水稻锌指蛋白基因RZF5和RZF71的克隆与功能分析
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摘要
水稻是重要的粮食作物，也是单子叶植物研究的模式植物。随着水稻基因组计划的完成，解析基因功能，进而根据其功能在作物遗传改良中加以利用日益成为后基因组学的重要研究内容。高盐、干旱和低温等环境胁迫是影响植物生长和作物高产的重要限制因素。本研究选择与逆境胁迫相关的TFIIIA型锌指蛋白基因进行研究，以期获得在作物抗逆性改良方面具有应用潜力的候选基因。本研究取得了以下研究进展：
利用生物信息学手段和RT-PCR方法从水稻幼苗组织中分离了两个新的TFIIIA型锌指蛋白基因RZF5和RZF71，其编码产物都含有两个典型的C2H2型锌指结构。RZF5编码171个氨基酸残基，分子量为17.95 kDa；RZF71编码产物为250个氨基酸残基，分子量为25 kDa。组织表达分析表明，RZF5和RZF71在水稻根、茎、叶和幼穗组织中均有不同程度的表达，RZF5在各组织中的表达丰度相差不大，而RZF71在根中表达丰度相对较高。诱导表达分析表明，在150 mmol·L~(-1) NaCl和20％PEG6000胁迫的水稻幼苗中，RZF5和RZF71的表达都显著增强，低温(4℃)和0.1 mmol·L~(-1) ABA处理对两个基因的表达量影响不大。
基因的电子定位结果显示RZF5位于水稻第1染色体长臂，RZF71位于第12染色体长臂。对RZF5和RZF71基因起始密码子上游1，500bp的基因组序列进行分析表明，它们的启动子区舍有多个逆境胁迫相关元件，其中RZF5的启动子区含有1个脱落酸应答元件ABRE(ABA response element)、2个干旱应答元件CRT／DREs(C-repeat／dehydration-response element)、5个MYB识别位点MRSs(sequence homologous to MYB recognition site)和1个W-box；在RZF71的启动子区含有1个干旱应答元件DRE、3个MYB识别位点序列MRSs、1个CBF表达诱导子ICE(inducer of CBF expression)识别元件和1个W-box。
将RZF5基因启动子区与GUS报告基因融合后转化水稻愈伤组织，GUS报告基因的表达受到150 mmol·L~(-1)NaCl和20％PEG6000的诱导，受低温(4℃)诱导不明显。将RZF5和RZF71基因编码区与GFP融合后构建融合表达载体，通过农杆菌介导法分别转化洋葱表皮细胞和水稻愈伤组织细胞，结果表明，融合蛋白仅在细胞核中表达。
构建了RZF5基因过量表达的表达载体，通过叶盘法将RZF5转化烟草，获得18株过量表达RZF5的转基因烟草植株，其中16株(T_0代)明显矮化，为对照株高(105cm)的40.9％-57.1％。对T_1代转基因烟草幼苗进行抗逆性分析表明：100 mmol·L~(-1)和200 mmol·L~(-1)NaCl处理15d后，植株的根长为未处理植株的36.1％和17.4％，与非转基因植株(15.3％、1.4％)相比差异极显著；在300 mmol·L~(-1)和500 mmol·L~(-1)甘露醇处理下，转基因植株的根长为未处理植株的98.0％和67.6％，与非转基因植株(62.2％、24.3％)相比差异极显著；在4℃条件下，转基因植株与非转基因植株的生长都受到明显抑制，其与正常温度下生长的植株相比相对鲜重分别为22.2％和22.4％无明显差异。
通过农杆菌介导法将RZF5和RZF71两个基因分别转化水稻，获得过量表达、抑制表达和发夹结构引起RNA干涉的转基因植株，25株过量表达RZF5的转基因水稻中，16株植林(T_0代)的平均株高为54.5cm，为非转基因植株株高的49.5％-61.6％。过量表达RZF71的转基因植株(T_0代)农艺性状没有明显变化。对转基因水稻的耐逆性分析表明，3个过量表达RZF5转基因水稻(T_1代)株系在冷胁迫36h后，电解质渗透率分别为8.54％、11.20％和1.70％与对照(84.18％)相比差异达到极显著水平；在含150mmol·L~(-1)NaCl的培养基上三个株系出苗率分别为79.5％、82.5％和92.5％，与对照(62.5％)相比差异极显著；4-5叶期的T_1代转基因幼苗用150 mmol·L~(-1)的NaCl处理7d后，其相对株高、相对干重和相对鲜重都明显高于对照。
花药培养作为一种组织培养技术，不仅已成为研究生理、生化和遗传常用的重要方法，而且对于生理学及分子生物学中基因调控、表达机理的阐明有重要意义；花药培养建立的双单倍体群体，是AFLP、RAPD、SSR等分子标记选择育种和基因图谱构建的理想材料。提高花药培养效率，一直是科技工作者进行研究的主题。本研究以黑壳子粳／苏御糯、铁杆青／苏御糯和薄稻／苏御糯3个籼粳杂交F_1及其亲本和粳稻品种武运粳8号为试验材料，在M_8基本培养基中，加入不同浓度的植物激素类物质和有机营养物，建立籼粳杂交F_1高效花药培养技术体系。结果表明：M_8+2mg·L~(-1)2，4-D+1mg·L~(-1) NAA+0.2 mg·L~(-1) KT为最佳诱导培养基，籼粳杂交F_1花药愈伤组织诱导率达4.1％-4.5％；在该培养基中添加水解酪蛋白或酵母提取物，能明显提高诱导率，增幅达20.5％-27.3％；M_8+2.5mg·L~(-1) 6-BA+0.5 mg·L~(-1)KT为最佳分化培养基，籼粳杂交F_1愈伤组织的绿苗分化率达14.3％-25.0％。
