质外体过氧化氢与水通道蛋白PIP1；4对Hpa1蛋白诱导拟南芥抗病性的调控作用
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摘要
Harpin是由革兰氏阴性植物病原细菌产生的性质和功能相似的一类蛋白质,影响病原菌在寄主植物上的致病性和非寄主植物上的过敏反应(HR)。外施的harpin能诱导植物抗病、抗虫和抗旱,并且也能促进植物的生长。Harpin的这些多效性是由于激活了多种不同的信号通路,包括植物激素的转导、活性氧簇信号、离子通道、细胞程序性死亡、系统获得抗性(SAR)或水杨酸和蛋白质激酶级联等。到目前为止,harpin几乎从所有的革兰氏阴性植物病原细菌中被鉴定出来。Harpin能激活植物生长和防卫反应信号通路,但是,在植物中,一种harpin蛋白在一个信号通路中如何能被识别?一个特定的harpin在植物中的受体是什么?这个信号如何被转导到细胞内?能把细菌效应蛋白,特别是Ⅲ型效应子从植物细胞外转运到细胞内的植物转位子是什么?目前都还不清楚。
本研究使用HpalXoo,它由水稻白叶枯病原细菌(Xanthomonas oryzae pv. oryzae)产生,分子量15.6kD,比其他病原细菌的harpin(最大44kD)都小,但生物活性强。本研究着重剖析HpalXoo在植物中的亚细胞定位与后续反应之间的关系,质外体的H_2O_2如何参与植物防卫反应,HpalXoo蛋白在信号通路中如何被植物中的蛋白识别以及识别之后对植物产生的影响。
1.转基因植物产生的HpalXoo蛋白在质外体定位诱导H2O2产生与向细胞质转移及对抗病性的影响
已有的研究证明,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,包括产生于质外体的H2O2信号的转导,但是,一种harpin蛋白在一个信号通路中如何被识别以及质外体的H2O2如何参与防卫反应,还不清楚。本章中,我们研究了来源于水稻白叶枯病原细菌(Xanthomonas oryzae pv. oryzae)的harpin蛋白HpalXoo在拟南芥中是否影响了H2O2的产生以及植物的抗病性。把HpalXoo和融合了质外体定位信号基因(S)的HpalXoo (SHpalxoo)转入拟南芥,分别产生了转基因拟南芥植株HETAt和SHETAt。本研究发现,转基因拟南芥抗丁香假单胞番茄致病变种菌株DC3000(Pst DC3000)和大白菜软腐病菌菌株RL4(Pcc RL4)。HpalXoo定位于HETAt的细胞质和SHETAt的质外体和细胞膜,H2O2积累于HETAt的细胞质和SHETA的质外体和细胞质。已经知道,质外体中的H2O2是由质膜上的烟酰胺腺嘌呤二核苷酸磷酸氧化酶(NOX)产生的,这是植物防卫反应的一个普遍特征。H2O2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。当HpalXoo处理WT植物时,HpalXoo定位于质外体并在质外体中诱导H2O2的产生,但是质外体和细胞质中都有H2O2的积累。在SHETAt和HpalXoo处理的野生型(WT)拟南芥中,如果抑制质外体中H2O2的产生,细胞质中就没有H2O2的积累并且植物失去抗病能力。这些结果说明,HpalXoo诱导的产生于质外体的H202在参与植物调节抗病过程中需要从质外体运转到细胞质中。
2. HpalXoo促进转基因拟南芥生长的作用不依赖于H2O2
已有的研究证明,harpin能激活植物的生长信号通路。但是,一种harpin蛋白在一个信号通路中如何被识别以及harpin蛋白诱导的H2O2是否参与生长信号通路,还不清楚。在第一章中,我们产生了转基因拟南芥植株HETAt和SHETAt。本研究发现,转基因拟南芥植株HETAt和SHETAt的生长快于对照植株,并且植株体内的与生长相关基因表达增强。在SHETAt中,H2O2的积累依赖于质膜NADPH氧化酶活性,并且H2O2的积累不影响植株的生长。在HETAt中,H2O2含量的上升与NADPH氧化酶无关,H2O2的产生也不影响植株的生长。外施的HpalXoo能促进野生型拟南芥(WT)的生长,HpalXoo诱导的产生于质外体的H2O2以及积累于质外体和细胞内的H202与促生长无关。这些实验结果说明,HpalXoo能够激活植物的生长信号通路,并且这个信号通路与HpalXoo诱导的H2O2无关。
3. Hpalxoo与PIP1；4蛋白在拟南芥细胞膜上的互作验证
前两章已经证明,转入拟南芥的HpalXoo定位于植物的细胞外和细胞内,诱导植物的防卫反应,并能促进植物生长。基于HpalXoo的这些有益表型,本章中,为了试图解析HpalXoo的作用机理,以HpalXoo为诱饵对拟南芥cDNA文库进行筛选,初步筛选到6个阳性克隆,其中包括植物水通道蛋白PIP1；4。通过普通的酵母双杂交体系(Yeast two-hybrid system, Y2H)、膜酵母双杂交体系(Membrane yeast two-hybrid system, MYTH)、Pull-down assay、双分子荧光互补技术(Bimolecular Fluorescence Complementation, BiFC)和荧光染料的观察证实了HpalXoo与PIP1；4的互作,并且互作发生在植物的细胞膜上。而N端缺失53个氨基酸的HpalXoo蛋白(ANT)则不能与PIP1；4发生相互作用。
