水稻黄单胞菌Hpa1与AvrBs2蛋白调控植物生长与抗病性的研究
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摘要
革兰氏阴性植物病原细菌Harpin与Avr蛋白代表Ⅲ型泌出蛋白的不同类型,对致病性与植物抗病防卫反应起不同作用。Avr蛋白属于致病效应因子,需要从细菌细胞转移到植物细胞,才能行使致病功能。Harpin蛋白作为转运子起作用,帮助致病效应蛋白转位。同时,无论是Harpin蛋白还是Avr蛋白,对致病性与植物抗病防卫反应都有双重影响,而Harpin蛋白还有促进植物生长的作用。对这种多重功能的分子机制,目前还不完全清楚。本文选用以前了解比较多的Harpin蛋白Hpal,研究其促进植物生长与抗病防卫反应的分子机理,探讨蛋白结构与亚细胞定位对其功能的影响。另外,还研究了水稻黄单胞菌(Xanthomonas oryzae)AvrBs2的致病作用,为今后探讨Harpin与Avr蛋白之间的功能关系提供基础。
1.Hpal利用乙烯和赤霉素信号调控植物生长与生长相关基因表达
前人研究表明,Harpin通过乙烯信号及其调控因子EIN5来促进植物生长,这一效应主要涉及植物伸展素蛋白(expansin, EXP)的作用。植物EXP构成一个很大的蛋白家族,其成员受不同激素的调控,松弛细胞壁、促进细胞伸展,影响植物不同器官或组织的生长。EXP蛋白家族何种成员受何种激素信号调控,还不完全清楚。因此,本文研究了水稻白叶枯病菌Hpa1诱导EXP基因表达与促进植物生长的关系,着重探讨乙烯与赤霉素在这个过程中所起的作用。研究结果表明,当用Hpa1分别处理拟南芥、番茄、烟草和水稻时,植物的生长速度加快,同时叶片含氮量、叶绿素含量及叶绿素a和叶绿素b的比率提高。对Hpa1诱导的EXP基因表达分析结果显示：有11个组织特异性的EXP基因被Hpa1诱导表达,其中有4个与Hpa1诱导的信号通路相关,还有4个是最新鉴定的在烟草或水稻叶片中特异表达的基因。Hpa1处理能够诱导乙烯信号通路关键基因的上调表达,而阻断乙烯信号的响应,能够抑制Hpa1诱导的EXP基因表达和植物促生长作用。在水稻中,Hpa1诱导EXP基因表达和促生长效应与GA3作用类似,而阻断赤霉素的生物合成,Hpa1诱导的EXP基因表达和促进植物生长的作用也会被抑制。由此说明,Hpa1诱导EXP基因表达和促进植物生长需要ET和GA信号的参与。
2.Hpa1蛋白生物学功能行使与促进植物生长的作用必需其N-端序列
水稻黄单胞菌的Hpa1蛋白,能够促进植物生长,增强植物防卫反应与抗逆性,但机制还没研究清楚。已有研究表明,Hpal N-端氨基酸对其在非寄主烟草上产生HR是必需的。为了研究Hpa1N-端氨基酸对其生物学功能及植物生长的影响,我们分别构建了全长Hpal和Hpal N-端缺失突变体的原核表达载体,得到了全长Hpa1蛋白和缺失N端的Hpa1蛋白(Hpa1ΔNT).分别用Hpa1和Hpa1ΔNT处理植物,发现HpalΔNT丧失了激发烟草产生HR的能力,同时也大大减弱了植物的抗病防卫反应及抗逆性。对植物生长能力的测定结果表明,Hpa1能够促进植物的营养生长,但不影响生殖生长；Hpa1处理能增强植物生长相关基因的表达和植物叶片细胞的伸长生长；Hpa1处理能促进植物叶片叶的CO2的传导及光合作用,而HpalΔNT则大大削弱了Hpa1所诱导的促生长效应。由此说明,Hpal N-端对其生物学功能的行使及促进植物生长是必需的。
3.质外体和细胞质Hpal对H2O2的产生与抗病性发挥不同作用
革兰氏阴性植物病原细菌分泌的Harpin蛋白能够激活植物防卫反应信号通路,诱导植物产生抗病性,其中包括Harpin诱导的活性氧爆发,尤其是质外体中H2O2的产生。但是,Harpin蛋白在防卫反应信号通路中是如何被识别的以及产生于植物细胞质外体中的H2O2是如何参与防卫反应的,目前还不是很清楚。本研究以水稻白叶枯病菌的Harpin蛋白Hpa1为研究对象,解析其亚细胞定位是否影响了H2O2的产生以及H2O2是如何参与调控植物抗病性的。分别把Hpal和融合了质外体定位信号基因(S)的Hpal(SHpa1)转入拟南芥,产生了转基因拟南芥植株HETAt (Hpal-expressing transgenic Arabidopsis thaliana)&SHETAt (SHpal-expressing transgenic Arabidopsis thaliana)。研究发现,转基因拟南芥对丁香假单胞菌Pst DC3000和大白菜软腐病菌Pcc RL4具有抗性。亚细胞定位显示,Hpa1定位于HETAt的细胞质及SHETAt的质外体和细胞膜。通过对植物中H2O2的检测发现,H2O2主要积累于HETAt的细胞质及SHETA的质外体和细胞质。药理学试验结果表明,H2O产生于HETAt的细胞质和SHETAt的质外体,并且产生于质外体的H2O2依赖于NOX。外源喷施Hpa1处理野生型拟南芥时,其主要定位于细胞质外体并能诱导H2O2在质外体产生,且质外体H2O2能转运至细胞质。用DPI处理喷施Hpa1的野生型拟南芥和SHETAt植物后,抑制了质外体H202的产生,且细胞质中H2O2的积累也大大减少,同时也削弱了植物的抗病能力。以上结果表明,受Hpa1诱导且在质外体产生的H2O2需要从质外体运转到细胞质中才能参与植物防卫反应过程。
4.水稻条斑病菌Ⅲ型效应因子AvrBs2的病理功能研究
水稻黄单胞菌通过Ⅲ型分泌系统将大量的效应因子注入寄主细胞,使寄主产生抗(感)病性。虽然越来越多的Ⅲ型效应蛋白包括Avr蛋白被鉴定出来,但是关于其蛋白结构与生物学功能的研究目前还不是很多。AvrBs2是第一个被证明能够增强病原细菌在寄主植物中繁殖的Ⅲ型效应蛋白,能够增强野油菜黄单胞菌Xanthomonas campestris pv. vesicatoria(Xcv)的毒性,但是高度保守的AvrBs2效应因子在其它黄单胞菌中的生物学功能目前研究的还不是很清楚,尤其是水稻黄单胞菌。本文拟研究Xoc菌株RS105的AvrBs2效应因子的蛋白结构与生物学功能。通过同源交换,我们获得了avrBs2xoc基因插入突变体。接种试验结果表明,avrBs2突变体和野生型虽然在烟草上引起过敏反应的能力没有明显差异,但突变体却显著降低了病原菌在水稻IR24上的致病性,同时增强了植物病程相关蛋白基因PR-laΔPR-1b的表达。由此我们判断,AvrBs2xoc是一个致病效应因子,且能抑制寄主植物的免疫反应。
