条锈菌诱导的小麦β-1，3-葡聚糖酶基因的表达特征分析及条锈菌基因功能验证体系建立的探索
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摘要
小麦条锈病是由条形柄锈菌小麦专化型(Puccinia striiformis Westend. f. sp. tritici Eriks.,Pst)引起的世界范围内小麦生产上危害最为严重的一类病害,已成为影响小麦生产可持续性发展的限制因素。长期以来,国内外的锈病工作者围绕该病害开展了各项系统的研究,已在条锈病流行规律、综合防治等方面取得了重大成就。在寄主和病原菌互作方面,也已从组织学,细胞化学及生物化学等方面展开了一系列的讨论。但关于互作的分子机理以及病原真菌的基因功能方面尚未展开深入的研究。本论文则围绕小麦条锈菌及其与小麦互作的体系,主要开展了两大部分具体三方面的研究工作。
1.条锈菌诱导的小麦β-1,3-葡聚糖酶基因的表达特征分析
对条锈菌诱导的一种小麦β-1,3-葡聚糖酶基因TaGlu进行了基因组全长的克隆、生物信息学分析及在互作过程中的表达特征分析。TaGlu编码334个氨基酸组成的蛋白前体TaGLU,属于Glycoside hydrolase-family 17,具有Glycoside hydrolase catalytic core结构域,含有一个28个氨基酸构成的信号肽和一个跨膜区。进化树分析表明TaGLU和来源于小麦、大麦、水稻的碱性不具C末端延伸的葡聚糖酶亲缘关系较近。
在转录水平上,TaGlu受条锈菌的诱导表达,且亲和与非亲和组合中的表达变化趋势基本一致,只是在表达强度上,非亲和组合要高于亲和组合。此外,SA、MeJA和ET均能诱导TaGlu的快速积累,暗示TaGlu是通过多途径参与抗病防御反应。
以TaGlu的原核表达产物作为抗原,通过免疫实验家兔制备了抗血清,并纯化获得了该酶的特异性多克隆抗体Anti-TaGLU。ELISA分析表明制备的抗体具有较高的效价。Western blot结果显示,TaGLU在接种后24 h的亲和及非亲和组合中均有积累,但在未接种对照中没有积累或积累量很小检测不到,且没有和其他已报道不同分子量的小麦葡聚糖酶发生免疫学反应。TaGLU的免疫胶体金标记发现,在接种条锈菌后,该酶主要分布于寄主细胞壁和病原菌的吸器外间质中,且非亲和反应的标记密度要远高于亲和组合,表明TaGLU参与了小麦的抗条锈反应。
2.条锈菌无毒基因筛选体系建立的初探
前期借助生物信息学分析,从小麦条锈菌夏孢子、萌发夏孢子及吸器等cDNA文库中获得了大量候选无毒基因序列。但由于条锈菌是严格专性寄生菌,无法获得纯培养,缺乏行之有效的转化方法,因此无法用传统遗传学进一步明确这些基因的功能。本研究基于多数无毒基因产物在含相应抗病基因的寄主中产生HR的思路,利用小麦内生菌实现候选基因在植株体内的表达,通过产生HR表型变化,从而达到分离鉴定无毒基因的目的。
从小麦品种AVS、YR5、YR8、YR39拔节期叶片中共分离得到4种内生细菌WS、WR、YR和PR。通过对它们的基本生长特性研究,选择了具有三型分泌系统(TTSS)的Pseudomonas aeruginosa(WS)作为体系的介导细菌。构建了含有水稻病原菌Xanthomonas oryzae pv. oryzicola无毒基因avrRxo1的Gateway表达载体,转化至WS后渗透含有相应抗性基因Rxo1的玉米叶片,观察到了明显的过敏性坏死反应。说明该系统可以成功的在叶片中表达外源的无毒基因,并诱发相应的抗性植株产生HR。该研究为条锈菌Avr基因的鉴定及进一步利用无毒基因进行相应R基因的分离奠定基础。
3.根癌农杆菌介导的小麦条锈菌遗传转化体系构建的探索
小麦条锈菌是担子菌亚门一种严格专性寄生的病原真菌,无法在人工培养基上完成其生活史。建立一套适用于基因功能分析的稳定遗传转化体系对深入开展小麦条锈菌的功能基因组学研究显得尤为迫切。本研究首次选择根癌农杆菌介导的方法,进行了小麦条锈菌遗传转化的探索。
研究构建了由小麦杆锈菌(Puccinia graminis f. sp. tritici,Pgt)elongation factor 1α基因启动子Ef-1α控制的以潮霉素B为选择标记、红色荧光蛋白(RED)为报告基因的双元载体,建立了四种共培养方法,获得了一批检测呈阳性、外源标记基因可能成功插入条锈菌基因组的转化后代。明确了根癌农杆菌转化方法在小麦条锈菌遗传转化体系构建上的可行性,为稳定遗传转化体系的建立并通过其进行功能基因组学方面的研究提供方法和思路。
Stripe rust, caused by Puccinia striiformis Westend. f. sp. tritici Eriks. (Pst), is a serious fungal disease of wheat worldwide. Epidemics of the disease cause huge yield losses and downgrading in grain quality. Great achievements have been made in epidemiology and disease control. Systemic research on the mechanism between wheat-Pst has been extensively carried out at the histology, cytology, and biochemical aspects. However, little was known about the molecular interaction between the host and microbe as well as the gene function of the fungi. In this present thesis, two major parts including three aspects related to Puccinia striiformis and the wheat-Pst interaction system were represented.
1. Expression profile of a wheatβ-1,3-glucanase gene induced by Pst
