植物黄酮类化合物槲皮素与转录因子AtMYB44诱导和调控植物防卫反应的研究
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摘要
转录因子在植物识别病原物后激活防卫反应的过程中起着重要的作用,它通过调控防卫基因的表达实现防卫反应。植物抗病性激发子能够诱导植物对多种病害的抗性。但是它们如何启动抗病信号通路的,如何参与抗病反应的,目前其机制不清楚。因此,本博士论文着重解析转录因子AtMYB44在植物抗病反应中的作用和槲皮素(quercetin)诱导的抗病机制,探讨了AtMYB44基因启动子甲基化对该基因的表达及其对抗病反应的影响。为阐明转录因子和黄酮类化合物参与防卫反应的机制及与激素或非激素信号传导途径的交叉提供研究基础。
1. AtMYB44转基因拟南芥的产生及鉴定
MYB类转录因子在植物胁迫应答过程中起到重要的调控作用,能够使植物提高抗病能力。AtMYB44属于拟南芥中R2R3 MYB亚基22转录因子家族。其T-DNA插入突变体atmyb44和野生型拟南芥相比能够增强PP2C编码基因的盐诱导表达,减少干旱或盐胁迫容忍性。atmyb44突变体使桃蚜驱避率降低。接种Pseudomonas syringae pv. tomato DC3000 (PstDC3000)后,和野生型相比,atmyb44突变体表现抗病性降低。为了分析转录因子AtMYB44在植物防卫反应中的作用,我们构建了双元载体pCAMBI1301::44::6His,成功地将AtMYB44基因引入到载体中,并将该质粒转入农杆菌A.tumefaciens EHA105中,转化拟南芥生态型Col-0,构建了过表达植株35S-M,抗生素抗性筛选和分子生物学验证后获得过表达植株纯合体。采用同样的方法将AtMYB44基因和GFP基因转入野生型拟南芥Col-0中,构建了过表达植株44GFP。同样我们将AtMYB44基因ATG上游的2000bp(44P2000)引入了pBI121载体,构建了双元载体pB121::44P2000::GUS,转基因验证筛选后,由农杆菌A.tumefaciens EHA105介导,转化拟南芥生态型Col-0,获得转基因植株纯合体44PGUS。随后我们又对T-DNA插入突变体atmyb44进行了卡那霉素筛选和分子鉴定,成功获得纯合体atmyb44。我们还构建了携带双元载体的工程菌A.tumefaciens EHA105(pCAMBI1301:44P2000::44CDS::GFP::His),转入拟南芥拟南芥突变体atmyb44,将44P2000::44CDS:GFP引入突变体中,经筛选鉴定后获得纯化的互补转基因拟南芥Catmyb44.过表达拟南芥35S-M和44GFP、启动子转基因拟南芥44PGUS、互补拟南芥Catmyb44和突变体atmyb44纯合体获得为后续实验准备了实验材料。
2.转录因子AtMYB44诱导表达及亚细胞定位
转录因子是在胁迫反应和生长发育中基因表达改变的重要调控因子。核定位信号区是使转录因子能够进入细胞核,从而调控下游基因的表达,最终决定生物表型。MYB家族就是一群能够结合特定DNA序列的转录因子.AtMYB44基因可以被非生物胁迫诱导增强表达,例如,创伤、冷胁迫、盐胁迫和一些激素诱导,增强植物的抗逆反应或植物发育生长。本实验构建了AtMYB44基因的原核表达工程菌E.coli BL21 (pET30a::AtMYB44),并利用AtMYB44蛋白PAGE电泳条带制备了AtMYB44蛋白的兔抗。我们利用A.tumefaciens EHA105 (pCAMBI1301::44P2000::44 CDS::GFP::His)和A.tumefaciens EHA105 (pBI121::44::GFP)在洋葱表皮上对AtMYB44进行了HrpNEa蛋白和乙烯利(ACC)诱导表达和亚细胞定位,同时利用转基因拟南芥Catmyb44和44GFP对AtMYB44进行了亚细胞定位分析,结果表明AtMYB44定位在细胞核内,也验证了44P2000的活性。而在HrpNEa蛋白和ACC诱导下,和对照相比,GFP能够更强的表达。对44PGUS组织化学染色实验分析表明,AtMYB44在拟南芥的各个组织中进行表达,AtMYB44基因的RT-PCR也验证了这一结果,而且发现HrpNEa和槲皮素能够在拟南芥的各个组织中诱导AtMYB44基因表达。我们对Catmyb44的10个品系和35S-M的5个品系进行了AtMYB44基因诱导表达,筛选到最佳转基因植物。这些结果暗示了AtMYB44基因能够被一些生物或非生物胁迫诱导表达,而在植物体内发挥其生物学功能。
3.转录因子AtMYB44在拟南芥中对Pseudomonas syringae抗病性的作用
植物受病原菌侵染或用病原物激发子处理植物时,植物激活的防卫反应中涉及大量基因的转录调控。AtMYB44属于拟南芥中R2R3-MYB亚基22转录因子家族。大量的证据暗示植物R2R3 MYB转录因子卷入到植物防卫中,当然防卫反应包括植物防御病染菌时宿主基因的转录调控。我们分析了AtMYB44转录因子在拟南芥对PstDC3000基本防卫中的作用。AtMYB44的T-DNA插入突变体atmyb44接种PstDC3000后,与野生型相比,3d后叶片细菌数量增加,病害症状显著加重,水杨酸调控的PR1和PR2基因表达减弱,AtMYB44能够在野生型拟南芥上被PstDC3000诱导表达。AtMYB44基因过表达植株35S-M-1和35S-M-4接种PstDC3000后,与野生型相比,3d后叶片细菌数量减少约1倍,病害症状显著减弱,水杨酸调控的PRl和PR2基因表达增强,AtMYB44被PstDC3000诱导的表达明显上调；而互补植株Catmyb44-2、Catmyb44-7和Catmyb44-9接种PstDC3000后,与野生型相比症状和抗性基因表达没有变化。从细胞水平上分析了野生型WT、转基因35S-M和Catmyb44拟南芥植株对病原菌产生的防卫反应,分子水平上分析了AtMYB44受环境胁迫(SA、MeJA、ET、ABA、HrpNEa、槲皮素)的诱导表达,这些结果表明AtMYB44正调控植物对PstDC3000的基本防卫反应。通过分析AtMYB44在基本防卫信号通路突变NahG与nprl上的受病原细菌诱导表达,表明水杨酸信号通路正调控AtMYB44表达。AtMYB44具有转录激活,能在植物防卫反应中调控下游基因表达,参与植物对病原细菌的抗性。
4.槲皮素诱导的过氧化氢在拟南芥中对Psedomonas syringae pv. tomato DC3000抗病性的作用
