玉米抗旱相关基因的克隆与功能分析
摘要
水分胁迫是主要的非生物胁迫因子之一,严重影响植物生长及农作物生产。本研究在我们前期研究的干旱胁迫诱导的差减文库中克隆了三个受干旱诱导表达的基因:ZmFBL2、ZmCKS2和ZmBTF3b。
ZmFBL2基因编码蛋白ZmFBL2含有一个F-box结构域和一个富含亮氨酸结构域(LRR)。F-box结构域最初是在细胞周期蛋白F的N末端被鉴定的,命名为F-box。F-box蛋白在N末端都含有保守的F-box结构域(约含有40-50个氨基酸)。F-box蛋白在C末端含有一个或多个高度可变的蛋白-蛋白互作结构域,如富含亮氨酸的结构域LRR、Kelch、WD40(Trp和Asp)和三十四肽重复单位TPR,这些结构域可以对底物进行特异的结合。过去的几年中,高等植物拟南芥、矮牵牛、水稻、玉米中已经鉴定出数个F-box蛋白。这些F-box蛋白在植物花发育、自交不亲、种子发育、生理节奏调节、防卫反应、生长素信号途径、茉莉酸信号途径、水杨酸信号途径以及非生物胁迫响应等过程起重要作用。本研究克隆得到玉米中受干旱诱导表达的一个F-box蛋白ZmFBL2,并研究其结构与功能。
ZmCKS2含有一个CKS(cyclin-dependent kinase regulatory subunit)结构域,该结构域在真核生物中高度保守;ZmBTF3b是一个含有169个氨基酸的蛋白,含有一个保守的NAC (nascent polypeptide-associated complex)结构域。
主要研究结果如下:
1、应用同源克隆的方法获得ZmFBL2基因的cDNA全长,结构分析发现,ZmFBL2蛋白N端含有典型的F-box结构域,C端含有LRR结构域;通过PCR扩增并获得ZmFBL2基因组DNA及启动子序列;ZmFBL2基因组含有四个外显子和三个内含子;ZmFBL2启动子含有早期应答脱水胁迫、响应高盐、冷、水杨酸、ABA、氧化胁迫等非生物胁迫的顺式作用元件以及胚、胚乳、种子发育、光诱导、昼夜节律调节相关的调控元件。
2、组织差异性表达分析结果表明,ZmFBL2在玉米幼叶、穗位叶、开花期根及雄穗中表达量较高,其中穗位叶中表达量最高;在脱水和PEG处理下,ZmFBL2在地上部分和地下部分均表现显著上调表达趋势;在冷处理条件下,ZmFBL2在地下部分的表达量无显著的变化趋势,但是在地上部分表达量有显著的上调表达趋势。在NaCl处理条件下,ZmFBL2在地上部分及地下部分均显示上调表达趋势,但未达到显著上调表达水平;在响应4种外源激素信号分子时,ZmFBL2对MJ处理无显著的诱导表达变化,而在应答2,4-D和ABA两个外源激素时均表现出上调表达模式;对SA处理表现出下调表达趋势。
3、通过同源克隆,获得玉米中10个MSK(maize skp1-like gene)基因,分别命名为MSK1-10。酵母双杂交结果表明,F-box蛋白(1-368氨基酸)可以与MSK1、MSK2、MSK9及ZmMADS6、ZmMADS7互作。F2(1-75氨基酸)可以与MSK1、MSK2、MSK5、MSK9、CKS及ZmMADS4、ZmMADS6、ZmMADS7互作。而F1(76-368氨基酸)、F3(1-38氨基酸)不能与任何MSK蛋白或ZmMADS蛋白互作。β-半乳糖苷酶显色检验分析也验证了以上互作结果的可靠性。
4、应用20%PEG处理在营养土中生长至10天的过表达转基因株系及野生型拟南芥,结果表明过表达转基因株系存活率明显高于野生型;在含有0.3μM ABA、150mM NaCl的MS固体培养基上,过表达转基因株系存活率明显高于野生型;而在含有350mM甘露醇MS固体培养基上,表达转基因株系与野生型存活率差异不明显。
5、在本研究的过程中还通过同源克隆的方法克隆得到干旱诱导表达的ZmCKS2和ZmBTF3b基因。ZmCKS2基因编码的蛋白ZmCKS2是细胞周期依赖性蛋白激酶的亚基。该基因启动子序列中含有多个与逆境胁迫相关的顺式作用元件;ZmCKS2基因在幼穗中表达量最高,并受干旱胁迫和茉莉酸甲酯处理上调表达,受冷胁迫和外源ABA处理下调表达。说明该基因可能在玉米应答逆境胁迫过程中起重要作用。ZmBTF3b基因编码的蛋白ZmBTF3b含有保守的新生多肽复合体(NAC)结构域;酵母单杂交试验表明该蛋白C末端具有转录激活功能。该基因启动子序列中有多个与逆境胁迫相关的顺式作用元件;表达模式分析表明该基因在花丝、幼穗、雄穗、幼根和开花期根中表达量相对较高,叶子中表达量相对较低,并且在根部受脱水干旱和PEG处理上调表达,受冷胁迫和NaCl处理下调表达。结果表明,该基因可能与玉米在应答逆境胁迫过程中起重要作用。
Water stress is one of the major environmental stresses that seriously affect plant growth and crop yield worldwide. In this study we cloned three drought induced genes ZmFBL2, ZmCKS2 and ZmBTF3b from our previous drought related SSH library.