To investigate the role ofβ-1,3-glucanase (EC 3.2.1.39) in the resistance response of wheat (cv. Suwon 11) to stripe rust, a wheatβ-1,3-glucanase gene induced by Pst, designated as TaGlu, was cloned and characterized. TaGlu was predicted to encode a basic protein of 334 amino acids. Quantitative real-time PCR analysis revealed that the transcript of TaGlu was induced during both compatible and incompatible interactions with Pst, but the transcription level was much higher in the incompatible interaction than that in the compatible interaction. TaGlu also showed noticeable induction of gene expression in young green leaf tissues treated with salicylic acid, methyl jasmonate or ethylene. Immunogold labeling assay showed that the gold particles were localized mainly in the host cell wall and over the extrahaustorial matrix, and the labeling densities were found significantly higher in the incompatible interaction than those in the compatible interaction.
2. Primary study of development of Pst avirulence gene screening system
A large quantity sequences of putative avirulence genes have been selected from different cDNA libraries of Pst based on their bioinformatics analysis. However, their functions were not identified due to the reason that gene function investigation system of Pst is unavailable. To overcome the limitations of existing methods to identify Pst avirulence genes, a strategy based on functional expression that exhibit HR-based resistance was repersented in this study.
Four gram-negative bacteria endophytes (WS, WR, YR and PR) were isolated from four wheat adult plants. WS, which identified as Pseudomonas aeruginosa, was then used as a“microorganism-vector”to delivery and express Pst genes in plant leaves based on its growth characteristics and the the type III secretion system. A gateway expression vector containing Xanthomonas oryzae pv. oryzicola avirulence gene avrRxo1 was constructed and transformed into WS. After infiltration of the endophyte into the leaves of the cron cultivar which have related resistant gene Rxo1, an obvious HR was observed. The result indicated this system was able to delivery and express foreign gene in plant leaves. However, considering the problem of stability, the system still needs improvements before screening Pst avirulence genes.
3. Primary study of Agrobacterium-mediated transformation of Puccinia striiformis
The biotrophic fungus, Pst, can not be cultured on the artificial medium and does not have known alternate host. It is difficult to use conventional genetic approaches to study the gene function. In order to facilitate the employment of molecular genetic techniques with Pst, the establishment of a transformation system for this fungus is required. Previous attempts of the Ps transformation were all based on particle bombardment.