Rice is one of the most important food crops as well as a model plant formonocotyledon research. With the development of rice genome project, it becomesincreasingly a hotspot to analyze the genes function and to apply them in geneticengineering research. Environmental stresses such as cold, high salinity and drought aremajor obstacles affecting plant growth and crop productivity. Our research focused onidentifying the TFIIIA type zinc finger protein genes involved in abiotic stress, andexpecting to provide new targets for producing tolerance-enhanced transgenics. Noveltiesachieved in this study are as follows:
Two novel TFIIIA-type zinc finger protein genes RZF5 and RZF71 was identifiedfrom rice (Oryza sativa L. subs. japonica) by bioinformatics method and RT-PCR approach.Sequence analysis showed that RZF5 and RZF71 both contained two typical C2H2 zincfinger domains. RZF5 encodes a 17.95 kDa polypeptide with 171 amino acids and RZF71encodes a 25 kDa polypeptide with 250 amino acids. RT-PCR analysis showed that RZF5and RZF71 were constitutively expressed in various rice tissues including root, stem, leaveand spike, and strongly induced by 150 mmol·L~(-1)NaCl and 20％PEG6000. But both geneswere not induced by low temperature (4℃) and 0.1 mmol·L~(-1) ABA treatments.
Data analysis of rice genome shows that RZF5 is located on the long arm ofchromosome 1 and RZF71 is located on long arm of chromosome 12. About 1500bpsequences of the upstreams form the translation start codons of RZF5 gene and RZF71 genewere analyzed by MapInspector program, respectively. The results show that there are nineputative cis-acting elements related to abiotic stress in RZF5 promoter regions, includingone ABRE (abscisic acid response element), two CRT/DREs(C-repeat/dehydration-response elements), five MRSs (sequence homologous to MYBrecognition sites) and one W-box were found within RZF5 promoter, and there were sixelements in RZF71 including one CRT/DRE, three MRSs, one ICE (inducer of CBFexpression) element and one W-box.