4. HpalXoo与PIP1；4蛋白互作促进拟南芥的生长与诱导其抗病的初步研究
证实HpalXoo与PIP1；4互作之后,为了研究这两个蛋白的互作对植物产生的影响,从拟南芥研究中心购买了8种atpipl;4拟南芥(CS803583、CS870828、CS872202、 CS876999、CS879846、CS879691、CS870571和SALK_147568)种子。这些突变体都是T2代杂合体,通过卡那霉素和Basta抗性筛选,从每一种突变体中都筛选到了纯合体。利用原核表达体系表达了HpalXoo和ΔNT并用蛋白纯化试剂盒纯化了这两个蛋白。通过表型观察发现,HpalXoo处理的WT拟南芥长势最好,而HpalXoo处理的atpip1;4拟南芥与ANT或水处理的WT和atpip1;4拟南芥生长表型一致,长势都弱于HpalXoo处理的WT。Pst DC3000接种实验发现,HpalXoo处理的WT拟南芥抗病最强,而HpalXoo处理的atpip1;4拟南芥与ΔNT或水处理的WT和atpip1;4拟南芥易感病。这些试验结果说明,PIP1；4与HpalXoo的互作不仅能够促进拟南芥的生长,也能够诱导拟南芥对病原菌的抗性。
Harpin proteins produced by gram-negative phytopathogenic bacteria generally affect virulence to host plants and the hypersensitive reaction (HR) of nonhost plants. When applied to plants, any harpins can induce resistance to pathogens and insects, confer drought tolerance, and enhance plant growth. The diverse effects are attributable to the activation of distinct signaling pathways that involve transduction of phytohormones, reactive oxygen species (ROS) signals, ion channels, programmed cell death, systemic acquired resistance (SAR) or salicylic acid (SA) signal pathway and protein kinase cascade, etc. Currently, harpins have been identified from almost all the Gram-negative phytopathogenic bacteria. Harpin have been shown to activate plant growth and defense signaling pathways, but the recognition of a harpin connecting with a pathway in plants, what a plant receptor of a particular harpin protein is, the transduction of the signal to a cellular pathway and what plant translocator (s) for plant-cytoplasmic trafficking of bacterial effector proteins is (are), especially type-Ⅲ effectors have long been in question.
The Hpalxoo protein produced by the rice bacterial leaf blight pathogen Xanthomonas oryzae pv. oryzae belongs to the harpin group of proteins. This study focuses on analysis of the relationship between cellular location of Hpalxoo and subsequent responses in plants, the apoplastic H2O2participation in defenses, the Hpalxoo recognition by a plant receptor and the impact of perception.
