摘要
植物在整个生长过程中都会受到各种病菌的侵染,为了生存及保护自己植物进化出了自己独特的免疫系统。是否能够迅速准确地启动免疫反应从而成功抵抗病菌侵染,主要依赖于植物能否适时而准确地调控自身防卫相关基因的转录表达。植物察觉病菌侵染后,会将胁迫信号传递到细胞核内,激活转录因子并将防卫信号放大并且传递,进而诱发下游一系列防卫相关基因的表达。在众多的植物基因的转录调控机制中转录因子和染色质修饰介导的基因转录调控是植物基因转录调控最重要的两大调控机制。染色质修饰主要包括围绕核小体的组蛋白修饰和依赖ATP提供能量的染色质重塑复合体蛋白介导的染色质构型调控。本研究证明转录因子和染色质修饰相关基因介导的基因转录调控参与拟南芥抗病防卫反应。
1.组蛋白H2B单泛素化调控拟南芥中抗病基因的表达
抗病基因调控是植物免疫反应的最重要调控环节。本章研究发现拟南芥E3连接酶基因HUB1(HISTONE MONOUBIQUITINATION1)和HUB2参与自体免疫突变体bon1中R(Resistance)基因SNC1(SUPPRESSOR OF nprl-1, CONSTITUTIVE1)的表达调控。SNC1在bon1中组成性上调表达,同时NC1基因位点H2B单泛素化(H2Bub)的水平比野生型显著地增加。突变HUB1或者其同源基因HUB2都可以显著地抑制bon1的自体免疫反应。SNC1在bon1中的上调表达与其基因位点H2Bub修饰相关。同时我们发现在野生型拟南芥中病菌侵染后SNC1和HUB1的表达及SNC1基因位点的H2Bubi的水平都会增加。以上结果证明H2Bub在自体免疫突变体bon1的R基因调控中起着关键的作用,并且H2Bub很有可能参与了病菌侵染后R基因的表达调控。
2.拟南芥CHD亚类染色质重塑基因CHR5在拟南芥免疫反应中的功能研究
依赖ATP提供能量的染色质重塑蛋白复合体可以通过水解ATP获得能量改变染色质的构型,染色质构型改变是真核生物调控基因转录表达的重要机制之一。本研究通过对一个拟南芥CHD类染色质重塑基因CHR(Chromatin Remodeling5)的鉴定和功能分析,证明其与植物免疫反应相关。通过对自体免疫突变体bon1潜在的抑制突变体的遗传筛选发现CHR5功能缺失突变体可以显著地抑制bon1的免疫反应。bon1中组成性激活表达的NB-LRR类的R基因SNC1及PR基因及植物防卫激素水杨酸(SA)的累积,都显著地被CHR5基因突变抑制。bon1突变体对丁香假单胞菌番茄致病变种Pst DC3000(Pseudomonas syringae pv. tomato DC3000)的组成性抗性也被CHR5基因突变明显的抑制。同时揭示了CHR5与HUB1和DDM1在bon1相关的免疫反应调控中的相互关系。通过对chr5突变体的对病菌Psm E4326(Pseudomonas. syringae pv. maculicola ES4326)的抗病实验证明CHR5正调控拟南芥对病菌的基本防卫反应。最后通过RNAseq对CHR5基因突变体进行了全基因组转录表达分析发现CHR5主要与一些植物防卫反应相关的基因的转录表达相关,并证明CHR5可能主要参与植物的胁迫应答反应。
3. AtMYB44通过水杨酸信号通路正调控拟南芥对丁香假单胞菌的抗性
AtMYB44属于植物R2R3型第22亚类的MYB转录因子。研究证明AtMYB44是一个重要的胁迫响应基因。本研究发现AtMYB44基因的表达能够被植物激素水杨酸(salicylic acid,SA)处理和半活体营养型病菌丁香假单胞菌番茄致病变种(Pseudomonas syringae pv. tomato DC3000, Pst DC3000)侵染显著地诱导表达,且AtMYB44的诱导表达需要SA的积累及NPR1蛋白的参与。AtMYB44过表达的转基因拟南芥比野生型显著地更抗pst DC3000的侵染,且病菌侵染后过表达植物会有更强PR1激活表达并产生更多的活性氧。相反AtMYB44基因功能缺失突变体atmyb44’比野生型更感病,同时病菌侵染后atmyb44突变体中PR1的激活表达也会减弱。进一步的遗传杂交实验证明,AtMYB44过表达植物的所有抗病表型都需要SA积累和NPR1蛋白的参与。上述结果证明AtMYB44通过依赖于NPR1的SA防卫信号通路正调控拟南芥对Pst DC3000的抗病性。
Due to their sessile lifestyle, plants are in constant conflict with pathogens throughout their life cycles. In order to fend off a wide range of microbial pathogen attacks, plants have developed sophisticated immunity system to defend themselves. Plant immunity responses against pathogen infection largely depend on the plant's capacity to reprogram rapidly and specifically the transcription of a large number of the host genes. In a pathogen stress type-dependent manner, external signals are trans-located into the nucleus to activate transcription factors, resulting in the increased expression of particular sets of defense-related genes. Among mechanisms of transcriptional regulation, transcription factors and chromatin modification accomplished through the activity of histone-modifying enzymes and ATP dependent chromatin-remodeling complexes are emerging as two key processes in the orchestration of plant immunity responses.
