棉花丝裂原活化蛋白激酶基因GbMPK3的功能鉴定
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摘要
丝裂原活化蛋白激酶(MAPK)信号级联广泛存在于各种真核生物体内,在细胞信号传导的过程中具有重要作用。本研究从海岛棉Hai7124根系中分离到一个在受黄萎病菌侵染时上调表达的MAPK类基因GbMPK3,并分别在本氏烟和陆地棉YZ1中对其在植物抗非生物胁迫和生物胁迫中的功能和调控机理进行了分析,取得的主要结果如下：
1. GbMPK3的克隆及序列分析
从海岛棉Hai7124受黄萎病菌V991处理的RNA-seq数据库中分离到一条在黄萎病菌侵染时表达量明显变化的EST序列(CL1Contig1773),通过RACE和Genome-walking获得该基因的全长和启动子序列。氨基酸序列比对和系统进化树分析发现其编码的蛋白具有MAPK的典型特征,属于MAPK基因家族TEY亚家族A组的成员,并与拟南芥MPK3的同源度为68%,因此将其命名为GbMPK3。将GbMPK3蛋白的N端融合GFP,瞬时转化烟草表皮细胞,结果显示GbMPK3蛋白主要定位于细胞核。
2. GbMPK3的表达模式分析利用RT-PCR和qPCR分析了GbMPK3的表达模式。结果表明GbMPK3的表达水平较低且具有组织特异性,在根系、花瓣、花药和纤维中表达量相对较高。GbMPK3在棉花根系中的表达还受到冷、高温、盐、甲基紫精(MV)和脱水等非生物逆境、水杨酸(SA)等激素以及黄萎病菌侵染的诱导。
3.超量表达GbMPK3提高了烟草对干旱和氧化胁迫的耐受性构建了GbMPK3基因超量表达载体并分别转化烟草和棉花。通过抗性筛选和分子鉴定分别获得了高量表达GbMPK3的转基因烟草和棉花株系。对野生型和转基因烟草在成株期进行干旱胁迫处理,发现转基因烟草具有更强的耐旱性。表现为转基因系复水之后的株高和存活率显著高于野生型。生理生化分析发现超表达GbMPK3提高了烟草在干旱胁迫下抗氧化酶APX的基因表达和酶活性,从而减少了烟草细胞内的活性氧积累。用MV处理烟草种子和成株期烟草叶片,发现MV对野生型和转基因系都造成了氧化胁迫,但是野生型对MV更敏感。MV处理之后野生型烟草种子的萌发率和成株期叶盘的叶绿素含量均显著低于转基因植株。RT-PCR分析结果显示,GbMPK3超表达的烟草在MV处理后抗氧化酶APX的基因表达量高于野生型。推测超量表达GbMPK3增强烟草对干旱和氧化胁迫的耐受性可能与提高抗氧化酶的表达和活性有关。
4.超量表达GbMPK3影响了烟草对烟草花叶病毒(TMV)和灰霉病菌的抗性对超量表达GbMPK3基因的烟草转基因系和野生型进行病原菌接种处理。转基因烟草对TMV的抗性增强,超表达GbMPK3降低了TMV在烟草中扩散和繁殖的速率；而接种灰霉病菌后转基因烟草则表现出更感病的表型。
5.超量表达GbMPK3降低了烟草和棉花对黄萎病菌的抗性对超量表达GbMPK3的烟草和棉花转基因材料进行黄萎病菌接种实验,发现转基因烟草和棉花的抗病性都显著降低。受黄萎病菌侵染后,野生型和转基因材料都表现出叶片萎蔫、黄化等黄萎病的典型病征,但野生型的叶片黄化与萎蔫出现的时期晚于转基因系,具体表现为野生型的发病率和病指显著低于转基因系。利用VIGS方法抑制棉花中GbMPK3的表达,对GbMPK3沉默的棉花进行接种实验。接种黄萎病菌后,对照材料和抑制GbMPK3表达的材料之间没有观察到抗病性的差异,两者在叶片黄化和脱落程度上无明显区别。说明抑制(GbMPK3表达对棉花抗黄萎病的影响不大,而超量表达GbMPK3显著减弱了植物对黄萎病菌的抗病性。
6. GbMPK3影响黄萎病菌侵染下棉花SA相关基因的表达qPCR分析显示,无论是GbMPK3抑制表达、GbMPK3超量表达还是野生型的棉花根系中SA相关基因,如WRKY70、 PR1和PR5等,在接种5天后相对于接水处理的对照组都显著上调。与野生型相比,GbMPK3超量表达转基因系中WRKY70,PR1和PR5等基因的表达水平显著升高；而抑制GbMPK3表达并未影响PRl和PR5的表达。这些结果说明棉花品种YZ1受黄萎病菌侵染后SA路径被激活,GbMPK3的超量表达增强了SA信号路径的激活水平。现有的一些研究结果表明,JA可能正调控植物对黄萎病菌的抗性,SA和JA信号路径通过WRKY70、 NPR1等关键因子相互拮抗。因此,SA也可能影响棉花对黄萎病菌的抗性,但SA调控棉花抗性的具体机理、超量表达GbMPK3降低陆地棉对黄萎病菌的抗性与其显著升高的SA信号传导水平是否关联还有待进一步的研究。
Mitogen-activated protein kinase (MAPK) cascades are highly conserved signaling modules found in all eukaryotes, and they play essential roles in cell signal transduction. A mitogen-activated protein kinase gene, GbMPK3, was induced upon Verticillium dahliae infection. In this work, GbMPK3was isolated from sea-island cotton Hai7124and its functional roles in response to different abiotic and biotic stress conditions were analyzed in both tobacco (Nicotiana benthamiana) and upland cotton YZ1. The main results of this work were as follows:
1. Cloning and characterization of GbMPK3An EST (CLlContig1773) putatively encoded a MAPK, GbMPK3, was isolated based on the results of RNA-seq of sea-island cotton Hai7124inoculation with V. dahliae. The full-length of cDNA and DNA sequences were obtained through RACE and Genome-walking. Sequence similarity search and phylogenetic analysis revealed that GbMPK3was classified as a group A MAPK. The35S-GFP::GbMPK3constructs were completed and introduced into tobacco leaves to investigate the subcellular localization of GbMPK3, and the result indicated that GbMPK3protein mainly accumulated in the nucleus.2. Expression profile of GbMPK3
The expression level of GbMPK3in cotton tissues under normal condition was evaluated by RT-PCR, and the results showed that GbMPK3expression level was low in all tissues and preferentially expressed in roots, petals, anthers and fibers. qPCR was employed to determine whether the expression of GbMPK3was affected by hormones and stresses, and GbMPK3was up-regulated upon cold (4℃), heat (42℃),10μmol/L MV,200mmol/L NaCl, dehydration, hormones (such as1mmol/L SA) and V. dahliae treatments in our study.
3. Overexpression of GbMPK3enhances drought and oxidative stress tolerance in tobacco
GbMPK3was introduced into tobacco and cotton to generate transgenic plants with constitutively higher expression of GbMPK3. Homozygous (T2) transgenic tobacco were conferred with enhanced drought tolerance, reduced water loss during drought treatment, improved plant height and survival rates after re-watering. Additionally, the gene expression levels and enzymatic activity of antioxidant enzymes were more strongly induced with depressed hydrogen peroxide accumulation in GbMPK3-overexpressing tobacco compared with wild-type under drought condition. Furthermore, observation of seed germination and leaf morphology showed that tolerance of transgenic plants to MV was improved due to increased antioxidant enzyme expression, suggesting that GbMPK3may positively regulate drought tolerance through enhanced reactive oxygen species scavenging ability.
4. Overexpression of GbMPK3affects tobacco disease resistance to Tobacco mosaic virus (TMV) and Botrytis cinerea
Wilt-type and transgenic tobacco were inoculated with different types of plant pathogens. Transgenic tobacco enhances resistance against TMV with reduced spreading rate and reproduction of TMV. Yet overexpression of GbMPK3significantly reduces tobacco resistance to the fungal pathogen Botrytis cinerea.