About 1500 bp sequence in the promoter region of the RZF5 gene was fused to GUSreporter gene in pCAMBIA1301 vector and then was transformed into rice callus by Agrobacterium-mediated transformation. Histochemical analysis revealed, that the GUSactivity was induced both in the 150 mmol·L~(-1) NaCl and 20％PEG6000 treatments, but notin the cold (4℃) treatment. To investigate the subcellular localizations of RZF5 and RZF71protein, RZF5 and RZF71 gene were fused in the green fluorescence protein (GFP) reportergene, respectively. These fusion genes were introduced to onion epidermal cells and neecallus cells respectively. The results showed that the fusion proteins were localized in thenucleus, respectively, while the control was distributed throughout the cells.
To find out the functions of these two zinc finger proteins, we generated RZF5-sensetransgenic tobacco plants by Agrobacterium-rnediated transformation. Among eighteenplants of the T_0 generation, the growth of sixteen plants was obviously suppressedcompared to the non-transgenic ones. And the transgenic plants of the T_1 generation wereanalyzed for abiotic stresses. The measurements showed that the relative root length of thetransgenic seedlings were elongated 36.1％and 17.4％compared to the normal one under100 mmol·L~(-1) and 200 mmol·L~(-1) NaCl treatments, respectively, whereas the non-transgenicones could only elongated 15.3％and 1.4％, respectively. Under 300 mmol·L~(-1) and 500mmol·L~(-1) Mannitol stress, the relative root length of the transgenic seedlings elongated98.0％and 67.6％, respectively, whereas the non-transgenic ones could only elongated62.2％and 24.3％. There was no significant difference between the transgenic seedlings andthe non-transgenic ones under controlled cold stress. Thus, the overexpression of RZF5 inthe transgenic tobacoo plants confered tolerance to high salt and osmotic stress.
Transgenic RZF5 (sense, antisense, hairpin) rice plants of were generated byAgrobacterium-mediated transformation. The growth of RZF5-sense transgenic plants wasretarded. Average plant height of sixteen T_0 generation of RZF5-sense transgenic plantswere 40.9％to 57.1％, to that of non-transgenic plants. But transgenlc RZF71 (sense,antisense, hairpin) plants exhibited neither growth inhibition nor visible phenotypicalterations. Transgenic lines were analyzed for cold and NaCl stress in the T_1 generation.Electrolyte leakage rate and plant morphology were compared between transgenic lines anduntransformed after clod stress 36 h. The results showed the growth of three independentRZF5-sense lines were no visible cold symptom and electrolyte leakage rate (8.54％、11.20％and 1.70％) lower than those of untransformed (84.18％). Data collected forpercentage of germination showed three independent transgenic RZF5-sense lines weremore tolerant than untransformed and germination rate of three transgenic lines were79.5％, 82.5％and 92.5％, whereas untransformed were only 62.5％under 150 mmol·L~(-1) NaCl stress. Thus overexpression of RZF5 conferred tolerance to cold and salt stress whiletransgenic RZF71 rice plants did not. These results suggest RZF5 may be quite useful inenhancing plant tolerance to abiotic stress.
Anther culture has become a powerful method to research plant physiology,biochemistry and genetics. And it has been playing important roles to research genesregulation and expression mechanism by anther culture in physiology and molecularbiology. Doubled haploids population based on anther culture is perfect material formolecular marker assistant breeding and genetic mapping. It is a topic that how to improveanther culture efficiency for scientists. In this reseach, the anthers of F_1 progenies fromthree crosses between japonica and indica rice varieties and their parents, and one japonicavariety Wuyunjing 8 were used to induce the calli in the M_8 medium supplemented withdifferent concentrations and combinations of plant hormones and organic nutriments. Ahighly efficient system of anther culture was established for F_1 progenies between japonicaand indica rice varieties. The results showed that the highest rate of callus induction(4.1％-4.5在for japonica/indica F_1 hybrid ) were obtained in the M_8 medium supplementedwith 2,4-D (2 mg.L~(-1)), NAA (1mg.L~(-1)) and KT (0.2 mg.L~(-1)). The induction rate could befurther increased up to 20.5％-27.3％in this medium added with hydrolyzed casein andyeast extract; It could be obtained, the 14.3％-25.0％of plant regeneration rate(japonica/indica F_1) in the M_8 medium added with 6-BA (2.5 mg'L~(-1)) and KT (0.5mg.L~(-1)).
