汉麻<i>CsOLS</i>基因启动子的克隆及上游转录因子的筛选

doi:10.7668/hbnxb.20196830

摘要/Abstract

摘要：

汉麻中大麻素具有丰富的药理活性,汉麻橄榄醇合酶(CsOLS)基因编码的蛋白是催化大麻素前体物质橄榄酸(OA)合成的关键酶。为进一步研究CsOLS基因的表达调控,克隆CsOLS基因的启动子序列(CsOLSp),通过生物信息学分析其转录起始位点(TSS)和顺式元件;克隆约500 bp的CsOLSp截断片段CsOLSp1、CsOLSp2、CsOLSp3,构建诱饵载体,转化酵母细胞及完成自激活检测。以汉麻雌株花、叶等组织构建的cDNA文库为材料进行酵母单杂交筛库,对获得的转录因子进行一对一互作验证、结构域分析和分子对接分析。结果表明,克隆获得1 500 bp 的CsOLSp启动子,包含多种激素响应元件、光响应元件及转录因子结合位点。自激活检测表明,CsOLSp1和CsOLSp3可进行后续酵母文库筛选。经初筛与复筛,分别获得与CsOLSp1和CsOLSp3互作的蛋白数量为5,7个,其中转录因子各一个,分别为CsAHL14 和CsAHL15。一对一互作验证结果表明,CsAHL14可与CsOLSp1发生互作,CsAHL15 可与CsOLSp3发生互作。结构域分析显示,CsAHL14属于AHL家族B类成员,而CsAHL15属于AHL家族A类成员。分子对接分析结果表明,CsAHL14与TTATTTTTTAT序列具有较强的结合能力。推测转录因子 CsAHL14 可能通过特异性结合CsOLSp启动子中的TTATTTTTTAT顺式作用元件参与CsOLS基因的表达调控,在汉麻大麻素生物合成过程中发挥重要作用。

关键词: 汉麻, CsOLS基因, 启动子, 酵母文库筛选, 转录因子

Abstract:

The cannabinoids in hemp have rich pharmacological activities.The olivetol synthase(CsOLS)gene of hemp is a key enzyme that catalyzes the synthesis of olivetolic acid(OA),a precursor substance of cannabinoids.To further investigate the expression regulation of the CsOLS gene,the promoter sequence of the CsOLS gene(CsOLSp)was cloned.Its transcriptional initiation site(TSS)and cis-elements were analysed through bioinformatics.The truncated fragments about 500 bp of CsOLSp,namely CsOLSp1,CsOLSp2,and CsOLSp3,were cloned and constructed into yeast one-hybrid bait vectors,followed by transformation into yeast cells and autoactivation assays.A cDNA library constructed from female hemp flowers and leaves was used for yeast one-hybrid screening.Candidate proteins obtained from the screening were subjected to one-to-one interaction validation,domain analysis,and molecular docking analysis.The results showed that the 1 500 bp promoter sequence of CsOLS gene was successfully cloned,which contained multiple hormone-responsive elements,light-responsive elements,and transcription factor binding sites.Autoactivation assays indicated that CsOLSp1 and CsOLSp3 were suitable for subsequent yeast library screening.Through primary and secondary screening,the numbers of proteins interacting with CsOLSp1 and CsOLSp3 were determined to be 5 and 7 respectively.One transcription factor was obtained from each,namely CsAHL14 and CsAHL15.One-to-one interaction assays confirmed that CsAHL14 interacted with CsOLSp1,while CsAHL15 interacted with CsOLSp3.Domain analysis revealed that CsAHL14 belonged to class B of the AHL family,whereas CsAHL15 belonged to class A.Molecular docking analysis further demonstrated that CsAHL14 exhibited strong binding affinity to the TTATTTTTTAT sequence.It was speculated that the transcription factor CsAHL14 might participate in the expression regulation of the CsOLS gene by specifically binding to the TTATTTTTTAT cis-acting element in the CsOLSp promoter,and play an important role in the cannabinoid biosynthesis process of hemp.

Key words: Hemp, CsOLS gene, Promoter, Yeast library screening, Transcription factor

中图分类号:

赵艳, 杨将, 桑莹莹, 翟莹, 马天意, 张梅娟, 李珊珊. 汉麻CsOLS基因启动子的克隆及上游转录因子的筛选[J]. 华北农学报, 2026, 41(5): 83-91. doi: 10.7668/hbnxb.20196830.