Here, we introduced DNA into urediospores by using the Agrobacterium-mediated transformation system. The vector p13Ef-HYG-RED was builded up by inserting stem rust Ef-1αpromoter controled hph and Red genes fragment into pCAMBIA1302. 4 co-cultivation methods were established and got certain collections of putative transformants. These results could be very important for stable transformation system development of rust fungus.
1.条锈菌诱导的小麦β-1,3-葡聚糖酶基因的表达特征分析
对条锈菌诱导的一种小麦β-1,3-葡聚糖酶基因TaGlu进行了基因组全长的克隆、生物信息学分析及在互作过程中的表达特征分析。TaGlu编码334个氨基酸组成的蛋白前体TaGLU,属于Glycoside hydrolase-family 17,具有Glycoside hydrolase catalytic core结构域,含有一个28个氨基酸构成的信号肽和一个跨膜区。进化树分析表明TaGLU和来源于小麦、大麦、水稻的碱性不具C末端延伸的葡聚糖酶亲缘关系较近。
在转录水平上,TaGlu受条锈菌的诱导表达,且亲和与非亲和组合中的表达变化趋势基本一致,只是在表达强度上,非亲和组合要高于亲和组合。此外,SA、MeJA和ET均能诱导TaGlu的快速积累,暗示TaGlu是通过多途径参与抗病防御反应。
以TaGlu的原核表达产物作为抗原,通过免疫实验家兔制备了抗血清,并纯化获得了该酶的特异性多克隆抗体Anti-TaGLU。ELISA分析表明制备的抗体具有较高的效价。Western blot结果显示,TaGLU在接种后24 h的亲和及非亲和组合中均有积累,但在未接种对照中没有积累或积累量很小检测不到,且没有和其他已报道不同分子量的小麦葡聚糖酶发生免疫学反应。TaGLU的免疫胶体金标记发现,在接种条锈菌后,该酶主要分布于寄主细胞壁和病原菌的吸器外间质中,且非亲和反应的标记密度要远高于亲和组合,表明TaGLU参与了小麦的抗条锈反应。
2.条锈菌无毒基因筛选体系建立的初探
前期借助生物信息学分析,从小麦条锈菌夏孢子、萌发夏孢子及吸器等cDNA文库中获得了大量候选无毒基因序列。但由于条锈菌是严格专性寄生菌,无法获得纯培养,缺乏行之有效的转化方法,因此无法用传统遗传学进一步明确这些基因的功能。本研究基于多数无毒基因产物在含相应抗病基因的寄主中产生HR的思路,利用小麦内生菌实现候选基因在植株体内的表达,通过产生HR表型变化,从而达到分离鉴定无毒基因的目的。
从小麦品种AVS、YR5、YR8、YR39拔节期叶片中共分离得到4种内生细菌WS、WR、YR和PR。通过对它们的基本生长特性研究,选择了具有三型分泌系统(TTSS)的Pseudomonas aeruginosa(WS)作为体系的介导细菌。构建了含有水稻病原菌Xanthomonas oryzae pv. oryzicola无毒基因avrRxo1的Gateway表达载体,转化至WS后渗透含有相应抗性基因Rxo1的玉米叶片,观察到了明显的过敏性坏死反应。说明该系统可以成功的在叶片中表达外源的无毒基因,并诱发相应的抗性植株产生HR。该研究为条锈菌Avr基因的鉴定及进一步利用无毒基因进行相应R基因的分离奠定基础。
3.根癌农杆菌介导的小麦条锈菌遗传转化体系构建的探索
小麦条锈菌是担子菌亚门一种严格专性寄生的病原真菌,无法在人工培养基上完成其生活史。建立一套适用于基因功能分析的稳定遗传转化体系对深入开展小麦条锈菌的功能基因组学研究显得尤为迫切。本研究首次选择根癌农杆菌介导的方法,进行了小麦条锈菌遗传转化的探索。
研究构建了由小麦杆锈菌(Puccinia graminis f. sp. tritici,Pgt)elongation factor 1α基因启动子Ef-1α控制的以潮霉素B为选择标记、红色荧光蛋白(RED)为报告基因的双元载体,建立了四种共培养方法,获得了一批检测呈阳性、外源标记基因可能成功插入条锈菌基因组的转化后代。明确了根癌农杆菌转化方法在小麦条锈菌遗传转化体系构建上的可行性,为稳定遗传转化体系的建立并通过其进行功能基因组学方面的研究提供方法和思路。
Stripe rust, caused by Puccinia striiformis Westend. f. sp. tritici Eriks. (Pst), is a serious fungal disease of wheat worldwide. Epidemics of the disease cause huge yield losses and downgrading in grain quality. Great achievements have been made in epidemiology and disease control. Systemic research on the mechanism between wheat-Pst has been extensively carried out at the histology, cytology, and biochemical aspects. However, little was known about the molecular interaction between the host and microbe as well as the gene function of the fungi. In this present thesis, two major parts including three aspects related to Puccinia striiformis and the wheat-Pst interaction system were represented.
