甘蓝型油菜酪氨酸代谢关键基因FAH的克隆、功能鉴定和表达分析

doi:10.13560/j.cnki.biotech.bull.1985.2023-0270

摘要/Abstract

摘要：

克隆甘蓝型油菜（Brassica napus L.）酪氨酸代谢关键基因FAH，对其进行功能验证和表达分析，为进一步解析FAH在甘蓝型油菜中的作用和功能提供理论依据。以甘蓝型油菜‘westar’为试材，克隆与拟南芥AtFAH同源性最高的甘蓝型油菜FAH基因BnaA06g38260D（BnaA06FAH）和BnaC05g49430D（BnaC05FAH），通过生物信息学分析其亲缘关系，构建过表达载体转化拟南芥突变体sscd1进行功能验证。克隆BnaA06FAH和BnaC05FAH启动子序列，利用PlantCare在线数据库分析启动子调控元件，构建启动子和GUS的融合载体，通过GUS组织化学染色分析其表达模式。结果显示，BnaA06FAH和BnaC05FAH与AtFAH的氨基酸序列相似性分别为93.11%和92.40%，2个基因过表达都可以完全抑制拟南芥突变体sscd1在短日照下模拟病斑的形成，暗示BnaA06FAH和BnaC05FAH都与AtFAH功能相似。BnaA06FAH和BnaC05FAH启动子除具有所必需的TATA-box和CAAT-box等基本顺式作用元件外，都含有多个与光诱导、激素响应和逆境胁迫响应元件以及多种与抗病相关的顺式作用元件，但与BnaA06FAH相比，BnaC05FAH与拟南芥AtFAH相同的顺式作用元件更多；GUS活性检测表明，BnaC05FAH启动子驱动的GUS基因的表达比BnaA06FAH强，并且两者驱动表达的组织部位不完全相同。因此，BnaA06FAH和BnaC05FAH启动子的作用部位和作用强度都存在明显差异。BnaA06FAH和BnaC05FAH都能够调控拟南芥sscd1模拟病斑的形成，但两者启动子驱动下游基因的强度和部位不同。

关键词: 甘蓝型油菜, FAH, 功能鉴定

Abstract:

The objective of this work is to clone the tyrosine metabolism key gene FAH in Brassica napus L., identify its function and analyze its expression, so as to provide theoretical evidence for further understanding the role and function of FAH in B. napus L. Two FAH genes BnaA06g38260D（BnaA06FAH）and BnaC05g49430D（BnaC05FAH）, which was the highest protein homology to AtFAH in Arabidopsis, were cloned from the B. napus L. variety‘westar’, the phylogenetic relationship was analyzed by bioinformatics, and the overexpression vector was constructed and transformed into Arabidopsis mutant sscd1 for function verification; the promoter sequences of two FAH genes BnaA06FAH and BnaC05FAH were cloned. PlantCARE was used to analyze the promoter cis-acting elements, and the fusion expression vector of gene promoter and GUS was constructed to analyze its expression patterns. As results, the amino acid sequence similarity of BnaA06FAH, BnaC05FAH and AtFAH was 93.11% and 92.40% respectively, the overexpressions of BnaA06FAH and BnaC05FAH completely inhibited the mimic lesion in sscd1 under short-day condition, suggesting both BnaA06FAH and BnaC05FAH had high structural and functional similarity with AtFAH. BnaA06FAH and BnaC05FAH promoters had the core elements of eukaryotic promoter TATA-box and CAAT-box, and also contained several cis-acting elements related to stress, hormone, stress response and a variety of cis-acting elements related to disease resistance, however, BnaC05FAH promoter shared more cis-acting elements with Arabidopsis AtFAH than with BnaA06FAH. GUS activity assays indicated BnaC05FAH promoter drove the expression of GUS gene stronger than BnaA06FAH, and the site of GUS gene expression drove by two different promoters were not completely consistent. Conclusively, there were differences in the sites and intensity where BnaA06FAH and BnaC05FAH promoters functioned. The results showed that both BnaA06FAH and BnaC05FAH played a role in regulating lesion mimic in the sscd1 mutant, but the expression of gene downstream and the expression site drove by BnaA06FAH and BnaC05FAH promoters was different.

Key words: Brassica napus L., fumarylacetoacetate hydrolase, functional identification

支添添, 周舟, 陈纪鹏, 韩成云. 甘蓝型油菜酪氨酸代谢关键基因FAH的克隆、功能鉴定和表达分析[J]. 生物技术通报, 2023, 39(10): 115-127.