ZmFBL2 is a F-box protein, contein a F-box domain and LRR domain. F-box domain was fistly found at the N-terminal region of cyclin F. At the N terminus of F-box proteins contained a conserved F-box motif (about 40~50 amino acid) and at the carboxy-terminal, F-box proteins often contained highly variable protein-protein interaction domains, such as WD40 repeat (Trp and Asp), Leu-rich repeat (LRR), tetratricopeptide repeat (TPR) and kelch repeat. In the past few years, many F-box proteins have been cloned in higher plants such as Arabidopsis, petunia, rice and maize. These F-box proteins played essential roles in flowering development, self-incompatibility, light signalling and clock control, plant hormone response pathways and abiotic stress. In this study, we isolated an drought induced gene ZmFBL2, a new F-box gene in maize. Its structure and functions was analized.
ZmCKS2 was predicted to contain a CKS (cyclin-dependent kinase regulatory subunit) structural domain, which was highly conserved in eukaryotes kingdom. ZmBTF3b encoded a putative transcription factor of 169 amino acids with a conserved NAC (nascent polypeptide-associated complex) domain.
The main results were as follows:
1. In this study we cloned the full-length cDNA of ZmFBL2, by using in silico and homology-based cloning techniques. The deduced protein of ZmFBL2 was predicted to contain a F-box structural domain, which was highly conserved in eukaryotes kingdom and a Leu-rich repeat (LRR) domains at the carboxy terminal. The promoter of ZmFBL2 was predicted to contain important regulatory elements including core promoter elements and drought, high salt, cold, SA, ABA, oxygen responsive, light regulation and embryo, endosperm, seeds development elements.
2. Real-time quantitative PCR (qRT-PCR) analysis revealed that ZmFBL2 was highly expressed in leaves at seedling stage, ear leaves, roots at flowering stage and tassels and was up-regulated under dehydration, PEG, cold, NaCl, 2,4-D and ABA treatment while was down-regulated under SA .
3. We cloned the full-length cDNA of ten MSK (maize skp1-like gene) genes by using in silico and homology-based cloning techniques. Yeast hybrid assay revealed that F-box protein interacted specifically with MSK1、MSK2、MSK9 and ZmMADS6、ZmMADS7 and the F-box domain was necessary.β-Galactosidase filter assays verified the reliability of the results.
4. Ten days old lins grown in organic fertilizer was treated by 20% PEG 6000. Survival rates of transgenic Arabidopsis was higher than WT Arabidopsis. On MS medium with ABA and NaCl, survival rates of transgenic lines was also higher than WT. But there was no obvious difference on MS medium with mannitol.
5. Two other drought induced genes ZmCKS2 and ZmBTF3b were cloned in this study. The deduced protein of ZmCKS2 was predicted to contain a CKS (cyclin-dependent kinase regulatory subunit) structural domain, which was highly conserved in eukaryotes kingdom. The promoter of ZmCKS2 was predicted to contain important regulatory elements including core promoter elements and drought, high salt, cold, ABA, light regulation, copper and oxygen responsive elements. Real-time quantitative PCR (qRT-PCR) analysis revealed that ZmCKS2 was highly expressed in ears while its expression was very low in roots and leaves at seedling stage and was up-regulated under drought or MeJA treatment but down-regulated under cold or ABA treatment. Under NaCl stress ZmCKS2 showed no obvious change. These results revealed that ZmCKS2 might be involved in multiple pathways of plants responding to different environmental conditions. The cloned ZmBTF3b gene encoded a putative transcription factor of 169 amino acids with a conserved NAC (nascent polypeptide-associated complex) domain. Transcriptional activity analysis showed that ZmBTF3b might be a functional transcription factor. Real-time quantitative PCR (qRT-PCR) analysis revealed that ZmBTF3b was highly expressed in silks, ears and tassels and was up-regulated under dehydration or PEG treatment while was down-regulated under cold, NaCl, ABA or SA. ZmBTF3b might have the transcriptional activation activity and play an essential role in response to abiotic stresses.