1. Expression profile of a wheatβ-1,3-glucanase gene induced by Pst
To investigate the role ofβ-1,3-glucanase (EC 3.2.1.39) in the resistance response of wheat (cv. Suwon 11) to stripe rust, a wheatβ-1,3-glucanase gene induced by Pst, designated as TaGlu, was cloned and characterized. TaGlu was predicted to encode a basic protein of 334 amino acids. Quantitative real-time PCR analysis revealed that the transcript of TaGlu was induced during both compatible and incompatible interactions with Pst, but the transcription level was much higher in the incompatible interaction than that in the compatible interaction. TaGlu also showed noticeable induction of gene expression in young green leaf tissues treated with salicylic acid, methyl jasmonate or ethylene. Immunogold labeling assay showed that the gold particles were localized mainly in the host cell wall and over the extrahaustorial matrix, and the labeling densities were found significantly higher in the incompatible interaction than those in the compatible interaction.
2. Primary study of development of Pst avirulence gene screening system
A large quantity sequences of putative avirulence genes have been selected from different cDNA libraries of Pst based on their bioinformatics analysis. However, their functions were not identified due to the reason that gene function investigation system of Pst is unavailable. To overcome the limitations of existing methods to identify Pst avirulence genes, a strategy based on functional expression that exhibit HR-based resistance was repersented in this study.
Four gram-negative bacteria endophytes (WS, WR, YR and PR) were isolated from four wheat adult plants. WS, which identified as Pseudomonas aeruginosa, was then used as a“microorganism-vector”to delivery and express Pst genes in plant leaves based on its growth characteristics and the the type III secretion system. A gateway expression vector containing Xanthomonas oryzae pv. oryzicola avirulence gene avrRxo1 was constructed and transformed into WS. After infiltration of the endophyte into the leaves of the cron cultivar which have related resistant gene Rxo1, an obvious HR was observed. The result indicated this system was able to delivery and express foreign gene in plant leaves. However, considering the problem of stability, the system still needs improvements before screening Pst avirulence genes.
3. Primary study of Agrobacterium-mediated transformation of Puccinia striiformis
The biotrophic fungus, Pst, can not be cultured on the artificial medium and does not have known alternate host. It is difficult to use conventional genetic approaches to study the gene function. In order to facilitate the employment of molecular genetic techniques with Pst, the establishment of a transformation system for this fungus is required. Previous attempts of the Ps transformation were all based on particle bombardment.
Here, we introduced DNA into urediospores by using the Agrobacterium-mediated transformation system. The vector p13Ef-HYG-RED was builded up by inserting stem rust Ef-1αpromoter controled hph and Red genes fragment into pCAMBIA1302. 4 co-cultivation methods were established and got certain collections of putative transformants. These results could be very important for stable transformation system development of rust fungus.
引文
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