ZHI Tian-tian, ZHOU Zhou, CHEN Ji-peng, HAN Cheng-yun. Cloning, Functional Identification and Expression Analysis of FAH, a Key Gene for Tyrosine Metabolism in Brassica napus L.[J]. Biotechnology Bulletin, 2023, 39(10): 115-127.

图/表 10

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引物名称Primer name	引物序列Primer sequence（5'-3'）
BnaC05FAH-ty-F	AACCTACCACTGCCCTTT
BnaC05FAH-ty-R	TTTCCACCAACTTCCTGC
BnaA06FAH-ty-F	ACCATACATCATCCGTTT
BnaA06FAH-ty-R	TCAAGGCAGTGAAGGTAAAA
BnaC05FAH-Xba I -F	GCTCTAGACCATGGCGTTGCTCAAGTCTTT
BnaC05FAH-Sac I -R	CGAGCTCCATCAAGGCAGTGAAGGTAAAA
BnaA06FAH-Xba I -F	GCTCTAGAATGGCGTTGCTCAAGTCTTTCG
BnaA06FAH-Sac I -R	CGAGCTCTCAAGGCAGTGAAGGTAAAAT
BnaC05FAHpro-Xba I -F	GCTCTAGATTAAGGTAATGCTTTGATTCG
BnaC05FAHpro-Bgl II -R	GGAAGATCTGGACGATAAACAAACAGAT
BnaA06FAHpro-Xba I -F	GCTCTAGAAGTTCCCTAACGTTGTCCTCACTTT
BnaA06FAHpro-Bgl II -R	GGAAGATCTTGTATGACGAAGTTTCCAAAGC
qBnaC05FAH-F	AACCTACCACTGCCCTTT
qBnaC05FAH-R	CGTATGACGATGTTTCCG
qBnaA06FAH-F	TTTGGAAACTTCACCGAG
qBnaA06FAH-R	GAGTGTGGAGCAACATCG
ACTIN2-F	AGCACTTGCACCAAGCAGCATG
ACTIN2-R	ACGATTCCTGGACCTGCCTCATC

引物名称Primer name	引物序列Primer sequence（5'-3'）
BnaC05FAH-ty-F	AACCTACCACTGCCCTTT
BnaC05FAH-ty-R	TTTCCACCAACTTCCTGC
BnaA06FAH-ty-F	ACCATACATCATCCGTTT
BnaA06FAH-ty-R	TCAAGGCAGTGAAGGTAAAA
BnaC05FAH-Xba I -F	GCTCTAGACCATGGCGTTGCTCAAGTCTTT
BnaC05FAH-Sac I -R	CGAGCTCCATCAAGGCAGTGAAGGTAAAA
BnaA06FAH-Xba I -F	GCTCTAGAATGGCGTTGCTCAAGTCTTTCG
BnaA06FAH-Sac I -R	CGAGCTCTCAAGGCAGTGAAGGTAAAAT
BnaC05FAHpro-Xba I -F	GCTCTAGATTAAGGTAATGCTTTGATTCG
BnaC05FAHpro-Bgl II -R	GGAAGATCTGGACGATAAACAAACAGAT
BnaA06FAHpro-Xba I -F	GCTCTAGAAGTTCCCTAACGTTGTCCTCACTTT
BnaA06FAHpro-Bgl II -R	GGAAGATCTTGTATGACGAAGTTTCCAAAGC
qBnaC05FAH-F	AACCTACCACTGCCCTTT
qBnaC05FAH-R	CGTATGACGATGTTTCCG
qBnaA06FAH-F	TTTGGAAACTTCACCGAG
qBnaA06FAH-R	GAGTGTGGAGCAACATCG
ACTIN2-F	AGCACTTGCACCAAGCAGCATG
ACTIN2-R	ACGATTCCTGGACCTGCCTCATC