ZmFBL2基因编码蛋白ZmFBL2含有一个F-box结构域和一个富含亮氨酸结构域(LRR)。F-box结构域最初是在细胞周期蛋白F的N末端被鉴定的,命名为F-box。F-box蛋白在N末端都含有保守的F-box结构域(约含有40-50个氨基酸)。F-box蛋白在C末端含有一个或多个高度可变的蛋白-蛋白互作结构域,如富含亮氨酸的结构域LRR、Kelch、WD40(Trp和Asp)和三十四肽重复单位TPR,这些结构域可以对底物进行特异的结合。过去的几年中,高等植物拟南芥、矮牵牛、水稻、玉米中已经鉴定出数个F-box蛋白。这些F-box蛋白在植物花发育、自交不亲、种子发育、生理节奏调节、防卫反应、生长素信号途径、茉莉酸信号途径、水杨酸信号途径以及非生物胁迫响应等过程起重要作用。本研究克隆得到玉米中受干旱诱导表达的一个F-box蛋白ZmFBL2,并研究其结构与功能。
ZmCKS2含有一个CKS(cyclin-dependent kinase regulatory subunit)结构域,该结构域在真核生物中高度保守;ZmBTF3b是一个含有169个氨基酸的蛋白,含有一个保守的NAC (nascent polypeptide-associated complex)结构域。
主要研究结果如下:
1、应用同源克隆的方法获得ZmFBL2基因的cDNA全长,结构分析发现,ZmFBL2蛋白N端含有典型的F-box结构域,C端含有LRR结构域;通过PCR扩增并获得ZmFBL2基因组DNA及启动子序列;ZmFBL2基因组含有四个外显子和三个内含子;ZmFBL2启动子含有早期应答脱水胁迫、响应高盐、冷、水杨酸、ABA、氧化胁迫等非生物胁迫的顺式作用元件以及胚、胚乳、种子发育、光诱导、昼夜节律调节相关的调控元件。
2、组织差异性表达分析结果表明,ZmFBL2在玉米幼叶、穗位叶、开花期根及雄穗中表达量较高,其中穗位叶中表达量最高;在脱水和PEG处理下,ZmFBL2在地上部分和地下部分均表现显著上调表达趋势;在冷处理条件下,ZmFBL2在地下部分的表达量无显著的变化趋势,但是在地上部分表达量有显著的上调表达趋势。在NaCl处理条件下,ZmFBL2在地上部分及地下部分均显示上调表达趋势,但未达到显著上调表达水平;在响应4种外源激素信号分子时,ZmFBL2对MJ处理无显著的诱导表达变化,而在应答2,4-D和ABA两个外源激素时均表现出上调表达模式;对SA处理表现出下调表达趋势。
3、通过同源克隆,获得玉米中10个MSK(maize skp1-like gene)基因,分别命名为MSK1-10。酵母双杂交结果表明,F-box蛋白(1-368氨基酸)可以与MSK1、MSK2、MSK9及ZmMADS6、ZmMADS7互作。F2(1-75氨基酸)可以与MSK1、MSK2、MSK5、MSK9、CKS及ZmMADS4、ZmMADS6、ZmMADS7互作。而F1(76-368氨基酸)、F3(1-38氨基酸)不能与任何MSK蛋白或ZmMADS蛋白互作。β-半乳糖苷酶显色检验分析也验证了以上互作结果的可靠性。
4、应用20%PEG处理在营养土中生长至10天的过表达转基因株系及野生型拟南芥,结果表明过表达转基因株系存活率明显高于野生型;在含有0.3μM ABA、150mM NaCl的MS固体培养基上,过表达转基因株系存活率明显高于野生型;而在含有350mM甘露醇MS固体培养基上,表达转基因株系与野生型存活率差异不明显。
5、在本研究的过程中还通过同源克隆的方法克隆得到干旱诱导表达的ZmCKS2和ZmBTF3b基因。ZmCKS2基因编码的蛋白ZmCKS2是细胞周期依赖性蛋白激酶的亚基。该基因启动子序列中含有多个与逆境胁迫相关的顺式作用元件;ZmCKS2基因在幼穗中表达量最高,并受干旱胁迫和茉莉酸甲酯处理上调表达,受冷胁迫和外源ABA处理下调表达。说明该基因可能在玉米应答逆境胁迫过程中起重要作用。ZmBTF3b基因编码的蛋白ZmBTF3b含有保守的新生多肽复合体(NAC)结构域;酵母单杂交试验表明该蛋白C末端具有转录激活功能。该基因启动子序列中有多个与逆境胁迫相关的顺式作用元件;表达模式分析表明该基因在花丝、幼穗、雄穗、幼根和开花期根中表达量相对较高,叶子中表达量相对较低,并且在根部受脱水干旱和PEG处理上调表达,受冷胁迫和NaCl处理下调表达。结果表明,该基因可能与玉米在应答逆境胁迫过程中起重要作用。
Water stress is one of the major environmental stresses that seriously affect plant growth and crop yield worldwide. In this study we cloned three drought induced genes ZmFBL2, ZmCKS2 and ZmBTF3b from our previous drought related SSH library.