以前研究证明槲皮素可以作为一种重要的抗氧化剂应用在人类的疾病治疗中。最近一些报道表明槲皮素是一种在特定条件下能作为促氧化剂作用。在这个研究中,我们展示了槲皮素能够诱导活性氧的产生并能增强拟南芥野生型对PstDC3000的抗性。当野生型拟南芥用槲皮素处理后再接种PstDC3000叶片就会出现多种防卫反应包括活性氧爆发、胼胝质沉积、过敏性细胞坏死、水杨酸的积累以及防卫基因PRl和PALl的表达变化等。这些防卫反应和植物对PstDC3000的抗性相一致的,因为再利用槲皮素处理植物中的病菌数量显著的少于对照植物。有趣的是当外施槲皮素后接种PstDC3000并同时用过氧化氢酶处理拟南芥上述的防卫反应和相关症状就消失了。而只用槲皮素处理或单独接种PstDC3000则只能在细胞或者分子水平上诱发有限的防卫反应。为了进一步的研究槲皮素在植物防卫反应中的作用,我们对不同的拟南芥突变体用槲皮素和PstDC3000单独处理或者先用槲皮素处理后再接种PstDC3000.发现在jar1, ein2和abil-2突变体中与野生型中结果是相似的,而在NahG和nprl中相应的防卫反应却消失了。上述结果表明槲皮素通过提高活性氧来增强拟南芥对PstDC3000的抗性,并且需要水杨酸信号通路和NPR1基因参与的。5.AtMYB44基因启动子DNA甲基化在拟南芥抗病防卫反应中的影响
DNA胞嘧啶甲基化作用对许多后生遗传过程都很重要,包括X染色体的失活、基因组印记、在致癌过程中的后生变化、转座子沉默、发育过程中特殊基因沉默和某种转基因的沉默。植物可以通过表观遗传来调控基因表达来适应环境压力,可以说生物体的遗传性状并不是完全由DNA决定的。在生物和非生物环境胁迫压力下,转录因子往往对生物体的遗传性状起到至关重要作用,而转录因子的表达也是受到DNA遗传修饰的调控。本实验分析了不同激素(ET、ABA、MeJA和SA)、HrpNEa、槲皮素和病原菌(PstDC3000和Erwinia carotovora subsp.carotovora (Ecc))诱导拟南芥表达转录因子AtMYB44,并检测了其启动子(44P2000)CG岛甲基化的变化,结果发现不同的处理CG岛甲基化的程度不同。5-氮杂-2’-脱氧胞苷(5-aza-2'deoxycytidine, 5-aza)能够去除DNA上的甲基化。野生型拟南芥Col-0、突变体atmyb44和互补植株Catmyb44-2被5-aza处理后,通过RT-PCR检测了AtMYB44基因的表达,初步确定了5-aza能够促进植物对AtMYB44基因的表达。而通过对被5-aza处理的转基因拟南芥P44GUS定性的或定量的GUS蛋白检测,发现5-aza影响着AtMYB44基因启动子发挥作用。PstDC3000接种于被5-aza处理的植物后,拟南芥发病表型、PRl和PAL基因的表达暗示了5-aza与抗病反应没有关系,仅仅是对转录因子AtMYB44表达发挥着作用。这些结果暗示了外界环境的胁迫会影响植物中AtMYB44基因上游启动子CG岛甲基化的数量变化,从而使AtMYB44基因发生不同程度的表达,继而激发了植物的抗病基因的表达,最终使植物具有抗病性。
总结
本研究利用转AtMYB44基因植物及外源槲皮素两种手段,对它们引发植物防卫反应信号传导作用及机制进行了解析。研究发现：(1)转录因子AtMYB44能够参与植物抗病防卫反应；转录因子AtMYB44能够被多种生物胁迫和非生物胁迫诱导表达,对病原细菌(PstDC3000)的抗性正向调控,并介导SA信号传导通路发挥作用。(2)槲皮素能够诱导AtMYB44在拟南芥中表达,并介导过氧化氢水平的升高增强拟南芥对病原细菌(PstDC3000)抗性。其抗病性机制是依赖SA信号和NPR1。(3)在生物和非生物胁迫中,转录因子AtMYB44的表达是靠其基因上游启动子甲基化来调控的。
创新点
(1)利用转基因拟南芥从细胞水平和分子水平研究了转录因子AtMYB44在植物防卫反应中的作用及其诱导表达的机理,最终确定了AtMYB44在植物防卫反应中功能及机制。
(2)分析了黄酮类化合物槲皮素在植物对PstDC3000的抗性中的作用,确定了槲皮素在植物防卫反应中的作用及机制。
Activation of defense responses against Pathogen infeetion is usually regulated through modulation of a large set of defense genes at transcriptional level by specific transcription factors. Elicitors of systemic acquired resistance (SAR) in plants can trigger plant disease resistance in Arabidopsis. But the underlying molecular mechanism, how they switch on signal pathway and regulate plant defense responses to pathogen attack, remains unclear. Studies in this Ph.D project aim at determination of functions of transcription factor AtMYB44 in plants defense responses against pathogenic bacteria and mechanism of quercetin inducing defense responses against pathogen. We investigated the effect of methylation of AtMYB44 promotor on expression of AtMYB44 and responses against pathogen attack. The results here will provide basic clues to explore the mechanism of the transcription factor and flovoniods mediated plant defense and growth pathways and crosstalk with hormonal and non-hormonal signaling pathways.