利用生物信息学手段和RT-PCR方法从水稻幼苗组织中分离了两个新的TFIIIA型锌指蛋白基因RZF5和RZF71，其编码产物都含有两个典型的C2H2型锌指结构。RZF5编码171个氨基酸残基，分子量为17.95 kDa；RZF71编码产物为250个氨基酸残基，分子量为25 kDa。组织表达分析表明，RZF5和RZF71在水稻根、茎、叶和幼穗组织中均有不同程度的表达，RZF5在各组织中的表达丰度相差不大，而RZF71在根中表达丰度相对较高。诱导表达分析表明，在150 mmol·L~(-1) NaCl和20％PEG6000胁迫的水稻幼苗中，RZF5和RZF71的表达都显著增强，低温(4℃)和0.1 mmol·L~(-1) ABA处理对两个基因的表达量影响不大。
基因的电子定位结果显示RZF5位于水稻第1染色体长臂，RZF71位于第12染色体长臂。对RZF5和RZF71基因起始密码子上游1，500bp的基因组序列进行分析表明，它们的启动子区舍有多个逆境胁迫相关元件，其中RZF5的启动子区含有1个脱落酸应答元件ABRE(ABA response element)、2个干旱应答元件CRT／DREs(C-repeat／dehydration-response element)、5个MYB识别位点MRSs(sequence homologous to MYB recognition site)和1个W-box；在RZF71的启动子区含有1个干旱应答元件DRE、3个MYB识别位点序列MRSs、1个CBF表达诱导子ICE(inducer of CBF expression)识别元件和1个W-box。
将RZF5基因启动子区与GUS报告基因融合后转化水稻愈伤组织，GUS报告基因的表达受到150 mmol·L~(-1)NaCl和20％PEG6000的诱导，受低温(4℃)诱导不明显。将RZF5和RZF71基因编码区与GFP融合后构建融合表达载体，通过农杆菌介导法分别转化洋葱表皮细胞和水稻愈伤组织细胞，结果表明，融合蛋白仅在细胞核中表达。
构建了RZF5基因过量表达的表达载体，通过叶盘法将RZF5转化烟草，获得18株过量表达RZF5的转基因烟草植株，其中16株(T_0代)明显矮化，为对照株高(105cm)的40.9％-57.1％。对T_1代转基因烟草幼苗进行抗逆性分析表明：100 mmol·L~(-1)和200 mmol·L~(-1)NaCl处理15d后，植株的根长为未处理植株的36.1％和17.4％，与非转基因植株(15.3％、1.4％)相比差异极显著；在300 mmol·L~(-1)和500 mmol·L~(-1)甘露醇处理下，转基因植株的根长为未处理植株的98.0％和67.6％，与非转基因植株(62.2％、24.3％)相比差异极显著；在4℃条件下，转基因植株与非转基因植株的生长都受到明显抑制，其与正常温度下生长的植株相比相对鲜重分别为22.2％和22.4％无明显差异。
通过农杆菌介导法将RZF5和RZF71两个基因分别转化水稻，获得过量表达、抑制表达和发夹结构引起RNA干涉的转基因植株，25株过量表达RZF5的转基因水稻中，16株植林(T_0代)的平均株高为54.5cm，为非转基因植株株高的49.5％-61.6％。过量表达RZF71的转基因植株(T_0代)农艺性状没有明显变化。对转基因水稻的耐逆性分析表明，3个过量表达RZF5转基因水稻(T_1代)株系在冷胁迫36h后，电解质渗透率分别为8.54％、11.20％和1.70％与对照(84.18％)相比差异达到极显著水平；在含150mmol·L~(-1)NaCl的培养基上三个株系出苗率分别为79.5％、82.5％和92.5％，与对照(62.5％)相比差异极显著；4-5叶期的T_1代转基因幼苗用150 mmol·L~(-1)的NaCl处理7d后，其相对株高、相对干重和相对鲜重都明显高于对照。
花药培养作为一种组织培养技术，不仅已成为研究生理、生化和遗传常用的重要方法，而且对于生理学及分子生物学中基因调控、表达机理的阐明有重要意义；花药培养建立的双单倍体群体，是AFLP、RAPD、SSR等分子标记选择育种和基因图谱构建的理想材料。提高花药培养效率，一直是科技工作者进行研究的主题。本研究以黑壳子粳／苏御糯、铁杆青／苏御糯和薄稻／苏御糯3个籼粳杂交F_1及其亲本和粳稻品种武运粳8号为试验材料，在M_8基本培养基中，加入不同浓度的植物激素类物质和有机营养物，建立籼粳杂交F_1高效花药培养技术体系。结果表明：M_8+2mg·L~(-1)2，4-D+1mg·L~(-1) NAA+0.2 mg·L~(-1) KT为最佳诱导培养基，籼粳杂交F_1花药愈伤组织诱导率达4.1％-4.5％；在该培养基中添加水解酪蛋白或酵母提取物，能明显提高诱导率，增幅达20.5％-27.3％；M_8+2.5mg·L~(-1) 6-BA+0.5 mg·L~(-1)KT为最佳分化培养基，籼粳杂交F_1愈伤组织的绿苗分化率达14.3％-25.0％。
Rice is one of the most important food crops as well as a model plant formonocotyledon research. With the development of rice genome project, it becomesincreasingly a hotspot to analyze the genes function and to apply them in geneticengineering research. Environmental stresses such as cold, high salinity and drought aremajor obstacles affecting plant growth and crop productivity. Our research focused onidentifying the TFIIIA type zinc finger protein genes involved in abiotic stress, andexpecting to provide new targets for producing tolerance-enhanced transgenics. Noveltiesachieved in this study are as follows:
Two novel TFIIIA-type zinc finger protein genes RZF5 and RZF71 was identifiedfrom rice (Oryza sativa L. subs. japonica) by bioinformatics method and RT-PCR approach.Sequence analysis showed that RZF5 and RZF71 both contained two typical C2H2 zincfinger domains. RZF5 encodes a 17.95 kDa polypeptide with 171 amino acids and RZF71encodes a 25 kDa polypeptide with 250 amino acids. RT-PCR analysis showed that RZF5and RZF71 were constitutively expressed in various rice tissues including root, stem, leaveand spike, and strongly induced by 150 mmol·L~(-1)NaCl and 20％PEG6000. But both geneswere not induced by low temperature (4℃) and 0.1 mmol·L~(-1) ABA treatments.
Data analysis of rice genome shows that RZF5 is located on the long arm ofchromosome 1 and RZF71 is located on long arm of chromosome 12. About 1500bpsequences of the upstreams form the translation start codons of RZF5 gene and RZF71 genewere analyzed by MapInspector program, respectively. The results show that there are nineputative cis-acting elements related to abiotic stress in RZF5 promoter regions, includingone ABRE (abscisic acid response element), two CRT/DREs(C-repeat/dehydration-response elements), five MRSs (sequence homologous to MYBrecognition sites) and one W-box were found within RZF5 promoter, and there were sixelements in RZF71 including one CRT/DRE, three MRSs, one ICE (inducer of CBFexpression) element and one W-box.
About 1500 bp sequence in the promoter region of the RZF5 gene was fused to GUSreporter gene in pCAMBIA1301 vector and then was transformed into rice callus by Agrobacterium-mediated transformation. Histochemical analysis revealed, that the GUSactivity was induced both in the 150 mmol·L~(-1) NaCl and 20％PEG6000 treatments, but notin the cold (4℃) treatment. To investigate the subcellular localizations of RZF5 and RZF71protein, RZF5 and RZF71 gene were fused in the green fluorescence protein (GFP) reportergene, respectively. These fusion genes were introduced to onion epidermal cells and neecallus cells respectively. The results showed that the fusion proteins were localized in thenucleus, respectively, while the control was distributed throughout the cells.