Harpin and Avr proteins are representative type III effectors secreted by Gram-negative phytopathogenic bacteria, that regulate the pathogenicity and the plant defense response to pathogens. Avr proteins are pathogenic effectors and must be translocated from bacteria into plant cell to pathopoiesis. Harpin protein function as a translocator to translocate the effectors. Whether the Harpin or Avr proteins are double effects on the pathogenicity and plant defense response to pathogens, and also the Harpin can induce plant growrh enhancement, however, the molecular mechanism is unclear. In this study, we chose the Hpal protein, focusing on analysis of the molecular mechanism that plant growth enhancement and pathogen defense response induced by Hpal and the effects of the protein structure and subcellular localization on its function. We also analysis the function of the type III effector protein AvrBs2produced by X. oryzae, and focusing on analysis of its effects on pathogenicity.
1. Expansin gene expression and plant growth regulated by ethylene and gibbrellin in response to Hpal
In early studys, ethylene signal and regulator of EIN5can induce plant growth enhancement in response to Harpin and the expansin protein act importantly on the effects. Expansins act to loose cell walls and modulate growth of the cell and plant mediated by some hormones. However, it's not clear whether a hormone is specific and distinct hormones interact to regulate EXP activity. In this study, we report the effect that expansin gene expression and plant growth enhancement (PGE) in response to Hpal, the type-Ⅲ effector produced by X. oryzae pv. oryzae and the roles of ethylene and gibbrellin in both responses. Our results show that when applied to Arabidopsis thaliana, tomato, tobacco and rice, Hpal can markedly increase these plants growth concomitantly with the leaf nitrogen, chlorophylls and chlorophyll a/b ratio, which can basally affect plant biosynthesis and productivity. RT-PCR assay of EXP genes revealed that eleven of organ-specific EXPs expression was affected in response to Hpal. Four of eleven EXPs behaved in consistence with the Hpal signaling process and the other four were newly identified as specifically expressed in leaves of tobacco or rice. Using genetic and chemical methods to block ET perception can arrested EXP expression and PGE in the four plants in response to Hpal.In rice, Hpal simulated GA3to induce EXP expression and PGE. Whether disrupting either of ET perception or GA biosynthesis, both effects were impaired. These results suggest that both events of EXP expression and PGE require GA and ET in plants responding to Hpal.