1. The production and identification of the atmyb44 transgenic Arabidopsis
Transcription factor MYB plays the important role of regulation in response of plants stress. It can enhance disease resistance of plants. AtMYB44 belongs to the R2R3 MYB subgroup 22 transcription factor family in Arabidopsis (Arabidopsis thaliana). The atmyb44 knockout mutant line exhibited enhanced salt-induced expression of PP2C-encoding genes and reduced drought/salt stress tolerance compared to wild-type plants. Green peach aphid repellency and resistance to Pseudomonas syringae pv. tomato DC3000 (PstDC3000) are decreased in atmyb44, compared to these in wild-type plants. To analyze the role of transcription factor AtMYB44 in plants defense response, we constructed unit binary vector pCAMBI1301::44::6His with AtMYB44 gene and the vector was transformed into Agrobacterium tumefaciens strain EHA105.Then the recombinant unit was introduced into Arabidopsis Col-0. Transgenic homozygous Arabidopsis (35S-M), which can overexpress AtMYB44 under the regulation of 35S promotor, were obtained by screening with antibiotic resistance and identifying with molecular biology. Unit doublet vector pBI121::44P2000::GUS with 44P2000 gene was build.44P2000 gene is upriver 2000bp of AtMYB44 gene ATG. Then the recombinant unit was introduced into Arabidopsis Col-0. Transgenic homozygous Arabidopsis (44PGUS), which can express GUS protein under the regulation of 44P2000, were obtained by screening with antibiotic resistance and identifying with molecular biology. Whereafter, homozygous mutant atmyb44, produced by inseting T-DNA into promotor of AtMYB44, were obtained by screening with antibiotic resistance and identifying with molecular biology. A.tumefaciens EHA105 (pCAMBI1301: 44P2000::44CDS::GFP::His) was also build and the recombinant unit was introduced into Arabidopsis mutant atmyb44. Complementary Arabidopsis for AtMYB44(Catmyb44) gene were constructed. Homozygous Arabidopsis Catmyb44 was obtained by screening with antibiotic resistance and identifying with molecular biology.35S-M and 44GFP,44PGUS, Catmyb44, and atmyb44 will be applied in follow-up experiments as subject.
2. Induced expression and subcellular localization of transcription factor AtMYB44
Transcription factors are critical regulators of the changes in gene expression that drive developmental processes and environmental stress responses. Nuclear localization signal can enter transcription factor into nucleus and transcription factor can regulate expression of downstream genes which can determine organism phenotype. MYB is a DNA-binding transcription factor family. AtMYB44 belongs to the R2R3 MYB subgroup 22 transcription factor family in Arabidopsis(Arabidopsis thaliana). Expression of AtMYB44 gene can be enhanced by induction of abiotic stress including wound, cold stress, salt stress, and auxin, which can promote resistance response and development of plants. In this study, E.coli BL21 (pET30a::AtMYB44), which can express AtMYB44 gene, was constructed and antibody of AtMYB44 was prepared in rabbit by using strip of AtMYB44 PAGE electrophoresis. Subcellular localization of AtMYB44 was analyzed in onion epidermis by A.tumefaciens EHA105 (pCAMBI1301::44P2000::44CDS::GFP::His) and A.tumefaciens EHA105 (pBI121::44::GFP) and in Catmyb44 and 44GFP. The result shows that AtMYB44 is localized in nucleus and 44P2000 can activate downstream gene. GFP protein could generate more accumulation in Catmyb44 under the condition of induction of HrpNEa protein and ACC compared with EVP. The result of histochemical stain of 44PGUS showed that expression of AtMYB44 existed in each tissue of Arabidopsis, which was identified by RT-PCR of AtMYB44. Moreover, HrpNEa and quercetin could induce expression of Atmyb44 in each tissue of Arabidopsis. Expression of AtMYB44 gene was induced in 10 Catmyb44 lines and 535S-M lines by HrpNEa protein and quercetin, and optimal transgenic plants were screened. The results suggest that Expression of AtMYB44 gene can be induced by some biotic and abiotic stress. And AtMYB44 can function in plants.