5. Overexpression of GbMPK3reduces V. dahliae resistance in tobacco and cotton
Wilt-type and transgenic plants were inoculated with V. dahliae to investigate the involvement of GbMPK3in plant defence. GbMPK3overexpression tobacco and cotton shows enhanced disease susceptiblity to V. dahliae strain V991. Both wilt-type and transgenic plants shows typical symptoms of verticillium wilt several days post V. dahliae infection, including wilting and yellowing leaves. Whereas the disease symptoms appeared early in GbMPK3-overexpression plants, resulting in elevated rate of diseased plants and disease index compared to wilt-type. The V. dahliae resistance was also analyzed in VIGS built GbMPK3-silencing plants. Silence of GbMPK3in cotton did not affect pathogen invasion, but slightly reduces yellowing and wilting leaves of diseased plants, as show by the decreased disease index. Therefore, while the effect of GbMPK3-silencing on cotton resistance to V. dahliae turns out to be weak, GbMPK3overexpression could significantly enhances V. dahliae susceptibility in both tobacco and cotton.
6. GbMPK3affects SA-related gene expression in cotton
qPCR analysis showed that the transcripts of SA-related genes (including the transcription factor WRKY70and two maker genes PR1and PR5) were accumulated in both wild-type, GbMPK3-silencing and overexpression cotton roots at day5post V. dahliae inoculation, suggested an up-regulation role of SA signaling in susceptible cotton cultivar YZ1. The elevated level of PR1and PR5was almost the same in wild-type and GbMPK3-silencing plants, yet transcripts of WRKY70, PR1and PR5were higher in GbMPK3overexpression cotton, indicating that GbMPK3overexpression may enhances V. dahliae induced SA signaling transduction. In previous studies, researches suggested JA signaling acted as a positively regulator in plants defense to V. dahliae. SA signaling is usually known antagonistic to JA signaling. Many important genes in regulating SA and JA signaling were reported, such as WRKY70and NPR1. Thus, SA may also participate in the complex signal networks of V. dahliae resistance in plants. But the mechanism of SA involved in cotton defense and the relation between elevated SA-related defence gene expression and reduced V. dahliae resistance still need to be studied further.
1. GbMPK3的克隆及序列分析
从海岛棉Hai7124受黄萎病菌V991处理的RNA-seq数据库中分离到一条在黄萎病菌侵染时表达量明显变化的EST序列(CL1Contig1773),通过RACE和Genome-walking获得该基因的全长和启动子序列。氨基酸序列比对和系统进化树分析发现其编码的蛋白具有MAPK的典型特征,属于MAPK基因家族TEY亚家族A组的成员,并与拟南芥MPK3的同源度为68%,因此将其命名为GbMPK3。将GbMPK3蛋白的N端融合GFP,瞬时转化烟草表皮细胞,结果显示GbMPK3蛋白主要定位于细胞核。
2. GbMPK3的表达模式分析利用RT-PCR和qPCR分析了GbMPK3的表达模式。结果表明GbMPK3的表达水平较低且具有组织特异性,在根系、花瓣、花药和纤维中表达量相对较高。GbMPK3在棉花根系中的表达还受到冷、高温、盐、甲基紫精(MV)和脱水等非生物逆境、水杨酸(SA)等激素以及黄萎病菌侵染的诱导。
