枸杞雄性不育花药差异蛋白质组分析及3个相关基因的克隆
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摘要
枸杞(Lycium barbarum L.)是一种重要的药食同源经济作物,杂种优势的利用是提高枸杞产量、缓解品质与产量矛盾、增强抗(耐)逆性的有效途径,而雄性不育是杂种优势利用的一种重要遗传工具和途径,明确枸杞雄性不育发生的机理,有重要的理论意义和应用价值。
本研究以枸杞雄性不育突变体宁杞5号和野生型宁杞1号为试验材料,采用双向荧光差异凝胶电泳技术和质谱技术对枸杞四分体早期花药进行了差异蛋白质组学研究。获得了可能与花药发育及雄性不育相关的差异表达蛋白,并对2个差异表达蛋白14-3-3和查尔酮合成酶(CHS)进行了cDNA克隆、生物信息学分析、相对定量表达及功能初步分析；克隆了枸杞R2R3类MYB基因LbMYB103的开放阅读框(ORF),并进行了基因组织表达分析与生物信息学分析。主要研究结果如下：
1.优化的枸杞花药总蛋白质提取方法和双向电泳体系,可有效解析枸杞花药蛋白质组。该体系是采用TCA-丙酮沉淀结合Tris-HCl法提取枸杞花药蛋白质,13cm胶条、350μg左右的上样量、含有2mol/L硫脲的裂解液,硝酸银染色,可得到重复性好、质量高的2-DE图谱,蛋白点主要集中在pH4～7,15.0～85.0kD范围内。
2.采用2D-DIGE技术,13cm pH4-7IPG胶条对枸杞花药蛋白质进行分离,从分离的1760多个蛋白点中选取差异表达量1.5倍以上的52个蛋白点(p<0.05)进行MALDI-TOF/TOF-MS/MS质谱分析,45个差异蛋白点成功鉴定为40个不同的蛋白质,成功率为86.0%。通过对这些鉴定蛋白注释功能的分析及分类,差异表达蛋白可分为参与：蛋白质合成和折叠(20%)、糖和能量代谢(18%)、信号转导(9%)、逆境响应(9%)、氨基酸代谢(7%)、功能未知的蛋白(7%)、花药发育(4%)、转录因子(3%)、脂肪酸代谢(2%)、核酸代谢(2%)和类黄酮合成相关的酶(2%)等。枸杞花药的发育是受一个复杂的网络调控的,多条代谢途径中多个基因的差异表达影响了花粉的育性。
3.经质谱鉴定,相对表达量有显著差异的蛋白点中,蛋白点14与26、27、28分别为谷氨酰胺合成酶(GS)和抗坏血酸过氧化物酶(APX),酶活性分析结果表明,雄性不育突变体花药GS和APX活性显著低于野生型,活性仅分别为野生型的22%和33%,酶活性变化和蛋白质差异表达变化的趋势一致。
4.枸杞花药发育及雄性不育相关差异表达蛋白基因的克隆、表达及生物信息学分析。
①采用RACE技术克隆了枸杞花药Lb14-3-3基因的cDNA序列,该序列长1005bp,其ORF长768bp,编码256个氨基酸。荧光定量结果表明：Lb14-3-3基因在花药、果实和花中优势表达,而在根茎叶中的表达量很低；在野生型和雄性不育突变体四分体早期花药中,Lb14-3-3基因转录水平和翻译水平的变化较相似,表现为在雄性不育突变体中表达量下降,推测Lb14-3-3在枸杞花药发育过程中具有调控功能。
②克隆了枸杞花药LbCHS基因的cDNA序列,该序列长1448bp,其ORF长1167bp,编码389个氨基酸。LbCHS属于CHS超基因家族。LbCHS基因在花药和花中优势表达,果实中的表达量很低,根茎叶中检测不到其转录本；在四分体早期花药中,LbCHS基因转录水平和翻译水平的变化较相似,表现为在雄性不育突变体中的相对表达量下降,由此初步推测LbCHS可能参与枸杞花药发育和花粉粒的形成。
5.分离了枸杞花药LbMYB103基因的ORF,共有975bp,编码324个氨基酸。Blastp分析表明,LbMYB103基因编码具有R2R3结构域的MYB转录因子家族蛋白。LbMYB103在花器官特异表达。在四分体早期花药中,表现为在雄性不育突变体中的相对表达量下降,初步推测LbMYB103可能参与调控枸杞花药的发育。
Wolfberry (Lycium barbarum L.) is one of important edible economic crops. The utilization of heterosis is the effective way to improve wolfberry production, ease the contradictions of quality and yield and enhance stress resistance (tolerance). Male sterility is an important genetic tool in utilization of heterosis. Therefore, understanding of male sterility mechanism has important theoretical significance and application value.