3. The roles of transcription factor AtMYB44 in defense resistance of Arabidopsis against Pseudomonas syringae
A common feature of plant defense responses is the transcriptional regulation of a large number of genes upon pathogen infection or treatment with pathogen elicitors. AtMYB44 belongs to the R2R3-MYB subgroup 22 transcription factor family in Arabidopsis (Arabidopsis thaliana). A large body of evidence suggests that plant R2R3 MYB transcription factors are involved in plant defense including transcriptional regulation of plant host genes in response to pathogen infection. However, there is only limited information about the roles of R2R3 MYB transcription factors in plant defense. We analyzed the role of the AtMYB44 transcription factor from Arabidopsis in plant defense against the bacterial pathogen Pseudomonas syringae (PstDC3000). T-DNA insertion mutant atmyb44 for AtMYB44 increased growth of PstDC3000 and displayed increased disease symptom severity as compared to wild-type plants. The atmyb44 mutant plants also displayed reduced expression of the SA-regulated PR1 and PR2 genes after the pathogen infection. PstDC3000 could induce expression of AtMYB44 in Arabidopsis. Overexpression Arabidopsis 35S-M-1, and-4 for AtMYB44 reduced growth of PstDC3000 and displayed decreased disease symptom severity as compared to wild-type plants. The 35S-M (1,4) plants also displayed increased expression of the SA-regulated PR1 and PR2 genes after the pathogen infection. Expression of AtMYB44 has obvious up-regulation by induction of PstDC3000. However, after complementary Arabidopsis Catmyb44-2,-7, and-9 for AtMYB44 were inoculated by PstDC3000, Catmyb44-2,-7, and-9 had the same disease symptom severity and expression of PR1 and PR1 as compared to wild-type plants. Analysis of cellular defense response against PstDC3000 in WT,35S-M, and Catmyb44 and expression of AtMYB44 under the condition of environmental stress (SA, MeJA, ET, ABA, HrpNEa, and quercetin) showed that stress-induced AtMYB44 functioned as a positive regulator of defense responses to PstDC3000. Analysis of PstDC3000-induced AtMYB44 in the defense signaling mutants NahG and nprl-1 further indicated that this gene is positively regulated by the salicylic acid (SA) signaling pathway. AtMYB44 was a transcriptional activator and was able to activate the expression of genes involved in plant defense.
4. Quercetin-induced H2O2 mediates the pathogen resistance against Psedomonas syringae pv. tomato DC3000 in Arabidopsis thaliana
Quercetin has been previously demonstrated to function as important antioxidant in human disease therapy. However, some reports show it is also prooxidant and can provoke hydrogen peroxide (H2O2) under certain conditions. In the current study, we demonstrated that quercetin induced the production of H2O2 and enhanced Arabidopsis thaliana (Arabidopsis) resistance against the virulent strain Pseudomonas syringae pv. tomato DC3000 (PstDC3000). When Arabidopsis was pretreated with quercetin before PstDC3000 inoculation, the leaves exhibited several defense responses, including H2O2 burst, callose deposition, hypersensitive cell death, and the expression of defense response genes PR1 (pathogenesis-related1) and PAL1(Phe ammonia-lyasel) expression. These responses were consistent with the enhancement of plant resistance to PstDC3000, which had significantly decreased population in the quercetin-pretreated plants than control plants. When we applied catalase and PstDC3000 together on the Arabidopsis pretreated with quercetin, the defense responses and other relevant symptoms vanished. Application of quercetin or inoculation of PstDC3000 on leaves of Arabidopsis could only induce limited defense responses in cellular or molecular level. In the further study, we treated the Arabidopsis mutants with quercetin or/and PstDC3000. These similar defense responses also occurred in the Arabidopsis mutant jar1, ein2 and abi1-2, while disappeared in the Arabidopsis mutant NahG and nprl. These studies illustrate that quercetin enhance Arabidopsis resistance against PstDC3000 infection by quercetin-mediated H2O2 elevation and depend SA and NPR1.
5. The effect of DNA methylation of atmyb44 promotor on resistance defense response of Arabidopsis
DNA cytosine methylation is important for many epigenetic processes including X chromosome inactivation, genomic imprinting, epigenetic changes during carcinogenesis, and silencing of transposons, of specific genes during development and of certain transgenes. To adapt plants to environment stress, gene expression in plants is controlled by epigenetics. Inheritable character of organism is not fully determined by DNA. Under the condition of bioctic and obiotic environment stress, transcription factor plays an important role on Inheritable character of organism, and expression of transcription factor is controlled by DNA genetic modification. In this study, expression of transcription factor AtMYB44 is analyzed in Arabidopsis treated by ET, ABA, MeJA, SA, HrpNEa, quercetin, PstDC3000, and E. carotovora subsp.carotovora (Ecc). And methylation of CG island in 44P2000.The result shows that different treatment leads to the discrimination of level of CG-island methylation.5-aza-2'deoxycytidine (5-aza) can remove methylation in DNA. Wild type Arabidopsis Col-0, mutant atmyb44, and complementary transgenic Arabidopsis Catmyb44 were treated by 5-aza. Then, expression of AtMYB44 was detected by RT-PCR. It was initially determined that 5-aza cold promoter expression of AtMYB44 gene in Arabidopsis. GUS protein from transgenic Arabidopsis P44GUS treated by 5-aza was detected qualitatively or quantitatively. It was found that 5-aza had an effect on the role of AtMYB44 promotor. After plants treated by 5-aza were inoculated by PstDC3000, Disease symptom development and expression of PR1 and PAL suggest that 5-aza has relation with resistance response against disease and plays a role in expression of AtMYB44. These results suggest that outside environmental stress can have an effect on the number of CG-island methylation in the upper stream promoter of AtMYB44 and the expression of AtMYB44 in different level provoked by methylation of CG island promotes expression of resistance and defense genes. In fine, plants possess resistance against pathogen.