3.超量表达GbMPK3提高了烟草对干旱和氧化胁迫的耐受性构建了GbMPK3基因超量表达载体并分别转化烟草和棉花。通过抗性筛选和分子鉴定分别获得了高量表达GbMPK3的转基因烟草和棉花株系。对野生型和转基因烟草在成株期进行干旱胁迫处理,发现转基因烟草具有更强的耐旱性。表现为转基因系复水之后的株高和存活率显著高于野生型。生理生化分析发现超表达GbMPK3提高了烟草在干旱胁迫下抗氧化酶APX的基因表达和酶活性,从而减少了烟草细胞内的活性氧积累。用MV处理烟草种子和成株期烟草叶片,发现MV对野生型和转基因系都造成了氧化胁迫,但是野生型对MV更敏感。MV处理之后野生型烟草种子的萌发率和成株期叶盘的叶绿素含量均显著低于转基因植株。RT-PCR分析结果显示,GbMPK3超表达的烟草在MV处理后抗氧化酶APX的基因表达量高于野生型。推测超量表达GbMPK3增强烟草对干旱和氧化胁迫的耐受性可能与提高抗氧化酶的表达和活性有关。
4.超量表达GbMPK3影响了烟草对烟草花叶病毒(TMV)和灰霉病菌的抗性对超量表达GbMPK3基因的烟草转基因系和野生型进行病原菌接种处理。转基因烟草对TMV的抗性增强,超表达GbMPK3降低了TMV在烟草中扩散和繁殖的速率；而接种灰霉病菌后转基因烟草则表现出更感病的表型。
5.超量表达GbMPK3降低了烟草和棉花对黄萎病菌的抗性对超量表达GbMPK3的烟草和棉花转基因材料进行黄萎病菌接种实验,发现转基因烟草和棉花的抗病性都显著降低。受黄萎病菌侵染后,野生型和转基因材料都表现出叶片萎蔫、黄化等黄萎病的典型病征,但野生型的叶片黄化与萎蔫出现的时期晚于转基因系,具体表现为野生型的发病率和病指显著低于转基因系。利用VIGS方法抑制棉花中GbMPK3的表达,对GbMPK3沉默的棉花进行接种实验。接种黄萎病菌后,对照材料和抑制GbMPK3表达的材料之间没有观察到抗病性的差异,两者在叶片黄化和脱落程度上无明显区别。说明抑制(GbMPK3表达对棉花抗黄萎病的影响不大,而超量表达GbMPK3显著减弱了植物对黄萎病菌的抗病性。
6. GbMPK3影响黄萎病菌侵染下棉花SA相关基因的表达qPCR分析显示,无论是GbMPK3抑制表达、GbMPK3超量表达还是野生型的棉花根系中SA相关基因,如WRKY70、 PR1和PR5等,在接种5天后相对于接水处理的对照组都显著上调。与野生型相比,GbMPK3超量表达转基因系中WRKY70,PR1和PR5等基因的表达水平显著升高；而抑制GbMPK3表达并未影响PRl和PR5的表达。这些结果说明棉花品种YZ1受黄萎病菌侵染后SA路径被激活,GbMPK3的超量表达增强了SA信号路径的激活水平。现有的一些研究结果表明,JA可能正调控植物对黄萎病菌的抗性,SA和JA信号路径通过WRKY70、 NPR1等关键因子相互拮抗。因此,SA也可能影响棉花对黄萎病菌的抗性,但SA调控棉花抗性的具体机理、超量表达GbMPK3降低陆地棉对黄萎病菌的抗性与其显著升高的SA信号传导水平是否关联还有待进一步的研究。
Mitogen-activated protein kinase (MAPK) cascades are highly conserved signaling modules found in all eukaryotes, and they play essential roles in cell signal transduction. A mitogen-activated protein kinase gene, GbMPK3, was induced upon Verticillium dahliae infection. In this work, GbMPK3was isolated from sea-island cotton Hai7124and its functional roles in response to different abiotic and biotic stress conditions were analyzed in both tobacco (Nicotiana benthamiana) and upland cotton YZ1. The main results of this work were as follows:
1. Cloning and characterization of GbMPK3An EST (CLlContig1773) putatively encoded a MAPK, GbMPK3, was isolated based on the results of RNA-seq of sea-island cotton Hai7124inoculation with V. dahliae. The full-length of cDNA and DNA sequences were obtained through RACE and Genome-walking. Sequence similarity search and phylogenetic analysis revealed that GbMPK3was classified as a group A MAPK. The35S-GFP::GbMPK3constructs were completed and introduced into tobacco leaves to investigate the subcellular localization of GbMPK3, and the result indicated that GbMPK3protein mainly accumulated in the nucleus.2. Expression profile of GbMPK3
The expression level of GbMPK3in cotton tissues under normal condition was evaluated by RT-PCR, and the results showed that GbMPK3expression level was low in all tissues and preferentially expressed in roots, petals, anthers and fibers. qPCR was employed to determine whether the expression of GbMPK3was affected by hormones and stresses, and GbMPK3was up-regulated upon cold (4℃), heat (42℃),10μmol/L MV,200mmol/L NaCl, dehydration, hormones (such as1mmol/L SA) and V. dahliae treatments in our study.