ZmFBL2 is a F-box protein, contein a F-box domain and LRR domain. F-box domain was fistly found at the N-terminal region of cyclin F. At the N terminus of F-box proteins contained a conserved F-box motif (about 40~50 amino acid) and at the carboxy-terminal, F-box proteins often contained highly variable protein-protein interaction domains, such as WD40 repeat (Trp and Asp), Leu-rich repeat (LRR), tetratricopeptide repeat (TPR) and kelch repeat. In the past few years, many F-box proteins have been cloned in higher plants such as Arabidopsis, petunia, rice and maize. These F-box proteins played essential roles in flowering development, self-incompatibility, light signalling and clock control, plant hormone response pathways and abiotic stress. In this study, we isolated an drought induced gene ZmFBL2, a new F-box gene in maize. Its structure and functions was analized.
ZmCKS2 was predicted to contain a CKS (cyclin-dependent kinase regulatory subunit) structural domain, which was highly conserved in eukaryotes kingdom. ZmBTF3b encoded a putative transcription factor of 169 amino acids with a conserved NAC (nascent polypeptide-associated complex) domain.
The main results were as follows:
1. In this study we cloned the full-length cDNA of ZmFBL2, by using in silico and homology-based cloning techniques. The deduced protein of ZmFBL2 was predicted to contain a F-box structural domain, which was highly conserved in eukaryotes kingdom and a Leu-rich repeat (LRR) domains at the carboxy terminal. The promoter of ZmFBL2 was predicted to contain important regulatory elements including core promoter elements and drought, high salt, cold, SA, ABA, oxygen responsive, light regulation and embryo, endosperm, seeds development elements.
2. Real-time quantitative PCR (qRT-PCR) analysis revealed that ZmFBL2 was highly expressed in leaves at seedling stage, ear leaves, roots at flowering stage and tassels and was up-regulated under dehydration, PEG, cold, NaCl, 2,4-D and ABA treatment while was down-regulated under SA .
3. We cloned the full-length cDNA of ten MSK (maize skp1-like gene) genes by using in silico and homology-based cloning techniques. Yeast hybrid assay revealed that F-box protein interacted specifically with MSK1、MSK2、MSK9 and ZmMADS6、ZmMADS7 and the F-box domain was necessary.β-Galactosidase filter assays verified the reliability of the results.
4. Ten days old lins grown in organic fertilizer was treated by 20% PEG 6000. Survival rates of transgenic Arabidopsis was higher than WT Arabidopsis. On MS medium with ABA and NaCl, survival rates of transgenic lines was also higher than WT. But there was no obvious difference on MS medium with mannitol.
5. Two other drought induced genes ZmCKS2 and ZmBTF3b were cloned in this study. The deduced protein of ZmCKS2 was predicted to contain a CKS (cyclin-dependent kinase regulatory subunit) structural domain, which was highly conserved in eukaryotes kingdom. The promoter of ZmCKS2 was predicted to contain important regulatory elements including core promoter elements and drought, high salt, cold, ABA, light regulation, copper and oxygen responsive elements. Real-time quantitative PCR (qRT-PCR) analysis revealed that ZmCKS2 was highly expressed in ears while its expression was very low in roots and leaves at seedling stage and was up-regulated under drought or MeJA treatment but down-regulated under cold or ABA treatment. Under NaCl stress ZmCKS2 showed no obvious change. These results revealed that ZmCKS2 might be involved in multiple pathways of plants responding to different environmental conditions. The cloned ZmBTF3b gene encoded a putative transcription factor of 169 amino acids with a conserved NAC (nascent polypeptide-associated complex) domain. Transcriptional activity analysis showed that ZmBTF3b might be a functional transcription factor. Real-time quantitative PCR (qRT-PCR) analysis revealed that ZmBTF3b was highly expressed in silks, ears and tassels and was up-regulated under dehydration or PEG treatment while was down-regulated under cold, NaCl, ABA or SA. ZmBTF3b might have the transcriptional activation activity and play an essential role in response to abiotic stresses.
引文
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