Conclusive remarks
Data obtained from studies described above have provided us the further understanding on mechanisms and functions of pathogen defense in plants by transgenic plant and exogenous quercetin. Firstly, transcription factor AtMYB44 can be induced by biotic and abiotic stress in plants defense responses against pathogen attack. It is found that AtMYB44 functions as positive regulation against PstDC3000 by SA mediated signaling pathway. Secondly, Quercetin can induce expression of AtMYB44 in Arabidopsis and enhance resistance against PstDC3000 by the level of H2O2. The mechanism of defense response against pathogen depends on SA and NPR1. Thirdly, expression of AtMYB44 is regulated by up-stream promotor methylation of AtMYB44 gene in biotic and abiotic stress.
1. AtMYB44转基因拟南芥的产生及鉴定
MYB类转录因子在植物胁迫应答过程中起到重要的调控作用,能够使植物提高抗病能力。AtMYB44属于拟南芥中R2R3 MYB亚基22转录因子家族。其T-DNA插入突变体atmyb44和野生型拟南芥相比能够增强PP2C编码基因的盐诱导表达,减少干旱或盐胁迫容忍性。atmyb44突变体使桃蚜驱避率降低。接种Pseudomonas syringae pv. tomato DC3000 (PstDC3000)后,和野生型相比,atmyb44突变体表现抗病性降低。为了分析转录因子AtMYB44在植物防卫反应中的作用,我们构建了双元载体pCAMBI1301::44::6His,成功地将AtMYB44基因引入到载体中,并将该质粒转入农杆菌A.tumefaciens EHA105中,转化拟南芥生态型Col-0,构建了过表达植株35S-M,抗生素抗性筛选和分子生物学验证后获得过表达植株纯合体。采用同样的方法将AtMYB44基因和GFP基因转入野生型拟南芥Col-0中,构建了过表达植株44GFP。同样我们将AtMYB44基因ATG上游的2000bp(44P2000)引入了pBI121载体,构建了双元载体pB121::44P2000::GUS,转基因验证筛选后,由农杆菌A.tumefaciens EHA105介导,转化拟南芥生态型Col-0,获得转基因植株纯合体44PGUS。随后我们又对T-DNA插入突变体atmyb44进行了卡那霉素筛选和分子鉴定,成功获得纯合体atmyb44。我们还构建了携带双元载体的工程菌A.tumefaciens EHA105(pCAMBI1301:44P2000::44CDS::GFP::His),转入拟南芥拟南芥突变体atmyb44,将44P2000::44CDS:GFP引入突变体中,经筛选鉴定后获得纯化的互补转基因拟南芥Catmyb44.过表达拟南芥35S-M和44GFP、启动子转基因拟南芥44PGUS、互补拟南芥Catmyb44和突变体atmyb44纯合体获得为后续实验准备了实验材料。
2.转录因子AtMYB44诱导表达及亚细胞定位
转录因子是在胁迫反应和生长发育中基因表达改变的重要调控因子。核定位信号区是使转录因子能够进入细胞核,从而调控下游基因的表达,最终决定生物表型。MYB家族就是一群能够结合特定DNA序列的转录因子.AtMYB44基因可以被非生物胁迫诱导增强表达,例如,创伤、冷胁迫、盐胁迫和一些激素诱导,增强植物的抗逆反应或植物发育生长。本实验构建了AtMYB44基因的原核表达工程菌E.coli BL21 (pET30a::AtMYB44),并利用AtMYB44蛋白PAGE电泳条带制备了AtMYB44蛋白的兔抗。我们利用A.tumefaciens EHA105 (pCAMBI1301::44P2000::44 CDS::GFP::His)和A.tumefaciens EHA105 (pBI121::44::GFP)在洋葱表皮上对AtMYB44进行了HrpNEa蛋白和乙烯利(ACC)诱导表达和亚细胞定位,同时利用转基因拟南芥Catmyb44和44GFP对AtMYB44进行了亚细胞定位分析,结果表明AtMYB44定位在细胞核内,也验证了44P2000的活性。而在HrpNEa蛋白和ACC诱导下,和对照相比,GFP能够更强的表达。对44PGUS组织化学染色实验分析表明,AtMYB44在拟南芥的各个组织中进行表达,AtMYB44基因的RT-PCR也验证了这一结果,而且发现HrpNEa和槲皮素能够在拟南芥的各个组织中诱导AtMYB44基因表达。我们对Catmyb44的10个品系和35S-M的5个品系进行了AtMYB44基因诱导表达,筛选到最佳转基因植物。这些结果暗示了AtMYB44基因能够被一些生物或非生物胁迫诱导表达,而在植物体内发挥其生物学功能。
3.转录因子AtMYB44在拟南芥中对Pseudomonas syringae抗病性的作用
植物受病原菌侵染或用病原物激发子处理植物时,植物激活的防卫反应中涉及大量基因的转录调控。AtMYB44属于拟南芥中R2R3-MYB亚基22转录因子家族。大量的证据暗示植物R2R3 MYB转录因子卷入到植物防卫中,当然防卫反应包括植物防御病染菌时宿主基因的转录调控。我们分析了AtMYB44转录因子在拟南芥对PstDC3000基本防卫中的作用。AtMYB44的T-DNA插入突变体atmyb44接种PstDC3000后,与野生型相比,3d后叶片细菌数量增加,病害症状显著加重,水杨酸调控的PR1和PR2基因表达减弱,AtMYB44能够在野生型拟南芥上被PstDC3000诱导表达。AtMYB44基因过表达植株35S-M-1和35S-M-4接种PstDC3000后,与野生型相比,3d后叶片细菌数量减少约1倍,病害症状显著减弱,水杨酸调控的PRl和PR2基因表达增强,AtMYB44被PstDC3000诱导的表达明显上调；而互补植株Catmyb44-2、Catmyb44-7和Catmyb44-9接种PstDC3000后,与野生型相比症状和抗性基因表达没有变化。从细胞水平上分析了野生型WT、转基因35S-M和Catmyb44拟南芥植株对病原菌产生的防卫反应,分子水平上分析了AtMYB44受环境胁迫(SA、MeJA、ET、ABA、HrpNEa、槲皮素)的诱导表达,这些结果表明AtMYB44正调控植物对PstDC3000的基本防卫反应。通过分析AtMYB44在基本防卫信号通路突变NahG与nprl上的受病原细菌诱导表达,表明水杨酸信号通路正调控AtMYB44表达。AtMYB44具有转录激活,能在植物防卫反应中调控下游基因表达,参与植物对病原细菌的抗性。