1. Apoplastic location of harpin protein HpalXoo induces apoplastic generation and cytoplasmic translocation of H2O2required for pathogen resistance in Arabidopsis
Harpin proteins secreted by Gram-negative phytopathogenic bacteria have been shown to activate plant defense pathways that involve transduction of hydrogen peroxide (H2O2) signal generated in the apoplast, but how a harpin is recognized connecting with a pathway and how the apoplastic H2O2anticipates in defenses are to be elucidated. Here we study if the cellular location of Hpalxoo, a harpin protein produced by rice bacterial leaf blight pathogen Xanthomonas oryzae pv. oryzae, impacts H2O2production and pathogen resistance in Arabidopsis thaliana. Transformation with the Hpalxoo gene and Hpalxoo fused to an apoplastic localization signal (SHpal Xoo) generated HpalXoo-and SHpal Xoo-expressing transgenic A. thaliana (HETAt and SHETAt) plants, respectively. HpalXoo was found to associate with the cytoplasm in HETAt and the apoplast in SHETAt, accompanying H2O2accumulation in cytoplasts and in apoplasts of SHETAt4as well. Apoplastic H2O2production via nicotinamide adenine dinucleotide phosphate (NADPH) oxidase (NOX) located in the plasma membrane has been shown as a common feature of plant defenses. In SHETAt, H2O2was generated in apoplasts in a NOX-dependent manner but accumulated to a greater extent in apoplasts than in cytoplasts. When applied to the parent plant, Hpalxoo located and induced H2O2generation in apoplasts but caused H2O2accumulation in both apoplasts and cytoplasts. In SHETAt and HpalXoo-treated parent plants, however, inhibition of apoplastic H2O2generation abrogated cytoplasmic H2O2accumulation and plant resistance to bacterial pathogens. These results suggest that the apoplastic H2O2is subject to a cytoplasmic translocation for participation in the regulation of pathogen resistance in the plant.
2. Growth enhancement of Hpalxoo transgenic Arabidopsis is not related to Hpalxoo-induced H2O2
The Hpalxoo protein produced by the rice bacterial leaf blight pathogen Xanthomonas oryzae pv. oryzae belongs to the harpin group of proteins secreted by Gram-negative phytopathogenic bacteria. In molecule mass, Hpal is smaller than other harpins, only15.6KD. harpin have been shown to activate plant growth signaling pathways, but how a harpin is recognized connecting with the pathway and how the harpin-induced H2O2participates in the pathway have long been in question. In the first chapter, transgenic plants HETAt and SHETAt were generated. In this chapter, HpalXoo and SHpalXoo Expression Transgenic Arabidopsis thaliana (HETAt and SHETAt) displayed growth and growth related genes enhancement. In SHETAt, H2O2accumulated in dependence of the outer membrane-associated NADPH oxidase activity, which didn't impact growth. In HETAt, the H2O2content elevated but was not related to NADPH oxidase, which also didn't contribute to growth. When Hpalxoo was applied to the parent plant, plant growth was enhanced, and H2O2of apoplasts and cytoplasts was not related to growth. These results suggested that growth pathways were activated in HETAt and SHETAt, and the pathway was not related to HpalXoo-induced H2O2.
3. Analysis of the interaction between Hpalxoo and PIP1;4in Arabidopsis membrane
The first two chapters have shown that Hpalxoo located in the extracellular and intracellular region, induced plant resistance to pathogenic bacteria Pst DC3000and Pcc RL4, enhanced plant growth and induced expression of defense and growth related genes in the transgenic plants. In this chapter, to resolve the acing mechanism of HpalXoo based on these valuable phenotypes of Hpalxoo, the Arabidopsis cDNA libraries were screened using HpalXoo as a bait and six interacting proteins were found including plant water channel protein PIP1;4. HpalXoo, but not ANT (deletion mutant, removed HpalXoo N-terminal53amino acids), could interact with PIP1;4by yeast two-hybrid assay, membrane yeast two-hybrid system, pull-down assay, bimolecular fluorescence complementation and fluorescence dys observation. And the interaction obviously localized at the cell membrane.
4. The preliminary study of the interaction between Hpalxoo and PIP1;4wich inducing growth enhancement and pathogen defenses in Arabidopsis
After confirming the interaction between HpalXoo and PIP1;4, eight kinds of atpip1;4Arabidopsis (CS803583, CS870828, CS872202, CS876999, CS879846, CS879691, CS870571and SALK_147568) seeds were purchased from the Arabidopsis Information Resource to study the impact of the interaction on plants and find the factors of plant affected. These mutants were T2-generation hybrids, and homozygotes were screened out from each mutant under the conditions of kanamycin and basta resistance. The two proteins Hpalxoo and ANT were successfully expressed and purified by prokaryotic expression system and protein purification kit. Hpalxoo-treated WT plants grew best compared with Hpalxoo-treated atpipl;4Arabidopsis,△NT or H2O-treated WT, and ANT or H2O-treated atpipl;4Arabidopsis. Hpalxoo-treated WT plants were resistant to Pst DC3000, but Hpalxoo-treated atpipl;4Arabidopsis, ANT or H2O-treated WT, and ANT or H2O-treated atpipl;4Arabidopsis were susceptible to Pst DC3000. All these tests suggested the interaction between Hpalxoo and PIP1;4could enhance plant growth and induce plant resistance to plant pathogens.