Using2D-DIGE and mass spectrometry technolgies, we studied differential proteome of male sterile wolfberry anther at the stage of early tetrad. We obtained differently expressed proteins which may be related with anther development and male sterility. Two of those differentially expressed proteins, Lb14-3-3and LbCHS genes cDNA were isolated, analyzied by bioinformatics and relatively quantified by qRT-PCR. Also, we cloned the R2R3MYB geae LbMYB103open reading frame (ORF) of wolfberry. The main results are as follows:
1. We optimized two-dimensional electrophoresis conditions and anther total protein extraction method of wolfberry. Using TCA-acetone precipitation combined Tris-HCl to extract wolfberry anther protein, with13cm pH4-7srips, about350μg loading quantity, lysis buffer containing2mol/L thiourea and silver nitrate staining can obtain reproducible, high-quality2-DE maps of anther protein of wolfberry. Anther proteins are mainly concentrated in the pH4-7and15.0-85.0kDa range. This2-DE condition can be effectively applied to analysis the wolfberry anther proteome.
2. Using2D-DIGE technology,13cm pH4-7IPG strips, over1760protein spots of wolfberry anther were separated, and52protein spots differentially expressed more than1.5folds (p<0.05) were selected to further analysis by MALDI-TOF/TOF-MS/MS, among which,45protein spots were successfully identified as40different proteins, with a success rate of86.0%. By analysis their function annotations of these successfully identified proteins, we classified these proteins. They were involved in protein synthesis and folding (20%), carbohydrate and energy metabolism (18%), signal transduction (9%), stress response (9%), amino acid metabolism (7%), unknown function (7%), anther development (4%), transcription factors (3%), fatty acid metabolism (2%), nucleic acid metabolism (2%) and flavonoid synthesis (2%). The wolfberry anther development is subject to a complex regulatory networks, multiple genes differentially expressed in multiple metabolic pathways affect pollen fertility.
3. Among significant differently expressed protein spots, spot No.14and26,27,28were identified as glutamine synthetase (GS) and ascorbate peroxidase (APX) respectively by mass spectrometry. Enzymes activity analysis results showed that in male sterile mutant anthers, GS and APX activities were only about22%and33%of that in the wild type, respectively.
4. Cloning and analysis of expressions of differentially expressed proteins related anther development and male sterility in wolfberry.
①By RACE technology, Lb14-3-3gene was cloned and its cDNA full-length was1005bp, its ORF was768bp, which encoded256amino acids. At transcription level, Lb14-3-3gene was predominantly expressed in anther, fruit and flower, and with a low expression in the roots, stems and leaves. The transcription level of male sterile mutant anther was lower than that of the wild type at the early tetrad stage, which was consistent with the change at protein level.
②The LbCHS gene was cloned with full-length of1448bp and its ORF was1167bp which encoded389ammo acids. LbCHS belongs to the CHS gene family. Gene expression analysis showed that LbCHS was highly expressed in reproductive organs of wolfberry, including flower and anthers, and low in the fruit. It can not be detected in vegetative organs. At early tetrad stage, LbCHS transcription level declined in male sterility relative to the wild type, which was similar to the trend at protein level.
5. The ORF sequence of LbMYB103gene was cloned with975bp encoding324amino acids. Blastp analysis showed that, LbMYB103genes encoded protein having R2R3domain of MYB transcription factor family. Quantitative real-time RT-PCR analysis indicated that LbMYB103mRNA accumulated with high level in flowers and anthers, and no transcripts detected in roots, stems, leaves and fruits. The relative expression level in male sterile mutant was declined relative to the wild type in early tetrads. From the preliminary analysis above, we speculate LbMYB103may be involved in the regulation of wolfberry anther development.
