玉米PLATZ基因<i>GRMZM2G006585</i>启动子的克隆及表达分析

doi:10.7668/hbnxb.20193576

摘要/Abstract

摘要：

高转录活性籽粒特异性启动子可调控目的基因在植物籽粒中特异性、高水平表达。为发掘玉米籽粒特异性启动子,以公开发表的玉米表达谱芯片数据为切入点,筛选出籽粒优势表达基因GRMZM2G006585,克隆其编码区上游约2 000 bp的DNA序列,命名为PZm2G006585。利用在线网站New PLACE和PlantCARE对其进行启动子顺式作用元件分析,发现其含有E-box、P-box等多个籽粒特异性相关元件,初步认为所克隆编码区上游序列为玉米来源的籽粒特异性启动子。为验证其功能,构建该启动子驱动β-葡萄糖苷酸酶基因(GUS)的表达载体并进行植物遗传转化。转基因水稻的GUS组织化学染色结果表明,该启动子驱动外源基因表达模式为籽粒特异、胚优势表达;转基因拟南芥单拷贝株系T₃种子中GUS活性检测结果显示,PZm2G006585驱动的GUS活性为909.52 nmol/(min·mg)。籽粒特异性启动子PZm2G006585的发掘和功能验证为驱动目标基因在玉米、水稻等单子叶植物籽粒中特异性表达提供了候选启动子资源。

关键词: 玉米, 籽粒特异性启动子, 植物生物反应器, PZm2G006585, 遗传转化

Abstract:

Grain specific promoter with high transcription activity can regulate the specific and high-level expression of target genes in plant grains.In order to explore specific promoters of maize grain,the dominant expression gene GRMZM2G006585 was screened from the published data of maize expression profile microarray.The DNA sequence about 2 000 bp upstream of its coding region was cloned and named PZm2G006585.By using the online websites New PLACE and PlantCARE to analyze its promoter elements,we found that it contained multiple grain specific related elements such as E-box and P-box.It was preliminarily considered that the upstream sequence of the cloned coding region was a grain specific promoter from maize.In order to verify its function,the expression vector of GUS gene was constructed and transformed into plants.GUS histochemical staining results of transgenic rice showed that the expression pattern of exogenous genes driven by the promoter was grain specific and embryo dominant.The results of GUS activity detection in T₃ seeds of transgenic Arabidopsis thaliana single copy line showed that the GUS activity driven by PZm2G006585 was 909.52 nmol/(min·mg).The discovery and functional verification of the grain-specific promoter PZm2G006585 can provide candidate promoter resource for specific expression of target genes in maize,rice and other monocotyledons.

Key words: Maize, Grain specific promoter, Plant bioreactor, PZm2G006585, Genetic transformation

王亚丽, 魏琦超, 李成伟. 玉米PLATZ基因GRMZM2G006585启动子的克隆及表达分析[J]. 华北农学报, 2023, 38(2): 21-30. doi: 10.7668/hbnxb.20193576.

WANG Yali, WEI Qichao, LI Chengwei. Cloning and Expression Analysis of Maize PLATZ Gene GRMZM2G006585 Promoter[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(2): 21-30. doi: 10.7668/hbnxb.20193576.

http://www.hbnxb.net/CN/Y2023/V38/I2/21

图/表 11

表1 表达谱分析用引物

Tab.1 Primers for expression profile analysis

引物名称 Primer name	引物序列(5'-3') Primer sequence(5'-3')
ZmTubulin_RT(F)	CCATCTATGACATCTGCCGCCGCTC
ZmTubulin_RT(R)	CAGGGAAGCCGTCAGGGATGAGATG
ZmGAPDH_RT(F)	ACCACCAACTGCCTTGCTCCAC
ZmGAPDH_RT(R)	TTGTGTACTTCTTCCCTTAGCGATGT
GRMZM2G006585_RT(F)	AATGGTATGGATCATGTGGCTAGG
GRMZM2G006585_RT(R)	TTTGGTGGTGGTTCTTCGTAGTT

表2 实时荧光定量PCR引物

Tab.2 Real time fluorescence quantitative PCR primers

引物名称 Primer name	引物序列(5'-3') Primer sequence(5'-3')
ZmLeg1_RT(F)	GTGACCAGCGAGGCGAGATC
ZmLeg1_RT(R)	GACTTGTGACGATGGGAACG
ZmZein27_RT(F)	GATCTTCGGCTTGGTCCTCC
ZmZein27_RT(R)	CGTCAGTTGCTGCGCTATCT

图1 RT-PCR检测不同组织(器官)中GRMZM2G006585基因的转录本丰度 R、ST、L、A、PO、H、SI、C、G分别代表根、茎、叶、花药、花粉、苞叶、花丝、穗轴、授粉后18 d的籽粒。

Fig.1 Transcriptional abundance of GRMZM2G006585 in different tissues(organs)detected by RT-PCR R,ST,L,A,PO,H,SI,C and G respectively represent root,stem,leaf, anther,pollen,husk,silk,cob and grain at 18 days after pollination.