To find out the functions of these two zinc finger proteins, we generated RZF5-sensetransgenic tobacco plants by Agrobacterium-rnediated transformation. Among eighteenplants of the T_0 generation, the growth of sixteen plants was obviously suppressedcompared to the non-transgenic ones. And the transgenic plants of the T_1 generation wereanalyzed for abiotic stresses. The measurements showed that the relative root length of thetransgenic seedlings were elongated 36.1％and 17.4％compared to the normal one under100 mmol·L~(-1) and 200 mmol·L~(-1) NaCl treatments, respectively, whereas the non-transgenicones could only elongated 15.3％and 1.4％, respectively. Under 300 mmol·L~(-1) and 500mmol·L~(-1) Mannitol stress, the relative root length of the transgenic seedlings elongated98.0％and 67.6％, respectively, whereas the non-transgenic ones could only elongated62.2％and 24.3％. There was no significant difference between the transgenic seedlings andthe non-transgenic ones under controlled cold stress. Thus, the overexpression of RZF5 inthe transgenic tobacoo plants confered tolerance to high salt and osmotic stress.
Transgenic RZF5 (sense, antisense, hairpin) rice plants of were generated byAgrobacterium-mediated transformation. The growth of RZF5-sense transgenic plants wasretarded. Average plant height of sixteen T_0 generation of RZF5-sense transgenic plantswere 40.9％to 57.1％, to that of non-transgenic plants. But transgenlc RZF71 (sense,antisense, hairpin) plants exhibited neither growth inhibition nor visible phenotypicalterations. Transgenic lines were analyzed for cold and NaCl stress in the T_1 generation.Electrolyte leakage rate and plant morphology were compared between transgenic lines anduntransformed after clod stress 36 h. The results showed the growth of three independentRZF5-sense lines were no visible cold symptom and electrolyte leakage rate (8.54％、11.20％and 1.70％) lower than those of untransformed (84.18％). Data collected forpercentage of germination showed three independent transgenic RZF5-sense lines weremore tolerant than untransformed and germination rate of three transgenic lines were79.5％, 82.5％and 92.5％, whereas untransformed were only 62.5％under 150 mmol·L~(-1) NaCl stress. Thus overexpression of RZF5 conferred tolerance to cold and salt stress whiletransgenic RZF71 rice plants did not. These results suggest RZF5 may be quite useful inenhancing plant tolerance to abiotic stress.
Anther culture has become a powerful method to research plant physiology,biochemistry and genetics. And it has been playing important roles to research genesregulation and expression mechanism by anther culture in physiology and molecularbiology. Doubled haploids population based on anther culture is perfect material formolecular marker assistant breeding and genetic mapping. It is a topic that how to improveanther culture efficiency for scientists. In this reseach, the anthers of F_1 progenies fromthree crosses between japonica and indica rice varieties and their parents, and one japonicavariety Wuyunjing 8 were used to induce the calli in the M_8 medium supplemented withdifferent concentrations and combinations of plant hormones and organic nutriments. Ahighly efficient system of anther culture was established for F_1 progenies between japonicaand indica rice varieties. The results showed that the highest rate of callus induction(4.1％-4.5在for japonica/indica F_1 hybrid ) were obtained in the M_8 medium supplementedwith 2,4-D (2 mg.L~(-1)), NAA (1mg.L~(-1)) and KT (0.2 mg.L~(-1)). The induction rate could befurther increased up to 20.5％-27.3％in this medium added with hydrolyzed casein andyeast extract; It could be obtained, the 14.3％-25.0％of plant regeneration rate(japonica/indica F_1) in the M_8 medium added with 6-BA (2.5 mg'L~(-1)) and KT (0.5mg.L~(-1)).
引文
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