本研究以枸杞雄性不育突变体宁杞5号和野生型宁杞1号为试验材料,采用双向荧光差异凝胶电泳技术和质谱技术对枸杞四分体早期花药进行了差异蛋白质组学研究。获得了可能与花药发育及雄性不育相关的差异表达蛋白,并对2个差异表达蛋白14-3-3和查尔酮合成酶(CHS)进行了cDNA克隆、生物信息学分析、相对定量表达及功能初步分析；克隆了枸杞R2R3类MYB基因LbMYB103的开放阅读框(ORF),并进行了基因组织表达分析与生物信息学分析。主要研究结果如下：
1.优化的枸杞花药总蛋白质提取方法和双向电泳体系,可有效解析枸杞花药蛋白质组。该体系是采用TCA-丙酮沉淀结合Tris-HCl法提取枸杞花药蛋白质,13cm胶条、350μg左右的上样量、含有2mol/L硫脲的裂解液,硝酸银染色,可得到重复性好、质量高的2-DE图谱,蛋白点主要集中在pH4～7,15.0～85.0kD范围内。
2.采用2D-DIGE技术,13cm pH4-7IPG胶条对枸杞花药蛋白质进行分离,从分离的1760多个蛋白点中选取差异表达量1.5倍以上的52个蛋白点(p<0.05)进行MALDI-TOF/TOF-MS/MS质谱分析,45个差异蛋白点成功鉴定为40个不同的蛋白质,成功率为86.0%。通过对这些鉴定蛋白注释功能的分析及分类,差异表达蛋白可分为参与：蛋白质合成和折叠(20%)、糖和能量代谢(18%)、信号转导(9%)、逆境响应(9%)、氨基酸代谢(7%)、功能未知的蛋白(7%)、花药发育(4%)、转录因子(3%)、脂肪酸代谢(2%)、核酸代谢(2%)和类黄酮合成相关的酶(2%)等。枸杞花药的发育是受一个复杂的网络调控的,多条代谢途径中多个基因的差异表达影响了花粉的育性。
3.经质谱鉴定,相对表达量有显著差异的蛋白点中,蛋白点14与26、27、28分别为谷氨酰胺合成酶(GS)和抗坏血酸过氧化物酶(APX),酶活性分析结果表明,雄性不育突变体花药GS和APX活性显著低于野生型,活性仅分别为野生型的22%和33%,酶活性变化和蛋白质差异表达变化的趋势一致。
4.枸杞花药发育及雄性不育相关差异表达蛋白基因的克隆、表达及生物信息学分析。
①采用RACE技术克隆了枸杞花药Lb14-3-3基因的cDNA序列,该序列长1005bp,其ORF长768bp,编码256个氨基酸。荧光定量结果表明：Lb14-3-3基因在花药、果实和花中优势表达,而在根茎叶中的表达量很低；在野生型和雄性不育突变体四分体早期花药中,Lb14-3-3基因转录水平和翻译水平的变化较相似,表现为在雄性不育突变体中表达量下降,推测Lb14-3-3在枸杞花药发育过程中具有调控功能。
②克隆了枸杞花药LbCHS基因的cDNA序列,该序列长1448bp,其ORF长1167bp,编码389个氨基酸。LbCHS属于CHS超基因家族。LbCHS基因在花药和花中优势表达,果实中的表达量很低,根茎叶中检测不到其转录本；在四分体早期花药中,LbCHS基因转录水平和翻译水平的变化较相似,表现为在雄性不育突变体中的相对表达量下降,由此初步推测LbCHS可能参与枸杞花药发育和花粉粒的形成。
5.分离了枸杞花药LbMYB103基因的ORF,共有975bp,编码324个氨基酸。Blastp分析表明,LbMYB103基因编码具有R2R3结构域的MYB转录因子家族蛋白。LbMYB103在花器官特异表达。在四分体早期花药中,表现为在雄性不育突变体中的相对表达量下降,初步推测LbMYB103可能参与调控枸杞花药的发育。
Wolfberry (Lycium barbarum L.) is one of important edible economic crops. The utilization of heterosis is the effective way to improve wolfberry production, ease the contradictions of quality and yield and enhance stress resistance (tolerance). Male sterility is an important genetic tool in utilization of heterosis. Therefore, understanding of male sterility mechanism has important theoretical significance and application value.