2. The Hpal Harpin needs nitroxyl terminus to promote vegetative growth and leaf photosynthesis in Arabidopsis
Hpal is a Harpin protein produced by Xanthomonas oryzae, an important bacterial pathogen of rice, and has the growth-promoting activity in plants. To understand the molecular basis for the function of Hpal, we generated an inactive variant protein, HpalΔNT, by deleting the nitroxyl-terminal region of the Hpal sequence and compared HpalΔNT with the full-length protein in terms of the effects on vegetative growth and related physiological responses in Arabidopsis. When Hpal was applied to plants, it acted to enhance the vegetative growth but did not affect the floral development. Enhanced plant growth was accompanied by induced expression of growth-promoting genes and cell growth in plant leaves. The growth-promoting activity of Hpal was further correlated with a physiological consequence shown as promotions in stomatal and mesophyll CO2conductance and leaf photosynthesis. On the contrary, plant growth, growth-promoting gene expression, and the physiological consequence changed little in response to the HaplΔNT treatment. These analyses suggest that Hpal requires the nitroxyl-terminus to facilitate CO2transport inside leaf cells and promote leaf photosynthesis and vegetative growth of the plant.
3. Apoplastic and cytoplasmic location of Harpin protein Hpal plays different roles in H2O2generation and pathogen resistance in Arabidopsis.
Harpins produced by Gram-negative phytopathogenic bacteria have been shown to activate plant defense pathways which involve the production of the reactive oxygen species, especially transduction of hydrogen peroxide signal generated in the apoplast. But how a Harpin is recognized in the pathway and how the apoplastic H2O2anticipates in defense responses are not clear. Here we use Hpal, a Harpin protein that produced by rice bacterial leaf blight pathogen X. oryzae pv. oryzae, to study if the cellular location of it impacts H2O2production and how the H2O2regulates pathogen resistance in nonhost Arabidopsis thaliana. Hpal gene and Hpal fused to an apoplastic localization signal (S) were transformed into Arabidopsis thaliana using the Agrobacterium tumefaciens and generated HETAt (Hpal-expressing transgenic Arabidopsis thaliana) and SHETAt (SHpal-expressing transgenic Arabidopsis thaliana) transgenic plants, respectively. Our results show that the HETAt and SHETAt plants are resistant to pathogen Pseudomonas syringae pv. tomato DC3000and Pectobacterium carotovora subsp. carotovora RL4. In HETAt, Hpal was found to associate with the cytoplasm and in SHETAt it was found in the apoplast and cytomembrane, which accompany with H2O2accumulation in cytoplasts in HETAt and in apoplasts of SHETAt as well. This study also demonstrate that H2O2was generated in cytoplasm in HETAt and was generated in a NOX-dependent manner in apoplasts in SHETAt. When applied to Arabidopsis, Hpal located and induced H2O2generation in apoplasts and caused H2O2accumulation in both apoplasts and cytoplasts. Treatment with the SHETAt and Hpal-treated parent plants using DPI, however, can inhibit apoplastic H2O2generation accompanying with less cytoplasmic H2O2accumulation and reduce plant resistance to bacterial pathogens. These results suggest that the apoplastic H2O2generation induced by Hpal is subject to translocation into cytoplasm for participation in the regulation of pathogen resistance in the plant.
4. The function study of Avrbs2protein that secreted through the type III secretion system of the Xanthomonas oryzae pv. oryzicola
Xanthomonas oryzae is the causal agent of bacterial disease in rice which can secrete many effectors into plant cells through the type III secretion apparatus. More and more type III effectors including Avr proteins have been indentificated, however, the function studys of many of them are unclear. AvrBs2is the first type III effector that has been shown to enhance bacterial multiplication within host tissue, a virulence contribution only observed in the strains of X. campestris pv.vesicatoria. However, the function of other Xanthomonads AvrBs2proteins is unclear though they are widely conserved in many Xanthomonads especially in Xanthomonas oryzae. Here, we examined the function of AvrBs2by inserted mutation in X. oryzae pv. oryzicola RS105. The mutation of avrBs2xoc resulted in reduced pathogenicity on host rice plants IR24and enhanced the expression of pathogen-related1a and1b protein relative to the wild-type strain RS105, but not affect the hypersensitive response (HR) in non-host Nicotiana benthamiana. These results suggest that the AvrBs2xoc effector protein is a virulence factor and suppress the host defence response in plant.
