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標題: 功能分析tzs基因及細胞分裂素對農桿菌生長、感染力及致病基因表現之影響
Functional studies of the tzs gene and cytokinin effects on Agrobacterium tumefaciens growth, virulence, and virulence gene expressions
作者: 鄭惇方
Cheng, Tun-Fang
關鍵字: 農桿菌
Agrobacterium tumefaciens
出版社: 生命科學系所
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摘要: 農桿菌(Agrobacterium tumefaciens)存在於土壤中,為一種植物性病原菌。農桿菌的Ti質體(tumor-inducing plasmid,Ti plasmid)上含有一段T-DNA (transfer DNA)會藉由感染過程,進而轉移並嵌入植物細胞染色體上。由於T-DNA上帶有生長激素(auxin)和細胞分裂素(cytokinin)的生合成基因(如:iaaH[indole-3-acetamide hydrolase]、iaaM[indoleacetic acid tryptophan monooxygenase]、ipt [isopentenyl transferase])。因此當T-DNA嵌入植物染色體後,這些基因被表現進而生合成植物賀爾蒙,導致植物細胞不正常增生而形成冠癭狀腫瘤(crown gall tumor)。農桿菌細胞膜上的VirA/VirG雙分子磷酸依賴系統(two-component phosphor-relay system),可以感受到植物受傷後分泌的醣類及酚類化合物,進而誘導開啟下游vir基因(virulence gene)的大量表現。這些vir基因產物會幫助T-DNA的產生、轉移並嵌入植物染色體中。在胭脂鹼型(nopaline type)農桿菌的Ti質體上,含有一段與ipt基因序列相似的tzs (trans-zeatin synthesis)基因,可以轉譯產生Tzs蛋白質,促使反玉米素生合成。tzs基因可以受到AS (acetosyringone)的誘導而大量表現,也會受到VirA/VirG雙分子磷酸依賴系統的調控而大量表現。Tzs蛋白質可與位在細胞膜上組成第四型分泌系統(Type IV secretion system,T4SS)的VirB5結合。又已知當tzs基因突變後,會導致農桿菌分泌反玉米素的含量減少、感染植物的能力下降及生長速度遲緩。在感染過程中,額外添加細胞分裂素,則能提升突變株感染植物的效率。並發現突變株培養在19oC、pH5.5的AB-MES培養基中,菌體內Vir蛋白質有較高的累積量,並有較高的virB啟動子活性。若在培養基中額外添加細胞分裂素,則突變株及野生株農桿菌中Vir蛋白質累積量,及virB啟動子活性皆有受到抑制。因此本研究為進一步探討tzs基因及其產物對農桿菌生長趨勢、感染植株能力及vir基因表現的影響。研究結果顯示,在19℃酸性AB-MES培養基中,tzs突變株的數個vir基因和tzs基因啟動子活性皆較野生株高,且在誘導培養的早期與野生株差異最明顯。另外,在AS誘導培養過程中額外添加細胞分裂素,可抑制數個vir基因和tzs基因啟動子活性。並發現當tzs基因功能喪失後,可能不會直接影響農桿菌第四型分泌系統的質體轉移效率,及附著在植物根上的能力。此外進一步構築四株tzs基因功能缺失突變株,並發現四株tzs突變株的表現性狀與tzs缺失株和tzs框移突變株相似,其感染力、生長速率皆低於野生株,而Vir蛋白質累積量則高於野生株。因此由本研究可知tzs基因及其產物可以藉由調控vir基因的表現,促使農桿菌改變生理代謝,進而影響農桿菌感染植物的能力。
Agrobacterium tumefaciens is a plant pathogen and exists in soil, which contains T-DNA (transfer DNA) in a Ti plasmid (tumor-inducing plasmid). The T-DNA are transferred and integrated into plant chromosome during A. tumefaciens infection process. There are auxin and cytokinin biosynthesis genes on the T-DNA, including iaaH (indole-3-acetamide hydrolase), iaaM (indoleacetic acid tryptophan monooxygenase), ipt (isopentenyl transferase). When the T-DNA integrates into plant chromosome, genes in the T-DNA region express and cause high levels of auxin and cytokinin productions, which increases plant cell numbers and sizes abnormally and finally causes crown gall disease. When plant wounded sites secrete monosaccharides and phenolic compounds, the VirA/VirG two component system in A. tumefaciens recognizes those signals, and induces virulence (vir) gene expressions. The Vir proteins involve in T-DNA production, transfer, and integration into plant chromosome. The tzs (trans-zeatin synthesis) gene which shares sequence similarity with the ipt gene only exists in the nopaline-type Ti plasmid. The Tzs protein mediates the biosynthesis of trans-zeatin in A. tumefaciens. The tzs gene expression is induced by acetosyringone (AS) and regulated by the VirA/VirG two component system. The tzs deletion and frame-shift mutants decreased trans-zeatin secretions, reduced virulence on several plant species and showed growth defects during infections. When exogenous cytokinins were added during infections, it restored virulence of the tzs mutants on plants. When grown in acidic AB-MES media with AS at 19oC, the Vir protein accumulations and virB promoter activities increased in the tzs mutants. When exogenous cytokinins were added during AS induction, the Vir protein accumulations and virB promoter activities were repressed. Results of thesis study showed that several vir genes promoter activities increased in the tzs mutants when grown in acidic AB-MES with AS at 19oC in comparison to wild-type, especially at the early stage of AS inductions. When exogenous cytokinins were added during AS inductions, several vir genes and tzs gene promoter activities decreased in the tzs mutant and the wild-type strains. In addition, RSF1010 plasmid transfer efficiencies by the type IV secretion system and bacterial attachment abilities on Arabidopsis roots were not significantly affected by the absence of the Tzs protein. We additionally generated four new tzs mutants, which phenotype are similar with tzs mutant. Four new tzs mutants reduced virulence on potato tuber discs, showed growth defects under AS inductions and increased Vir protein accumulations. Taken together, these data suggest that the tzs gene and it product, cytokinin, may be involved in regulation of vir gene expression, and therefore affect bacterial growth and virulence on plants.
其他識別: U0005-0102201311513200
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