1. Regulation of disease resistance gene expression by H2B monoubiquitination in Arabidopsis
Regulation of disease resistance (R) genes is one key event in plant immunity response. Here we show that two Arabidopsis E3ubiquitin ligase genes HUB1(HISTONE MONOUBIQUITINATION1) and HUB2are involved in the regulation of the R gene SNC1(SUPPRESSOR OFnpr1-1, CONSTITUTIVE1) in the autoimmune mutant bon1. The SNC1gene is up-regulated in bon1and the SNC1locus has a higher monoubiquitination of H2B. The loss of HUB1or HUB2function largely suppresses the autoimmune phenotypes in bon1. Thus the up-expression of SNC1in bon1is dependent on H2B modification at the SNC1locus. Furthermore, the expression of SNC1and HUB1are moderately up-regulated by pathogen infection and H2B monoubiquitination at SNC1is enhanced by pathogen infection. Together, this study indicates that H2B monoubiquitination is critical for R gene up-regulation in autoimmune mutants and it might also contribute to R gene regulation during immune responses.
2. Characterize and function studies of one plant helicase-like domain (CHD) chromatin remodeling factor chromatin remodeling5(CHR5) in Arabidopsis innate immunity
ATP-dependent chromatin remodeling complexes use the energy of ATP hydrolysis to alter the structure of chromatin, which is an important mechanism in the regulation of eukaryotic gene expression. In this study, we characterized the function of chromatin remodeling5(CHR5), a member of plant helicase-like domain(CHD) chromatin remodeling factor, in Arabidopsis innate immunity. We found that knock out CHR5gene can largely suppress the autoimmunity in bon1-1by a genetic screen of the potential suppressor of the autoimmunity mutant bon1-1.The constitutively expression of the SNC1and PR1gene and the accumulation of SA in bon1-1were all largely suppressed by CHR5mutation. And the constitutively pathogen resistance in bon1-1was also disappeared in the chr5bon1-1double mutants.While chr5single mutants exhibit defects in basal resistance against the bacterial pathogen P. syringae pv.maculicola ES4326(Psm E4326). Furthermore, we approved that CHR5may regulate the expression of SNC1in a locus specific manner. After RNAseq data analysis, we found that the expression many important defense response genes were significantly changed in chr5mutants. Our study indicates that CHR5is a negative regulator of plant immunity and may regulate gene expression at chromatin level.
3. AtMYB44positively modulates disease resistance to Pseudomonas syringae through the salicylic acid signalling pathway in Arabidopsis
AtMYB44which belongs to sub-group22of the R2R3MYB transcription factor family is one well known stress response related gene. Here, we found that the expression of AtMYB44is sensitive to the phytophormone salicylic acid (SA) treatment and the hemibiotrophic pathogen Pseudomonas syringae pv. tomato DC3000(Pst DC3000) infection. And the induction expression of AtMYB44by Pst DC3000requires both the SA accumulation and the NPR1gene. AtMYB44over-expression up-regulated PR genes induction expression and ROS accumulation by Pst DC3000, leading to enhanced resistance to the Pst DC3000. While the knockout mutant atmyb44shows opposite effects. Induction of PR1by Pst DC3000is reduced in atmyb44mutants. AtMYB44over-expression phenotypes, such as retarded growth, up-regelated PR1and resistance to Pst DC3000are reversed by SA depletion and NPR1mutation. Thus, AtMYB44positive regulates Arabidopsis defense response against Pst DC3000through SA signaling pathway.
引文
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