1.Hpal利用乙烯和赤霉素信号调控植物生长与生长相关基因表达
前人研究表明,Harpin通过乙烯信号及其调控因子EIN5来促进植物生长,这一效应主要涉及植物伸展素蛋白(expansin, EXP)的作用。植物EXP构成一个很大的蛋白家族,其成员受不同激素的调控,松弛细胞壁、促进细胞伸展,影响植物不同器官或组织的生长。EXP蛋白家族何种成员受何种激素信号调控,还不完全清楚。因此,本文研究了水稻白叶枯病菌Hpa1诱导EXP基因表达与促进植物生长的关系,着重探讨乙烯与赤霉素在这个过程中所起的作用。研究结果表明,当用Hpa1分别处理拟南芥、番茄、烟草和水稻时,植物的生长速度加快,同时叶片含氮量、叶绿素含量及叶绿素a和叶绿素b的比率提高。对Hpa1诱导的EXP基因表达分析结果显示：有11个组织特异性的EXP基因被Hpa1诱导表达,其中有4个与Hpa1诱导的信号通路相关,还有4个是最新鉴定的在烟草或水稻叶片中特异表达的基因。Hpa1处理能够诱导乙烯信号通路关键基因的上调表达,而阻断乙烯信号的响应,能够抑制Hpa1诱导的EXP基因表达和植物促生长作用。在水稻中,Hpa1诱导EXP基因表达和促生长效应与GA3作用类似,而阻断赤霉素的生物合成,Hpa1诱导的EXP基因表达和促进植物生长的作用也会被抑制。由此说明,Hpa1诱导EXP基因表达和促进植物生长需要ET和GA信号的参与。
2.Hpa1蛋白生物学功能行使与促进植物生长的作用必需其N-端序列
水稻黄单胞菌的Hpa1蛋白,能够促进植物生长,增强植物防卫反应与抗逆性,但机制还没研究清楚。已有研究表明,Hpal N-端氨基酸对其在非寄主烟草上产生HR是必需的。为了研究Hpa1N-端氨基酸对其生物学功能及植物生长的影响,我们分别构建了全长Hpal和Hpal N-端缺失突变体的原核表达载体,得到了全长Hpa1蛋白和缺失N端的Hpa1蛋白(Hpa1ΔNT).分别用Hpa1和Hpa1ΔNT处理植物,发现HpalΔNT丧失了激发烟草产生HR的能力,同时也大大减弱了植物的抗病防卫反应及抗逆性。对植物生长能力的测定结果表明,Hpa1能够促进植物的营养生长,但不影响生殖生长；Hpa1处理能增强植物生长相关基因的表达和植物叶片细胞的伸长生长；Hpa1处理能促进植物叶片叶的CO2的传导及光合作用,而HpalΔNT则大大削弱了Hpa1所诱导的促生长效应。由此说明,Hpal N-端对其生物学功能的行使及促进植物生长是必需的。
3.质外体和细胞质Hpal对H2O2的产生与抗病性发挥不同作用
革兰氏阴性植物病原细菌分泌的Harpin蛋白能够激活植物防卫反应信号通路,诱导植物产生抗病性,其中包括Harpin诱导的活性氧爆发,尤其是质外体中H2O2的产生。但是,Harpin蛋白在防卫反应信号通路中是如何被识别的以及产生于植物细胞质外体中的H2O2是如何参与防卫反应的,目前还不是很清楚。本研究以水稻白叶枯病菌的Harpin蛋白Hpa1为研究对象,解析其亚细胞定位是否影响了H2O2的产生以及H2O2是如何参与调控植物抗病性的。分别把Hpal和融合了质外体定位信号基因(S)的Hpal(SHpa1)转入拟南芥,产生了转基因拟南芥植株HETAt (Hpal-expressing transgenic Arabidopsis thaliana)&SHETAt (SHpal-expressing transgenic Arabidopsis thaliana)。研究发现,转基因拟南芥对丁香假单胞菌Pst DC3000和大白菜软腐病菌Pcc RL4具有抗性。亚细胞定位显示,Hpa1定位于HETAt的细胞质及SHETAt的质外体和细胞膜。通过对植物中H2O2的检测发现,H2O2主要积累于HETAt的细胞质及SHETA的质外体和细胞质。药理学试验结果表明,H2O产生于HETAt的细胞质和SHETAt的质外体,并且产生于质外体的H2O2依赖于NOX。外源喷施Hpa1处理野生型拟南芥时,其主要定位于细胞质外体并能诱导H2O2在质外体产生,且质外体H2O2能转运至细胞质。用DPI处理喷施Hpa1的野生型拟南芥和SHETAt植物后,抑制了质外体H202的产生,且细胞质中H2O2的积累也大大减少,同时也削弱了植物的抗病能力。以上结果表明,受Hpa1诱导且在质外体产生的H2O2需要从质外体运转到细胞质中才能参与植物防卫反应过程。
4.水稻条斑病菌Ⅲ型效应因子AvrBs2的病理功能研究
水稻黄单胞菌通过Ⅲ型分泌系统将大量的效应因子注入寄主细胞,使寄主产生抗(感)病性。虽然越来越多的Ⅲ型效应蛋白包括Avr蛋白被鉴定出来,但是关于其蛋白结构与生物学功能的研究目前还不是很多。AvrBs2是第一个被证明能够增强病原细菌在寄主植物中繁殖的Ⅲ型效应蛋白,能够增强野油菜黄单胞菌Xanthomonas campestris pv. vesicatoria(Xcv)的毒性,但是高度保守的AvrBs2效应因子在其它黄单胞菌中的生物学功能目前研究的还不是很清楚,尤其是水稻黄单胞菌。本文拟研究Xoc菌株RS105的AvrBs2效应因子的蛋白结构与生物学功能。通过同源交换,我们获得了avrBs2xoc基因插入突变体。接种试验结果表明,avrBs2突变体和野生型虽然在烟草上引起过敏反应的能力没有明显差异,但突变体却显著降低了病原菌在水稻IR24上的致病性,同时增强了植物病程相关蛋白基因PR-laΔPR-1b的表达。由此我们判断,AvrBs2xoc是一个致病效应因子,且能抑制寄主植物的免疫反应。
