	0.15±0.01*
8	桉叶油醇 Eucalyptol	7.78	0.64±0.08	0.71±0.06
9	反式-β-罗勒烯 Trans-β-Ocimene	8.00	0.77±0.03	0.74±0.03
10	茴香酮 Fenchone	8.71	0.23±0.04	1.02±0.05*
11	芳樟醇 Linalool	8.81	1.53±0.05	4.65±0.19*
12	（+）-2-莰酮（+）-2-Bornanone	9.60	0.28±0.03	0.71±0.12*
13	L-4-松油醇 L-terpinen-4-ol	10.06	0.21±0.03*	0.14±0.03
14	L-α-松油醇 L-α-Terpineol	10.25	0.09±0.01	0.10±0.02
15	草蒿脑 Estragole	10.35	17.43±1.33*	1.08±0.03
16	顺式肉桂酸甲酯 Methyl cis-cinnamate	11.87	0.17±0.05	-
17	丁香酚 Eugenol	12.59	20.10±5.75	46.58±1.11*
18	肉桂酸甲酯 Methyl cinnamate	12.94	51.39±5.28*	34.43±0.28
19	β-榄香烯 β-Elemene	13.09	0.13±0.01	0.28±0.01*
20	甲基丁香酚 Methyl eugenol	13.16	1.48±0.23	3.73±0.04*
21	反式-α-佛手柑油烯 Trans-α-Bergamotene	13.64	0.84±0.09	3.73±0.04*
22	α-愈创木烯 α-Guaiene	13.71	-	0.09±0.01
23	蛇麻烯 Humulene	13.95	0.21±0.05	0.35±0.03*
24	顺式-4（15），5-衣兰油二烯 Cis-Muurola-4（15），5-diene	14.06	0.19±0.02	0.19±0.04
25	大根香叶烯D Germacrene D	14.29	0.95±0.11	0.97±0.06
26	β-环大根香叶烷 β-Cyclogermacrane	14.48	0.22±0.02	0.33±0.02*
27	δ-愈创木烯 δ-Guaiene	14.58	-	0.14±0.02
28	γ-愈创木烯 γ-Cadinene	14.68	0.62±0.03	0.83±0.04*
29	τ-杜松醇 τ-Cadinol	16.18	0.83±0.08	0.77±0.06
总鉴定率 Total identification rate （%）			99.42	99.49

表3 过表达OgUNE10对丁香罗勒叶片挥发性化合物种类和相对含量的影响

Table 3 Effects of OgUNE10 overexpressionon the types and relative content of volatile compounds in clove basil leaves

序号 No.	化合物 Compound	保留时间 Retention time （min）	相对含量 Relative content （%）
序号 No.	化合物 Compound	保留时间 Retention time （min）	35S：：GFP	35S：：OgUNE10-GFP
1	3-己烯醛 3-Hexenal	4.05	0.27±0.01	0.38±0.08*
2	α-蒎烯 α-Pinene	6.19	0.23±0.04	0.25±0.04
3	莰烯 Camphene	6.45	-	0.10±0.01
4	香桧烯 Sabinene	6.84	0.13±0.03	-
5	β-蒎烯 β-Pinene	6.90	0.24±0.04	0.27±0.02
6	β-月桂烯 β-Myrcene	7.08	0.14±0.01	0.19±0.03
7	D-枞油烯 D-sylvestrene	7.73	0.10±0.01	0.15±0.01*
8	桉叶油醇 Eucalyptol	7.78	0.64±0.08	0.71±0.06
9	反式-β-罗勒烯 Trans-β-Ocimene	8.00	0.77±0.03	0.74±0.03
10	茴香酮 Fenchone	8.71	0.23±0.04	1.02±0.05*
11	芳樟醇 Linalool	8.81	1.53±0.05	4.65±0.19*
12	（+）-2-莰酮（+）-2-Bornanone	9.60	0.28±0.03	0.71±0.12*
13	L-4-松油醇 L-terpinen-4-ol	10.06	0.21±0.03*	0.14±0.03
14	L-α-松油醇 L-α-Terpineol	10.25	0.09±0.01	0.10±0.02
15	草蒿脑 Estragole	10.35	17.43±1.33*	1.08±0.03
16	顺式肉桂酸甲酯 Methyl cis-cinnamate	11.87	0.17±0.05	-
17	丁香酚 Eugenol	12.59	20.10±5.75	46.58±1.11*
18	肉桂酸甲酯 Methyl cinnamate	12.94	51.39±5.28*	34.43±0.28
19	β-榄香烯 β-Elemene	13.09	0.13±0.01	0.28±0.01*
20	甲基丁香酚 Methyl eugenol	13.16	1.48±0.23	3.73±0.04*
21	反式-α-佛手柑油烯 Trans-α-Bergamotene	13.64	0.84±0.09	3.73±0.04*
22	α-愈创木烯 α-Guaiene	13.71	-	0.09±0.01
23	蛇麻烯 Humulene	13.95	0.21±0.05	0.35±0.03*
24	顺式-4（15），5-衣兰油二烯 Cis-Muurola-4（15），5-diene	14.06	0.19±0.02	0.19±0.04
25	大根香叶烯D Germacrene D	14.29	0.95±0.11	0.97±0.06
26	β-环大根香叶烷 β-Cyclogermacrane	14.48	0.22±0.02	0.33±0.02*
27	δ-愈创木烯 δ-Guaiene	14.58	-	0.14±0.02
28	γ-愈创木烯 γ-Cadinene	14.68	0.62±0.03	0.83±0.04*
29	τ-杜松醇 τ-Cadinol	16.18	0.83±0.08	0.77±0.06
总鉴定率 Total identification rate （%）			99.42	99.49

图6 OgUNE10表达及PAL基因表达分析

Fig. 6 Expression analysis of OgUNE10 and PAL genes

图7 甲酯转移酶和甲酯酶编码基因表达分析

Fig. 7 Expression analysis of genes encoding methyltransferase and methylesterase

图8 萜烯合成相关基因的表达分析OgEFS1：（-）-葑醇合成酶1；OgMD1：甘露醇脱氢酶；OgCyt P450：细胞色素氧化酶P450；OgG8H1：香叶醇8-羟化酶；OgiPR1：（-）-异胡椒酮还原酶1；OgGD2：香叶醇脱氢酶2

Fig. 8 Expression analysis of genes related to terpenoids biosynthesisOgEFS1: (-)-Endo-fenchol synthase 1. OgMD1: Mannitol dehydrogenase 1. OgCyt P450: Cytochrome P450. OgG8H1: Geraniol 8-hydroxylase 1. OgiPR1: (-)-isopiperitenone reductase 1. OgGD2: Geraniol dehydrogenase 2

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bHLH转录因子UNE10克隆及其在丁香罗勒挥发性化合物合成调控中的功能

Cloning of bHLH Transcription Factor UNE10 and Its Regulatory Roles in the Biosynthesis of Volatile Compounds in Clove Basil

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