ZHAO Yan, YANG Jiang, SANG Yingying, ZHAI Ying, MA Tianyi, ZHANG Meijuan, LI Shanshan. Cloning of CsOLS Gene Promoter and Screening Upstream Transcription Factors in Hemp[J]. Acta Agriculturae Boreali-Sinica, 2026, 41(5): 83-91. doi: 10.7668/hbnxb.20196830.

http://www.hbnxb.net/CN/Y2026/V41/I5/83

图/表 12

图1 CsOLSp截断示意图

Fig.1 Truncated schematic diagram of CsOLSp

图2 CsOLSp序列的PCR扩增及质粒双酶切结果 M.DNA Marker 15000;1.CsOLSp启动子的PCR扩增条带;2.重组质粒pMD18-T-CsOLSp双酶切鉴定。

Fig.2 PCR amplification of the CsOLSp sequence and double restriction enzyme digestion of the recombinant plasmid M.DNA Marker 15000;1.PCR amplification product of the CsOLSp promoter;2.The double restriction enzyme digestion of the recombinant plasmid pMD18-T-CsOLSp.

图3 CsOLSp序列的TSS和顺式元件分析 A.TSS的预测,并标注+1;B.顺式元件预测(除AT-rich顺式元件);C.AT-rich顺式元件预测。

Fig.3 Analysis of the TSS and cis-elements of the CsOLSp sequence A.Prediction of the TSS,with the +1 position indicated;B.Prediction of cis-elements excluding AT-rich elements;C.Prediction of AT-rich cis-elements.

图4 CsOLSp1、CsOLSp2、CsOLSp3片段的PCR扩增及质粒双酶切结果 M.DNA Marker 15000;1,3,5.分别为CsOLSp1、CsOLSp2、CsOLSp3片段的PCR扩增条带;2,4,6.分别为重组质粒pAbAi-CsOLSp1、pAbAi-CsOLSp2、pAbAi-CsOLSp3的双酶切鉴定。

Fig.4 PCR amplification of CsOLSp1,CsOLSp2 and CsOLSp3 fragments and double-enzyme digestion analysis of recombinant plasmids M.DNA Marker 15000;1,3,5.The PCR amplification products of the CsOLSp1,CsOLSp2,and CsOLSp3 fragments respectively;2,4,6.The double-enzyme digestion identification of the recombinant plasmids pAbAi-CsOLSp1,pAbAi-CsOLSp2,and pAbAi-CsOLSp3 respectively.

图5 诱饵菌株的PCR鉴定 M.DNA Marker 2000;1—3.pAbAi-CsOLSp1、pAbAi-CsOLSp2、pAbAi-CsOLSp3的酵母菌落PCR。

Fig.5 PCR identification of bait strains M.DNA Marker 2000;1—3.Yeast colony PCR of pAbAi-CsOLSp1,pAbAi-CsOLSp2,and pAbAi-CsOLSp3.

图6 诱饵载体自激活检测 p53-AbAi.Y1HGold[p53-AbAi]阳性对照菌株;pAbAi-CsOLSp1、pAbAi-CsOLSp2、pAbAi-CsOLSp3.Y1HGold[pAbAi- CsOLSp1]诱饵菌株、Y1HGold[pAbAi-CsOLSp2]诱饵菌株、Y1HGold[pAbAi-CsOLSp3]诱饵菌株。

Fig.6 Self-activation assay of bait constructs p53-AbAi.The positive control strain Y1HGold[p53-AbAi];pAbAi-CsOLSp1,pAbAi-CsOLSp2,pAbAi-CsOLSp3.The bait strains of Y1HGold[pAbAi-CsOLSp1],Y1HGold[pAbAi-CsOLSp2],Y1HGold[pAbAi-CsOLSp3].