Using2D-DIGE and mass spectrometry technolgies, we studied differential proteome of male sterile wolfberry anther at the stage of early tetrad. We obtained differently expressed proteins which may be related with anther development and male sterility. Two of those differentially expressed proteins, Lb14-3-3and LbCHS genes cDNA were isolated, analyzied by bioinformatics and relatively quantified by qRT-PCR. Also, we cloned the R2R3MYB geae LbMYB103open reading frame (ORF) of wolfberry. The main results are as follows:
1. We optimized two-dimensional electrophoresis conditions and anther total protein extraction method of wolfberry. Using TCA-acetone precipitation combined Tris-HCl to extract wolfberry anther protein, with13cm pH4-7srips, about350μg loading quantity, lysis buffer containing2mol/L thiourea and silver nitrate staining can obtain reproducible, high-quality2-DE maps of anther protein of wolfberry. Anther proteins are mainly concentrated in the pH4-7and15.0-85.0kDa range. This2-DE condition can be effectively applied to analysis the wolfberry anther proteome.
2. Using2D-DIGE technology,13cm pH4-7IPG strips, over1760protein spots of wolfberry anther were separated, and52protein spots differentially expressed more than1.5folds (p<0.05) were selected to further analysis by MALDI-TOF/TOF-MS/MS, among which,45protein spots were successfully identified as40different proteins, with a success rate of86.0%. By analysis their function annotations of these successfully identified proteins, we classified these proteins. They were involved in protein synthesis and folding (20%), carbohydrate and energy metabolism (18%), signal transduction (9%), stress response (9%), amino acid metabolism (7%), unknown function (7%), anther development (4%), transcription factors (3%), fatty acid metabolism (2%), nucleic acid metabolism (2%) and flavonoid synthesis (2%). The wolfberry anther development is subject to a complex regulatory networks, multiple genes differentially expressed in multiple metabolic pathways affect pollen fertility.
3. Among significant differently expressed protein spots, spot No.14and26,27,28were identified as glutamine synthetase (GS) and ascorbate peroxidase (APX) respectively by mass spectrometry. Enzymes activity analysis results showed that in male sterile mutant anthers, GS and APX activities were only about22%and33%of that in the wild type, respectively.
4. Cloning and analysis of expressions of differentially expressed proteins related anther development and male sterility in wolfberry.
①By RACE technology, Lb14-3-3gene was cloned and its cDNA full-length was1005bp, its ORF was768bp, which encoded256amino acids. At transcription level, Lb14-3-3gene was predominantly expressed in anther, fruit and flower, and with a low expression in the roots, stems and leaves. The transcription level of male sterile mutant anther was lower than that of the wild type at the early tetrad stage, which was consistent with the change at protein level.
②The LbCHS gene was cloned with full-length of1448bp and its ORF was1167bp which encoded389ammo acids. LbCHS belongs to the CHS gene family. Gene expression analysis showed that LbCHS was highly expressed in reproductive organs of wolfberry, including flower and anthers, and low in the fruit. It can not be detected in vegetative organs. At early tetrad stage, LbCHS transcription level declined in male sterility relative to the wild type, which was similar to the trend at protein level.
5. The ORF sequence of LbMYB103gene was cloned with975bp encoding324amino acids. Blastp analysis showed that, LbMYB103genes encoded protein having R2R3domain of MYB transcription factor family. Quantitative real-time RT-PCR analysis indicated that LbMYB103mRNA accumulated with high level in flowers and anthers, and no transcripts detected in roots, stems, leaves and fruits. The relative expression level in male sterile mutant was declined relative to the wild type in early tetrads. From the preliminary analysis above, we speculate LbMYB103may be involved in the regulation of wolfberry anther development.
引文
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