本研究使用HpalXoo,它由水稻白叶枯病原细菌(Xanthomonas oryzae pv. oryzae)产生,分子量15.6kD,比其他病原细菌的harpin(最大44kD)都小,但生物活性强。本研究着重剖析HpalXoo在植物中的亚细胞定位与后续反应之间的关系,质外体的H_2O_2如何参与植物防卫反应,HpalXoo蛋白在信号通路中如何被植物中的蛋白识别以及识别之后对植物产生的影响。
1.转基因植物产生的HpalXoo蛋白在质外体定位诱导H2O2产生与向细胞质转移及对抗病性的影响
已有的研究证明,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,包括产生于质外体的H2O2信号的转导,但是,一种harpin蛋白在一个信号通路中如何被识别以及质外体的H2O2如何参与防卫反应,还不清楚。本章中,我们研究了来源于水稻白叶枯病原细菌(Xanthomonas oryzae pv. oryzae)的harpin蛋白HpalXoo在拟南芥中是否影响了H2O2的产生以及植物的抗病性。把HpalXoo和融合了质外体定位信号基因(S)的HpalXoo (SHpalxoo)转入拟南芥,分别产生了转基因拟南芥植株HETAt和SHETAt。本研究发现,转基因拟南芥抗丁香假单胞番茄致病变种菌株DC3000(Pst DC3000)和大白菜软腐病菌菌株RL4(Pcc RL4)。HpalXoo定位于HETAt的细胞质和SHETAt的质外体和细胞膜,H2O2积累于HETAt的细胞质和SHETA的质外体和细胞质。已经知道,质外体中的H2O2是由质膜上的烟酰胺腺嘌呤二核苷酸磷酸氧化酶(NOX)产生的,这是植物防卫反应的一个普遍特征。H2O2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。当HpalXoo处理WT植物时,HpalXoo定位于质外体并在质外体中诱导H2O2的产生,但是质外体和细胞质中都有H2O2的积累。在SHETAt和HpalXoo处理的野生型(WT)拟南芥中,如果抑制质外体中H2O2的产生,细胞质中就没有H2O2的积累并且植物失去抗病能力。这些结果说明,HpalXoo诱导的产生于质外体的H202在参与植物调节抗病过程中需要从质外体运转到细胞质中。
2. HpalXoo促进转基因拟南芥生长的作用不依赖于H2O2
已有的研究证明,harpin能激活植物的生长信号通路。但是,一种harpin蛋白在一个信号通路中如何被识别以及harpin蛋白诱导的H2O2是否参与生长信号通路,还不清楚。在第一章中,我们产生了转基因拟南芥植株HETAt和SHETAt。本研究发现,转基因拟南芥植株HETAt和SHETAt的生长快于对照植株,并且植株体内的与生长相关基因表达增强。在SHETAt中,H2O2的积累依赖于质膜NADPH氧化酶活性,并且H2O2的积累不影响植株的生长。在HETAt中,H2O2含量的上升与NADPH氧化酶无关,H2O2的产生也不影响植株的生长。外施的HpalXoo能促进野生型拟南芥(WT)的生长,HpalXoo诱导的产生于质外体的H2O2以及积累于质外体和细胞内的H202与促生长无关。这些实验结果说明,HpalXoo能够激活植物的生长信号通路,并且这个信号通路与HpalXoo诱导的H2O2无关。
3. Hpalxoo与PIP1；4蛋白在拟南芥细胞膜上的互作验证
前两章已经证明,转入拟南芥的HpalXoo定位于植物的细胞外和细胞内,诱导植物的防卫反应,并能促进植物生长。基于HpalXoo的这些有益表型,本章中,为了试图解析HpalXoo的作用机理,以HpalXoo为诱饵对拟南芥cDNA文库进行筛选,初步筛选到6个阳性克隆,其中包括植物水通道蛋白PIP1；4。通过普通的酵母双杂交体系(Yeast two-hybrid system, Y2H)、膜酵母双杂交体系(Membrane yeast two-hybrid system, MYTH)、Pull-down assay、双分子荧光互补技术(Bimolecular Fluorescence Complementation, BiFC)和荧光染料的观察证实了HpalXoo与PIP1；4的互作,并且互作发生在植物的细胞膜上。而N端缺失53个氨基酸的HpalXoo蛋白(ANT)则不能与PIP1；4发生相互作用。
4. HpalXoo与PIP1；4蛋白互作促进拟南芥的生长与诱导其抗病的初步研究
证实HpalXoo与PIP1；4互作之后,为了研究这两个蛋白的互作对植物产生的影响,从拟南芥研究中心购买了8种atpipl;4拟南芥(CS803583、CS870828、CS872202、 CS876999、CS879846、CS879691、CS870571和SALK_147568)种子。这些突变体都是T2代杂合体,通过卡那霉素和Basta抗性筛选,从每一种突变体中都筛选到了纯合体。利用原核表达体系表达了HpalXoo和ΔNT并用蛋白纯化试剂盒纯化了这两个蛋白。通过表型观察发现,HpalXoo处理的WT拟南芥长势最好,而HpalXoo处理的atpip1;4拟南芥与ANT或水处理的WT和atpip1;4拟南芥生长表型一致,长势都弱于HpalXoo处理的WT。Pst DC3000接种实验发现,HpalXoo处理的WT拟南芥抗病最强,而HpalXoo处理的atpip1;4拟南芥与ΔNT或水处理的WT和atpip1;4拟南芥易感病。这些试验结果说明,PIP1；4与HpalXoo的互作不仅能够促进拟南芥的生长,也能够诱导拟南芥对病原菌的抗性。
Harpin proteins produced by gram-negative phytopathogenic bacteria generally affect virulence to host plants and the hypersensitive reaction (HR) of nonhost plants. When applied to plants, any harpins can induce resistance to pathogens and insects, confer drought tolerance, and enhance plant growth. The diverse effects are attributable to the activation of distinct signaling pathways that involve transduction of phytohormones, reactive oxygen species (ROS) signals, ion channels, programmed cell death, systemic acquired