图2 qRT-PCR检测籽粒不同发育时期GRMZM2G006585基因的相对转录本丰度 DAP3,6,9,12,18,24,32分别代表授粉后3,6,9,12,18,24,32 d。

Fig.2 Relative transcript abundance of GRMZM2G006585 at different stages of grain development detected by qRT-PCR DAP3,6,9,12,18,24,32 represent 3,6,9,12,18, 24,32 days after pollination,respectively.

图3 GRMZM2G006585编码区上游序列克隆 M.DNA 标准分子量DL5000+;1. PZm2G006585。

Fig.3 Cloning of upstream sequence of GRMZM2G006585 coding region M.DNA standard molecular weight DL5000+; 1.PZm2G006585.

图4 GRMZM2G006585编码区上游序列启动子顺式作用元件分析下划线表示顺式作用元件,+1表示转录起始位点。

Fig.4 Promoter cis-acting element analysis of upstream sequence of GRMZM2G006585 coding region Cis-acting elements are indicated by underscores,and the transcription site is indicated by +1.

图5 T₀转基因水稻多个器官的GUS染色 A.根;B.茎;C.叶;D.花。1.Pubi;2.缺失启动子;3.PZm2G006585;4.POsGluB-4;5.PZmZein27;6.PZmLeg1。

Fig.5 GUS staining of multiple organs in T₀ generation of transgenic rice A.Root; B.Stem; C.Leaf;D.Flower. 1.Pubi; 2.Promoter deletion;3.PZm2G006585; 4.POsGluB-4; 5.PZmZein27; 6.PZmLeg1.

图6 转基因水稻T₁籽粒不同发育阶段的GUS染色 A.DAF 3(开花后3 d)颖壳;B.DAF 7 颖果;C.DAF 7 颖壳;D.DAF 12 颖果;E.DAF 12 颖壳;F.DAF 40颖果;G.DAF 40颖壳。

Fig.6 GUS staining in different developmental stages of T₁ generation grains of transgenic rice 1.Pubi;2.缺失启动子;3.POsGluB-4;4.PZmZein27;5.PZmLeg1;6.PZm2G006585。 A.Testa(3 days after flowering, DAF 3); B.Caryopsis(DAF 7);C.Testa(DAF 7); D.Caryopsis (DAF 12); E.Testa(DAF 12); F.Caryopsis(DAF 40); G.Testa(DAF 40).1.Pubi; 2.Promoter deletion; 3. POsGluB-4; 4.PZmZein27; 5.PZmLeg1; 6.PZm2G006585.

图7 PZm2G006585在水稻籽粒中的表达模式 A1.缺失启动子(DAF 40); A2. PZm2G006585(开花后12 d,DAF 12); A3.PZm2G006585(DAF 20); B1.PZm2G006585(DAF 30); B2.PZm2G006585(DAF 35); B3.PZm2G006585(DAF 40)。

Fig.7 Expression pattern of PZm2G006585 in rice grains A1.Promoter deletion(DAF 40); A2. PZm2G006585(12 days after flowering, DAF 12); A3.PZm2G006585(DAF 20); B1.PZm2G006585(DAF 30); B2.PZm2G006585(DAF 35); B3.PZm2G006585(DAF 40).

图8 单拷贝纯合体拟南芥的筛选 A.转基因植株的筛选;B.部分T₁阳性株中GUS的 PCR鉴定(M.DNA分子质量标准; 1-47.转基因拟南芥株系编号;48.阴性对照);C.T₃单拷贝纯合体幼苗。

Fig.8 Screening of single copy homozygous Arabidopsis A.Screening of transgenic plants;B.PCR identification of GUS in some T₁ generation positive strains(M.DNA Marker; 1-47.Serial number of transgenic Arabidopsis;48.Negative control);C.T₃ generation single copy homozygotic seedlings.

图9 拟南芥中GUS活性分析不同字母代表0.05水平的差异显著性。

Fig.9 GUS activity analysis in Arabidopsis Different letters represent significant difference at the 0.05 level.

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玉米PLATZ基因GRMZM2G006585启动子的克隆及表达分析

Cloning and Expression Analysis of Maize PLATZ Gene GRMZM2G006585 Promoter

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