表1 酵母文库筛库结果

Tab.1 Result of yeast library screening

阳性克隆 Positive clone	同源基因登录号 Homologous gene accession number		功能注释 Functional annotation	出现次数 Occurrence frequency
CsOLSp1	1	XM_030637583.2	汉麻 L-抗坏血酸过氧化物酶2	17
	2	XM_030648159.2	汉麻 AT-hook基序核定位蛋白14	2
	3	XM_030637581.2	汉麻未表征蛋白LOC115709467	7
	4	XM_030626708.2	汉麻放氧增强蛋白2	1
	5	XM_030633900.2	汉麻未表征蛋白LOC115706300	1
CsOLSp3	1	XM_030624460.2	汉麻磷脂酰肌醇:神经酰胺肌醇磷酸转移酶1	5
	2	XM_030637932.2	汉麻ATP合酶O亚基	1
	3	XR_004012803.2	汉麻未表征非编码RNA	1
	4	XM_030634175.2	汉麻叶绿素a/b结合蛋白CP26	1
	5	XM_030643705.2	汉麻短链脱氢酶ptmH	1
	6	XM_030637842.2	汉麻含五肽重复序列蛋白At1g80270	1
	7	XM_061103180.1	汉麻 AT-hook 基序核定位蛋白15	1

图7 CsAHL14和CsAHL15片段的PCR扩增及质粒双酶切结果 M.DNA Marker 15000;1,3.CsAHL14和CsAHL15片段的PCR扩增条带;2,4.重组质粒pGADT7-CsAHL14和pGADT7-CsAHL15的双酶切鉴定。

Fig.7 PCR amplification of CsAHL14 and CsAHL15 fragments and double-enzyme digestion analysis of recombinant plasmids M.DNA Marker 15000;1,3.The PCR amplification products of the CsAHL14 and CsAHL15 fragments respectively;2,4.The double-enzyme digestion identification of the recombinant plasmids pGADT7-CsAHL14 and pGADT7-CsAHL15 respectively.

图8 一对一回转验证 p53-AbAi+pGADT7-53.阳性对照;pAbAi-CsOLSp1+pGADT7和pAbAi-CsOLSp3+pGADT7.阴性对照。

Fig.8 One-to-one interaction validation assay p53-AbAi + pGADT7-53.The positive control;pAbAi-CsOLSp1+pGADT7 and pAbAi-CsOLSp3+pGADT7.The negative controls.

图9 汉麻AHL家族结构域分析

Fig.9 Domain analysis of the AHL family in hemp

图10 CsAHL14和CsAHL15与DNA基序的分子对接分析 A.CsAHL14-DNA 复合物整体结构;B.CsAHL14 与 TTATTTTTTAT基序的结合模式;C.CsAHL15-DNA 复合物整体结构;D.CsAHL15 与 TAAATAAATATATT基序的结合模式。

Fig.10 Molecular docking analysis of CsAHL14 and CsAHL15 with DNA motifs A.Overall structure of the CsAHL14-DNA complex;B.Binding mode of CsAHL14 with the TTATTTTTTAT motif;C.Overall structure of the CsAHL15-DNA complex;D.Binding mode of CsAHL15 with the TAAATAAATATATT motif.

表2 蛋白质与核酸序列的对接结果

Tab.2 Docking results for proteins and nucleic acid sequences

蛋白质 Protein	核酸 Nucleic acid	蛋白质结合位点 Binding site of protein	核酸结合位点 Binding site of nucleic acid	保守基序 Conserved motifs	ipTM
CsAHL14	CsOLSp1	ARG-81、ARG-83、ARG-85、LYS-88	+T4、+T6、+T8、-T9	TTATTTTTTAT	0.90
		ARG-83、ARG-85、ARG-87、LYS-88	+A7、-T6、-A7、-T9、-T10、-T11	AAAAATACATAC	0.88
		ARG-81、ARG-83、ARG-85、LYS-88	+T8、-A5、-T6、-T8、-T9、-T10	TAAAAAATAAAA	0.87
		ARG-81、ARG-83、ARG-85、LYS-88	+T6、-A7、-T8、-T10、-T12	AAAAATATTAAT	0.83
		ARG-83、GLY-84、ARG-85、LYS-88	+A2、+T6、-A3、-A5、-T6、-T7	GAATATTA	0.80
CsAHL15	CsOLSp3	ARG-104、ARG-105、ARG-107、ARG-109、GLY-112	+A10、+T11、-T7、-A10、-T11	TAAATAAATATATT	0.69
		ARG-109、PRO-110、GLY-112	-T8、-T7	AAATATTA	0.65
		GLY-112、ASN-115	-T7、-A6	AATATATATA	0.64

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汉麻CsOLS基因启动子的克隆及上游转录因子的筛选

Cloning of CsOLS Gene Promoter and Screening Upstream Transcription Factors in Hemp

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