Harpin and Avr proteins are representative type III effectors secreted by Gram-negative phytopathogenic bacteria, that regulate the pathogenicity and the plant defense response to pathogens. Avr proteins are pathogenic effectors and must be translocated from bacteria into plant cell to pathopoiesis. Harpin protein function as a translocator to translocate the effectors. Whether the Harpin or Avr proteins are double effects on the pathogenicity and plant defense response to pathogens, and also the Harpin can induce plant growrh enhancement, however, the molecular mechanism is unclear. In this study, we chose the Hpal protein, focusing on analysis of the molecular mechanism that plant growth enhancement and pathogen defense response induced by Hpal and the effects of the protein structure and subcellular localization on its function. We also analysis the function of the type III effector protein AvrBs2produced by X. oryzae, and focusing on analysis of its effects on pathogenicity.
1. Expansin gene expression and plant growth regulated by ethylene and gibbrellin in response to Hpal
In early studys, ethylene signal and regulator of EIN5can induce plant growth enhancement in response to Harpin and the expansin protein act importantly on the effects. Expansins act to loose cell walls and modulate growth of the cell and plant mediated by some hormones. However, it's not clear whether a hormone is specific and distinct hormones interact to regulate EXP activity. In this study, we report the effect that expansin gene expression and plant growth enhancement (PGE) in response to Hpal, the type-Ⅲ effector produced by X. oryzae pv. oryzae and the roles of ethylene and gibbrellin in both responses. Our results show that when applied to Arabidopsis thaliana, tomato, tobacco and rice, Hpal can markedly increase these plants growth concomitantly with the leaf nitrogen, chlorophylls and chlorophyll a/b ratio, which can basally affect plant biosynthesis and productivity. RT-PCR assay of EXP genes revealed that eleven of organ-specific EXPs expression was affected in response to Hpal. Four of eleven EXPs behaved in consistence with the Hpal signaling process and the other four were newly identified as specifically expressed in leaves of tobacco or rice. Using genetic and chemical methods to block ET perception can arrested EXP expression and PGE in the four plants in response to Hpal.In rice, Hpal simulated GA3to induce EXP expression and PGE. Whether disrupting either of ET perception or GA biosynthesis, both effects were impaired. These results suggest that both events of EXP expression and PGE require GA and ET in plants responding to Hpal.