resistance (SAR) or salicylic acid (SA) signal pathway and protein kinase cascade, etc. Currently, harpins have been identified from almost all the Gram-negative phytopathogenic bacteria. Harpin have been shown to activate plant growth and defense signaling pathways, but the recognition of a harpin connecting with a pathway in plants, what a plant receptor of a particular harpin protein is, the transduction of the signal to a cellular pathway and what plant translocator (s) for plant-cytoplasmic trafficking of bacterial effector proteins is (are), especially type-Ⅲ effectors have long been in question.
The Hpalxoo protein produced by the rice bacterial leaf blight pathogen Xanthomonas oryzae pv. oryzae belongs to the harpin group of proteins. This study focuses on analysis of the relationship between cellular location of Hpalxoo and subsequent responses in plants, the apoplastic H2O2participation in defenses, the Hpalxoo recognition by a plant receptor and the impact of perception.
1. Apoplastic location of harpin protein HpalXoo induces apoplastic generation and cytoplasmic translocation of H2O2required for pathogen resistance in Arabidopsis
Harpin proteins secreted by Gram-negative phytopathogenic bacteria have been shown to activate plant defense pathways that involve transduction of hydrogen peroxide (H2O2) signal generated in the apoplast, but how a harpin is recognized connecting with a pathway and how the apoplastic H2O2anticipates in defenses are to be elucidated. Here we study if the cellular location of Hpalxoo, a harpin protein produced by rice bacterial leaf blight pathogen Xanthomonas oryzae pv. oryzae, impacts H2O2production and pathogen resistance in Arabidopsis thaliana. Transformation with the Hpalxoo gene and Hpalxoo fused to an apoplastic localization signal (SHpal Xoo) generated HpalXoo-and SHpal Xoo-expressing transgenic A. thaliana (HETAt and SHETAt) plants, respectively. HpalXoo was found to associate with the cytoplasm in HETAt and the apoplast in SHETAt, accompanying H2O2accumulation in cytoplasts and in apoplasts of SHETAt4as well. Apoplastic H2O2production via nicotinamide adenine dinucleotide phosphate (NADPH) oxidase (NOX) located in the plasma membrane has been shown as a common feature of plant defenses. In SHETAt, H2O2was generated in apoplasts in a NOX-dependent manner but accumulated to a greater extent in apoplasts than in cytoplasts. When applied to the parent plant, Hpalxoo located and induced H2O2generation in apoplasts but caused H2O2accumulation in both apoplasts and cytoplasts. In SHETAt and HpalXoo-treated parent plants, however, inhibition of apoplastic H2O2generation abrogated cytoplasmic H2O2accumulation and plant resistance to bacterial pathogens. These results suggest that the apoplastic H2O2is subject to a cytoplasmic translocation for participation in the regulation of pathogen resistance in the plant.