4.槲皮素诱导的过氧化氢在拟南芥中对Psedomonas syringae pv. tomato DC3000抗病性的作用
以前研究证明槲皮素可以作为一种重要的抗氧化剂应用在人类的疾病治疗中。最近一些报道表明槲皮素是一种在特定条件下能作为促氧化剂作用。在这个研究中,我们展示了槲皮素能够诱导活性氧的产生并能增强拟南芥野生型对PstDC3000的抗性。当野生型拟南芥用槲皮素处理后再接种PstDC3000叶片就会出现多种防卫反应包括活性氧爆发、胼胝质沉积、过敏性细胞坏死、水杨酸的积累以及防卫基因PRl和PALl的表达变化等。这些防卫反应和植物对PstDC3000的抗性相一致的,因为再利用槲皮素处理植物中的病菌数量显著的少于对照植物。有趣的是当外施槲皮素后接种PstDC3000并同时用过氧化氢酶处理拟南芥上述的防卫反应和相关症状就消失了。而只用槲皮素处理或单独接种PstDC3000则只能在细胞或者分子水平上诱发有限的防卫反应。为了进一步的研究槲皮素在植物防卫反应中的作用,我们对不同的拟南芥突变体用槲皮素和PstDC3000单独处理或者先用槲皮素处理后再接种PstDC3000.发现在jar1, ein2和abil-2突变体中与野生型中结果是相似的,而在NahG和nprl中相应的防卫反应却消失了。上述结果表明槲皮素通过提高活性氧来增强拟南芥对PstDC3000的抗性,并且需要水杨酸信号通路和NPR1基因参与的。5.AtMYB44基因启动子DNA甲基化在拟南芥抗病防卫反应中的影响
DNA胞嘧啶甲基化作用对许多后生遗传过程都很重要,包括X染色体的失活、基因组印记、在致癌过程中的后生变化、转座子沉默、发育过程中特殊基因沉默和某种转基因的沉默。植物可以通过表观遗传来调控基因表达来适应环境压力,可以说生物体的遗传性状并不是完全由DNA决定的。在生物和非生物环境胁迫压力下,转录因子往往对生物体的遗传性状起到至关重要作用,而转录因子的表达也是受到DNA遗传修饰的调控。本实验分析了不同激素(ET、ABA、MeJA和SA)、HrpNEa、槲皮素和病原菌(PstDC3000和Erwinia carotovora subsp.carotovora (Ecc))诱导拟南芥表达转录因子AtMYB44,并检测了其启动子(44P2000)CG岛甲基化的变化,结果发现不同的处理CG岛甲基化的程度不同。5-氮杂-2’-脱氧胞苷(5-aza-2'deoxycytidine, 5-aza)能够去除DNA上的甲基化。野生型拟南芥Col-0、突变体atmyb44和互补植株Catmyb44-2被5-aza处理后,通过RT-PCR检测了AtMYB44基因的表达,初步确定了5-aza能够促进植物对AtMYB44基因的表达。而通过对被5-aza处理的转基因拟南芥P44GUS定性的或定量的GUS蛋白检测,发现5-aza影响着AtMYB44基因启动子发挥作用。PstDC3000接种于被5-aza处理的植物后,拟南芥发病表型、PRl和PAL基因的表达暗示了5-aza与抗病反应没有关系,仅仅是对转录因子AtMYB44表达发挥着作用。这些结果暗示了外界环境的胁迫会影响植物中AtMYB44基因上游启动子CG岛甲基化的数量变化,从而使AtMYB44基因发生不同程度的表达,继而激发了植物的抗病基因的表达,最终使植物具有抗病性。
总结
本研究利用转AtMYB44基因植物及外源槲皮素两种手段,对它们引发植物防卫反应信号传导作用及机制进行了解析。研究发现：(1)转录因子AtMYB44能够参与植物抗病防卫反应；转录因子AtMYB44能够被多种生物胁迫和非生物胁迫诱导表达,对病原细菌(PstDC3000)的抗性正向调控,并介导SA信号传导通路发挥作用。(2)槲皮素能够诱导AtMYB44在拟南芥中表达,并介导过氧化氢水平的升高增强拟南芥对病原细菌(PstDC3000)抗性。其抗病性机制是依赖SA信号和NPR1。(3)在生物和非生物胁迫中,转录因子AtMYB44的表达是靠其基因上游启动子甲基化来调控的。
创新点
(1)利用转基因拟南芥从细胞水平和分子水平研究了转录因子AtMYB44在植物防卫反应中的作用及其诱导表达的机理,最终确定了AtMYB44在植物防卫反应中功能及机制。
(2)分析了黄酮类化合物槲皮素在植物对PstDC3000的抗性中的作用,确定了槲皮素在植物防卫反应中的作用及机制。
Activation of defense responses against Pathogen infeetion is usually regulated through modulation of a large set of defense genes at transcriptional level by specific transcription factors. Elicitors of systemic acquired resistance (SAR) in plants can trigger plant disease resistance in Arabidopsis. But the underlying molecular mechanism, how they switch on signal pathway and regulate plant defense responses to pathogen attack, remains unclear. Studies in this Ph.D project aim at determination of functions of transcription factor AtMYB44 in plants defense responses against pathogenic bacteria and mechanism of quercetin inducing defense responses against pathogen. We investigated the effect of methylation of AtMYB44 promotor on expression of AtMYB44 and responses against pathogen attack. The results here will provide basic clues to explore the mechanism of the transcription factor and flovoniods mediated plant defense and growth pathways and crosstalk with hormonal and non-hormonal signaling pathways.