2. The Hpal Harpin needs nitroxyl terminus to promote vegetative growth and leaf photosynthesis in Arabidopsis
Hpal is a Harpin protein produced by Xanthomonas oryzae, an important bacterial pathogen of rice, and has the growth-promoting activity in plants. To understand the molecular basis for the function of Hpal, we generated an inactive variant protein, HpalΔNT, by deleting the nitroxyl-terminal region of the Hpal sequence and compared HpalΔNT with the full-length protein in terms of the effects on vegetative growth and related physiological responses in Arabidopsis. When Hpal was applied to plants, it acted to enhance the vegetative growth but did not affect the floral development. Enhanced plant growth was accompanied by induced expression of growth-promoting genes and cell growth in plant leaves. The growth-promoting activity of Hpal was further correlated with a physiological consequence shown as promotions in stomatal and mesophyll CO2conductance and leaf photosynthesis. On the contrary, plant growth, growth-promoting gene expression, and the physiological consequence changed little in response to the HaplΔNT treatment. These analyses suggest that Hpal requires the nitroxyl-terminus to facilitate CO2transport inside leaf cells and promote leaf photosynthesis and vegetative growth of the plant.
3. Apoplastic and cytoplasmic location of Harpin protein Hpal plays different roles in H2O2generation and pathogen resistance in Arabidopsis.
Harpins produced by Gram-negative phytopathogenic bacteria have been shown to activate plant defense pathways which involve the production of the reactive oxygen species, especially transduction of hydrogen peroxide signal generated in the apoplast. But how a Harpin is recognized in the pathway and how the apoplastic H2O2anticipates in defense responses are not clear. Here we use Hpal, a Harpin protein that produced by rice bacterial leaf blight pathogen X. oryzae pv. oryzae, to study if the cellular location of it impacts H2O2production and how the H2O2regulates pathogen resistance in nonhost Arabidopsis thaliana. Hpal gene and Hpal fused to an apoplastic localization signal (S) were transformed into Arabidopsis thaliana using the Agrobacterium tumefaciens and generated HETAt (Hpal-expressing transgenic Arabidopsis thaliana) and SHETAt (SHpal-expressing transgenic Arabidopsis thaliana) transgenic plants, respectively. Our results show that the HETAt and SHETAt plants are resistant to pathogen Pseudomonas syringae pv. tomato DC3000and Pectobacterium carotovora subsp. carotovora RL4. In HETAt, Hpal was found to associate with the cytoplasm and in SHETAt it was found in the apoplast and cytomembrane, which accompany with H2O2accumulation in cytoplasts in HETAt and in apoplasts of SHETAt as well. This study also demonstrate that H2O2was generated in cytoplasm in HETAt and was generated in a NOX-dependent manner in apoplasts in SHETAt. When applied to Arabidopsis, Hpal located and induced H2O2generation in apoplasts and caused H2O2accumulation in both apoplasts and cytoplasts. Treatment with the SHETAt and Hpal-treated parent plants using DPI, however, can inhibit apoplastic H2O2generation accompanying with less cytoplasmic H2O2accumulation and reduce plant resistance to bacterial pathogens. These results suggest that the apoplastic H2O2generation induced by Hpal is subject to translocation into cytoplasm for participation in the regulation of pathogen resistance in the plant.
4. The function study of Avrbs2protein that secreted through the type III secretion system of the Xanthomonas oryzae pv. oryzicola
Xanthomonas oryzae is the causal agent of bacterial disease in rice which can secrete many effectors into plant cells through the type III secretion apparatus. More and more type III effectors including Avr proteins have been indentificated, however, the function studys of many of them are unclear. AvrBs2is the first type III effector that has been shown to enhance bacterial multiplication within host tissue, a virulence contribution only observed in the strains of X. campestris pv.vesicatoria. However, the function of other Xanthomonads AvrBs2proteins is unclear though they are widely conserved in many Xanthomonads especially in Xanthomonas oryzae. Here, we examined the function of AvrBs2by inserted mutation in X. oryzae pv. oryzicola RS105. The mutation of avrBs2xoc resulted in reduced pathogenicity on host rice plants IR24and enhanced the expression of pathogen-related1a and1b protein relative to the wild-type strain RS105, but not affect the hypersensitive response (HR) in non-host Nicotiana benthamiana. These results suggest that the AvrBs2xoc effector protein is a virulence factor and suppress the host defence response in plant.
引文
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