2. Growth enhancement of Hpalxoo transgenic Arabidopsis is not related to Hpalxoo-induced H2O2
The Hpalxoo protein produced by the rice bacterial leaf blight pathogen Xanthomonas oryzae pv. oryzae belongs to the harpin group of proteins secreted by Gram-negative phytopathogenic bacteria. In molecule mass, Hpal is smaller than other harpins, only15.6KD. harpin have been shown to activate plant growth signaling pathways, but how a harpin is recognized connecting with the pathway and how the harpin-induced H2O2participates in the pathway have long been in question. In the first chapter, transgenic plants HETAt and SHETAt were generated. In this chapter, HpalXoo and SHpalXoo Expression Transgenic Arabidopsis thaliana (HETAt and SHETAt) displayed growth and growth related genes enhancement. In SHETAt, H2O2accumulated in dependence of the outer membrane-associated NADPH oxidase activity, which didn't impact growth. In HETAt, the H2O2content elevated but was not related to NADPH oxidase, which also didn't contribute to growth. When Hpalxoo was applied to the parent plant, plant growth was enhanced, and H2O2of apoplasts and cytoplasts was not related to growth. These results suggested that growth pathways were activated in HETAt and SHETAt, and the pathway was not related to HpalXoo-induced H2O2.
3. Analysis of the interaction between Hpalxoo and PIP1;4in Arabidopsis membrane
The first two chapters have shown that Hpalxoo located in the extracellular and intracellular region, induced plant resistance to pathogenic bacteria Pst DC3000and Pcc RL4, enhanced plant growth and induced expression of defense and growth related genes in the transgenic plants. In this chapter, to resolve the acing mechanism of HpalXoo based on these valuable phenotypes of Hpalxoo, the Arabidopsis cDNA libraries were screened using HpalXoo as a bait and six interacting proteins were found including plant water channel protein PIP1;4. HpalXoo, but not ANT (deletion mutant, removed HpalXoo N-terminal53amino acids), could interact with PIP1;4by yeast two-hybrid assay, membrane yeast two-hybrid system, pull-down assay, bimolecular fluorescence complementation and fluorescence dys observation. And the interaction obviously localized at the cell membrane.
4. The preliminary study of the interaction between Hpalxoo and PIP1;4wich inducing growth enhancement and pathogen defenses in Arabidopsis
After confirming the interaction between HpalXoo and PIP1;4, eight kinds of atpip1;4Arabidopsis (CS803583, CS870828, CS872202, CS876999, CS879846, CS879691, CS870571and SALK_147568) seeds were purchased from the Arabidopsis Information Resource to study the impact of the interaction on plants and find the factors of plant affected. These mutants were T2-generation hybrids, and homozygotes were screened out from each mutant under the conditions of kanamycin and basta resistance. The two proteins Hpalxoo and ANT were successfully expressed and purified by prokaryotic expression system and protein purification kit. Hpalxoo-treated WT plants grew best compared with Hpalxoo-treated atpipl;4Arabidopsis,△NT or H2O-treated WT, and ANT or H2O-treated atpipl;4Arabidopsis. Hpalxoo-treated WT plants were resistant to Pst DC3000, but Hpalxoo-treated atpipl;4Arabidopsis, ANT or H2O-treated WT, and ANT or H2O-treated atpipl;4Arabidopsis were susceptible to Pst DC3000. All these tests suggested the interaction between Hpalxoo and PIP1;4could enhance plant growth and induce plant resistance to plant pathogens.
引文
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