1. The production and identification of the atmyb44 transgenic Arabidopsis
Transcription factor MYB plays the important role of regulation in response of plants stress. It can enhance disease resistance of plants. AtMYB44 belongs to the R2R3 MYB subgroup 22 transcription factor family in Arabidopsis (Arabidopsis thaliana). The atmyb44 knockout mutant line exhibited enhanced salt-induced expression of PP2C-encoding genes and reduced drought/salt stress tolerance compared to wild-type plants. Green peach aphid repellency and resistance to Pseudomonas syringae pv. tomato DC3000 (PstDC3000) are decreased in atmyb44, compared to these in wild-type plants. To analyze the role of transcription factor AtMYB44 in plants defense response, we constructed unit binary vector pCAMBI1301::44::6His with AtMYB44 gene and the vector was transformed into Agrobacterium tumefaciens strain EHA105.Then the recombinant unit was introduced into Arabidopsis Col-0. Transgenic homozygous Arabidopsis (35S-M), which can overexpress AtMYB44 under the regulation of 35S promotor, were obtained by screening with antibiotic resistance and identifying with molecular biology. Unit doublet vector pBI121::44P2000::GUS with 44P2000 gene was build.44P2000 gene is upriver 2000bp of AtMYB44 gene ATG. Then the recombinant unit was introduced into Arabidopsis Col-0. Transgenic homozygous Arabidopsis (44PGUS), which can express GUS protein under the regulation of 44P2000, were obtained by screening with antibiotic resistance and identifying with molecular biology. Whereafter, homozygous mutant atmyb44, produced by inseting T-DNA into promotor of AtMYB44, were obtained by screening with antibiotic resistance and identifying with molecular biology. A.tumefaciens EHA105 (pCAMBI1301: 44P2000::44CDS::GFP::His) was also build and the recombinant unit was introduced into Arabidopsis mutant atmyb44. Complementary Arabidopsis for AtMYB44(Catmyb44) gene were constructed. Homozygous Arabidopsis Catmyb44 was obtained by screening with antibiotic resistance and identifying with molecular biology.35S-M and 44GFP,44PGUS, Catmyb44, and atmyb44 will be applied in follow-up experiments as subject.
2. Induced expression and subcellular localization of transcription factor AtMYB44
Transcription factors are critical regulators of the changes in gene expression that drive developmental processes and environmental stress responses. Nuclear localization signal can enter transcription factor into nucleus and transcription factor can regulate expression of downstream genes which can determine organism phenotype. MYB is a DNA-binding transcription factor family. AtMYB44 belongs to the R2R3 MYB subgroup 22 transcription factor family in Arabidopsis(Arabidopsis thaliana). Expression of AtMYB44 gene can be enhanced by induction of abiotic stress including wound, cold stress, salt stress, and auxin, which can promote resistance response and development of plants. In this study, E.coli BL21 (pET30a::AtMYB44), which can express AtMYB44 gene, was constructed and antibody of AtMYB44 was prepared in rabbit by using strip of AtMYB44 PAGE electrophoresis. Subcellular localization of AtMYB44 was analyzed in onion epidermis by A.tumefaciens EHA105 (pCAMBI1301::44P2000::44CDS::GFP::His) and A.tumefaciens EHA105 (pBI121::44::GFP) and in Catmyb44 and 44GFP. The result shows that AtMYB44 is localized in nucleus and 44P2000 can activate downstream gene. GFP protein could generate more accumulation in Catmyb44 under the condition of induction of HrpNEa protein and ACC compared with EVP. The result of histochemical stain of 44PGUS showed that expression of AtMYB44 existed in each tissue of Arabidopsis, which was identified by RT-PCR of AtMYB44. Moreover, HrpNEa and quercetin could induce expression of Atmyb44 in each tissue of Arabidopsis. Expression of AtMYB44 gene was induced in 10 Catmyb44 lines and 535S-M lines by HrpNEa protein and quercetin, and optimal transgenic plants were screened. The results suggest that Expression of AtMYB44 gene can be induced by some biotic and abiotic stress. And AtMYB44 can function in plants.