基因 Gene	顺式作用元件 Cis-acting element	核心序列 Core sequence	生物学功能 Biological function
BnaC05FAH BnaA06FAH AtFAH	CAAT-box	CCAAT/CAAT/CAAAT	启动子和增强子区的一般顺式作用元件 Common cis-acting element in promoter and enhancer regions
	TATA-box	ATTATA/TATACA/TATAAGAA	转录起始-30核心启动子元件 Core promoter element around -30 of transcription start
	as-1	TGACG	胁迫响应元件Stress responsive element
	CGTCA-motif	CGTCA	参与茉莉酸甲酯响应的顺式作用元件 Cis-acting regulatory element involved in the MeJA-responsiveness
	GT1-motif	GGTTAA	光响应元件Light responsive element
	O2-site	GTTGACGTGA	参与玉米醇溶蛋白代谢调控的顺式作用元件 Cis-acting regulatory element involved in zein metabolism regulation
	STRE	AGGGG	胁迫响应元件Stress responsive element
	TCT-motif	TCTTAC	部分光响应元件Part of a light responsive element
	TGACG-motif	TGACG	参与茉莉酸甲酯响应的顺式作用元件 Cis-acting regulatory element involved in the MeJA-responsiveness
BnaA06FAH AtFAH	ABRE	ACGTG/CACGTG	脱落酸响应元件Abscisic acid responsive element
	ABRE3a	TACGTG	脱落酸响应元件Abscisic acid responsive element
	ABRE4	CACGTA	脱落酸响应元件Abscisic acid responsive element
	DRE core	GCCGAC	脱水响应元件Dehydration responsive element
	G-box	TACGTG	光响应元件Light responsive element
	Sp1	GGGCGG	光响应元件Light responsive element
	WRE3	CCACCT	创伤响应元件Wounding responsive element

基因 Gene	顺式作用元件 Cis-acting element	核心序列 Core sequence	生物学功能 Biological function
BnaC05FAH BnaA06FAH AtFAH	CAAT-box	CCAAT/CAAT/CAAAT	启动子和增强子区的一般顺式作用元件 Common cis-acting element in promoter and enhancer regions
	TATA-box	ATTATA/TATACA/TATAAGAA	转录起始-30核心启动子元件 Core promoter element around -30 of transcription start
	as-1	TGACG	胁迫响应元件Stress responsive element
	CGTCA-motif	CGTCA	参与茉莉酸甲酯响应的顺式作用元件 Cis-acting regulatory element involved in the MeJA-responsiveness
	GT1-motif	GGTTAA	光响应元件Light responsive element
	O2-site	GTTGACGTGA	参与玉米醇溶蛋白代谢调控的顺式作用元件 Cis-acting regulatory element involved in zein metabolism regulation
	STRE	AGGGG	胁迫响应元件Stress responsive element
	TCT-motif	TCTTAC	部分光响应元件Part of a light responsive element
	TGACG-motif	TGACG	参与茉莉酸甲酯响应的顺式作用元件 Cis-acting regulatory element involved in the MeJA-responsiveness
BnaA06FAH AtFAH	ABRE	ACGTG/CACGTG	脱落酸响应元件Abscisic acid responsive element
	ABRE3a	TACGTG	脱落酸响应元件Abscisic acid responsive element
	ABRE4	CACGTA	脱落酸响应元件Abscisic acid responsive element
	DRE core	GCCGAC	脱水响应元件Dehydration responsive element
	G-box	TACGTG	光响应元件Light responsive element
	Sp1	GGGCGG	光响应元件Light responsive element
	WRE3	CCACCT	创伤响应元件Wounding responsive element

基因 Gene	顺式元件 Cis-acting element	序列 Sequence（5'-3'）	生物学功能 Biological function
BnaC05FAH	AC-I	（T/C）C（T/C）（C/T）ACC（T/C）ACC	木质部特异性表达顺式元件 Cis-acting regulatory element of xylem specific expression
	AE-box	AGAAACAA	光响应元件的一部分Part of a light responsive element
	AT1-motif	AATTATTTTTTATT	光响应元件的一部分Part of a light responsive element
	Box 4	ATTAAT	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
	ERE	ATTTTAAA	乙烯响应元件Ethylene responsive element
	GA-motif	ATAGATAA	光响应元件的一部分Part of a light responsive element
	WUN-motif	CAATTACAT	创伤响应元件Wounding responsive element
	MRE	AACCTAA	参与光响应的MYB结合位点 MYB binding site involved in light responsiveness
BnaA06FAH	A-box	CCGTCC	瞬时顺式调节元件Instantaneous cis-regulating element
	AAGAA-motif	gGTAAAGAAA	胚乳发育相关顺式作用元件 Cis-acting regulatory element involved in endosperm development
	GARE-motif	TCTGTTG	赤霉素响应元件Gibberellin responsive element
	TGA-element	TGACGTAA	生长素响应元件Auxin responsive element
	W-box	（T）TGAC（C/T）	病原诱导相关基因的顺式作用元件 Cis-acting regulatory element of pathogen-induction-related genes
AtFAH	ATCT-motif	AATCTAATCC	参与光响应的保守DNA模块的一部分 Part of a conserved DNA module involved in light responsiveness
	circadian	CAAAGATATC	参与昼夜节律控制的顺式作用调节元件 Cis-acting regulatory element involved in circadian rhythm control
	G-Box	CACGTG	参与光反应的顺式作用调节元件 Cis-acting regulatory element involved in light response