3. Overexpression of GbMPK3enhances drought and oxidative stress tolerance in tobacco
GbMPK3was introduced into tobacco and cotton to generate transgenic plants with constitutively higher expression of GbMPK3. Homozygous (T2) transgenic tobacco were conferred with enhanced drought tolerance, reduced water loss during drought treatment, improved plant height and survival rates after re-watering. Additionally, the gene expression levels and enzymatic activity of antioxidant enzymes were more strongly induced with depressed hydrogen peroxide accumulation in GbMPK3-overexpressing tobacco compared with wild-type under drought condition. Furthermore, observation of seed germination and leaf morphology showed that tolerance of transgenic plants to MV was improved due to increased antioxidant enzyme expression, suggesting that GbMPK3may positively regulate drought tolerance through enhanced reactive oxygen species scavenging ability.
4. Overexpression of GbMPK3affects tobacco disease resistance to Tobacco mosaic virus (TMV) and Botrytis cinerea
Wilt-type and transgenic tobacco were inoculated with different types of plant pathogens. Transgenic tobacco enhances resistance against TMV with reduced spreading rate and reproduction of TMV. Yet overexpression of GbMPK3significantly reduces tobacco resistance to the fungal pathogen Botrytis cinerea.
5. Overexpression of GbMPK3reduces V. dahliae resistance in tobacco and cotton
Wilt-type and transgenic plants were inoculated with V. dahliae to investigate the involvement of GbMPK3in plant defence. GbMPK3overexpression tobacco and cotton shows enhanced disease susceptiblity to V. dahliae strain V991. Both wilt-type and transgenic plants shows typical symptoms of verticillium wilt several days post V. dahliae infection, including wilting and yellowing leaves. Whereas the disease symptoms appeared early in GbMPK3-overexpression plants, resulting in elevated rate of diseased plants and disease index compared to wilt-type. The V. dahliae resistance was also analyzed in VIGS built GbMPK3-silencing plants. Silence of GbMPK3in cotton did not affect pathogen invasion, but slightly reduces yellowing and wilting leaves of diseased plants, as show by the decreased disease index. Therefore, while the effect of GbMPK3-silencing on cotton resistance to V. dahliae turns out to be weak, GbMPK3overexpression could significantly enhances V. dahliae susceptibility in both tobacco and cotton.
6. GbMPK3affects SA-related gene expression in cotton
qPCR analysis showed that the transcripts of SA-related genes (including the transcription factor WRKY70and two maker genes PR1and PR5) were accumulated in both wild-type, GbMPK3-silencing and overexpression cotton roots at day5post V. dahliae inoculation, suggested an up-regulation role of SA signaling in susceptible cotton cultivar YZ1. The elevated level of PR1and PR5was almost the same in wild-type and GbMPK3-silencing plants, yet transcripts of WRKY70, PR1and PR5were higher in GbMPK3overexpression cotton, indicating that GbMPK3overexpression may enhances V. dahliae induced SA signaling transduction. In previous studies, researches suggested JA signaling acted as a positively regulator in plants defense to V. dahliae. SA signaling is usually known antagonistic to JA signaling. Many important genes in regulating SA and JA signaling were reported, such as WRKY70and NPR1. Thus, SA may also participate in the complex signal networks of V. dahliae resistance in plants. But the mechanism of SA involved in cotton defense and the relation between elevated SA-related defence gene expression and reduced V. dahliae resistance still need to be studied further.
引文
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