3. The roles of transcription factor AtMYB44 in defense resistance of Arabidopsis against Pseudomonas syringae
A common feature of plant defense responses is the transcriptional regulation of a large number of genes upon pathogen infection or treatment with pathogen elicitors. AtMYB44 belongs to the R2R3-MYB subgroup 22 transcription factor family in Arabidopsis (Arabidopsis thaliana). A large body of evidence suggests that plant R2R3 MYB transcription factors are involved in plant defense including transcriptional regulation of plant host genes in response to pathogen infection. However, there is only limited information about the roles of R2R3 MYB transcription factors in plant defense. We analyzed the role of the AtMYB44 transcription factor from Arabidopsis in plant defense against the bacterial pathogen Pseudomonas syringae (PstDC3000). T-DNA insertion mutant atmyb44 for AtMYB44 increased growth of PstDC3000 and displayed increased disease symptom severity as compared to wild-type plants. The atmyb44 mutant plants also displayed reduced expression of the SA-regulated PR1 and PR2 genes after the pathogen infection. PstDC3000 could induce expression of AtMYB44 in Arabidopsis. Overexpression Arabidopsis 35S-M-1, and-4 for AtMYB44 reduced growth of PstDC3000 and displayed decreased disease symptom severity as compared to wild-type plants. The 35S-M (1,4) plants also displayed increased expression of the SA-regulated PR1 and PR2 genes after the pathogen infection. Expression of AtMYB44 has obvious up-regulation by induction of PstDC3000. However, after complementary Arabidopsis Catmyb44-2,-7, and-9 for AtMYB44 were inoculated by PstDC3000, Catmyb44-2,-7, and-9 had the same disease symptom severity and expression of PR1 and PR1 as compared to wild-type plants. Analysis of cellular defense response against PstDC3000 in WT,35S-M, and Catmyb44 and expression of AtMYB44 under the condition of environmental stress (SA, MeJA, ET, ABA, HrpNEa, and quercetin) showed that stress-induced AtMYB44 functioned as a positive regulator of defense responses to PstDC3000. Analysis of PstDC3000-induced AtMYB44 in the defense signaling mutants NahG and nprl-1 further indicated that this gene is positively regulated by the salicylic acid (SA) signaling pathway. AtMYB44 was a transcriptional activator and was able to activate the expression of genes involved in plant defense.
4. Quercetin-induced H2O2 mediates the pathogen resistance against Psedomonas syringae pv. tomato DC3000 in Arabidopsis thaliana
Quercetin has been previously demonstrated to function as important antioxidant in human disease therapy. However, some reports show it is also prooxidant and can provoke hydrogen peroxide (H2O2) under certain conditions. In the current study, we demonstrated that quercetin induced the production of H2O2 and enhanced Arabidopsis thaliana (Arabidopsis) resistance against the virulent strain Pseudomonas syringae pv. tomato DC3000 (PstDC3000). When Arabidopsis was pretreated with quercetin before PstDC3000 inoculation, the leaves exhibited several defense responses, including H2O2 burst, callose deposition, hypersensitive cell death, and the expression of defense response genes PR1 (pathogenesis-related1) and PAL1(Phe ammonia-lyasel) expression. These responses were consistent with the enhancement of plant resistance to PstDC3000, which had significantly decreased population in the quercetin-pretreated plants than control plants. When we applied catalase and PstDC3000 together on the Arabidopsis pretreated with quercetin, the defense responses and other relevant symptoms vanished. Application of quercetin or inoculation of PstDC3000 on leaves of Arabidopsis could only induce limited defense responses in cellular or molecular level. In the further study, we treated the Arabidopsis mutants with quercetin or/and PstDC3000. These similar defense responses also occurred in the Arabidopsis mutant jar1, ein2 and abi1-2, while disappeared in the Arabidopsis mutant NahG and nprl. These studies illustrate that quercetin enhance Arabidopsis resistance against PstDC3000 infection by quercetin-mediated H2O2 elevation and depend SA and NPR1.
5. The effect of DNA methylation of atmyb44 promotor on resistance defense response of Arabidopsis
DNA cytosine methylation is important for many epigenetic processes including X chromosome inactivation, genomic imprinting, epigenetic changes during carcinogenesis, and silencing of transposons, of specific genes during development and of certain transgenes. To adapt plants to environment stress, gene expression in plants is controlled by epigenetics. Inheritable character of organism is not fully determined by DNA. Under the condition of bioctic and obiotic environment stress, transcription factor plays an important role on Inheritable character of organism, and expression of transcription factor is controlled by DNA genetic modification. In this study, expression of transcription factor AtMYB44 is analyzed in Arabidopsis treated by ET, ABA, MeJA, SA, HrpNEa, quercetin, PstDC3000, and E. carotovora subsp.carotovora (Ecc). And methylation of CG island in 44P2000.The result shows that different treatment leads to the discrimination of level of CG-island methylation.5-aza-2'deoxycytidine (5-aza) can remove methylation in DNA. Wild type Arabidopsis Col-0, mutant atmyb44, and complementary transgenic Arabidopsis Catmyb44 were treated by 5-aza. Then, expression of AtMYB44 was detected by RT-PCR. It was initially determined that 5-aza cold promoter expression of AtMYB44 gene in Arabidopsis. GUS protein from transgenic Arabidopsis P44GUS treated by 5-aza was detected qualitatively or quantitatively. It was found that 5-aza had an effect on the role of AtMYB44 promotor. After plants treated by 5-aza were inoculated by PstDC3000, Disease symptom development and expression of PR1 and PAL suggest that 5-aza has relation with resistance response against disease and plays a role in expression of AtMYB44. These results suggest that outside environmental stress can have an effect on the number of CG-island methylation in the upper stream promoter of AtMYB44 and the expression of AtMYB44 in different level provoked by methylation of CG island promotes expression of resistance and defense genes. In fine, plants possess resistance against pathogen.
Conclusive remarks
Data obtained from studies described above have provided us the further understanding on mechanisms and functions of pathogen defense in plants by transgenic plant and exogenous quercetin. Firstly, transcription factor AtMYB44 can be induced by biotic and abiotic stress in plants defense responses against pathogen attack. It is found that AtMYB44 functions as positive regulation against PstDC3000 by SA mediated signaling pathway. Secondly, Quercetin can induce expression of AtMYB44 in Arabidopsis and enhance resistance against PstDC3000 by the level of H2O2. The mechanism of defense response against pathogen depends on SA and NPR1. Thirdly, expression of AtMYB44 is regulated by up-stream promotor methylation of AtMYB44 gene in biotic and abiotic stress.
引文
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