绿针假单胞菌GP72中2-羟基吩嗪（2-OH-PHZ）生物合成分子机理研究
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摘要
绿针假单胞菌GP72(Pseudomonas chlororaphis GP72),是从上海郊区甜椒根际土中分离筛选到的一株具有生物防治功能的假单胞菌属菌株,其分泌的吩嗪-1-羧酸(phenazine-1-carboxilic acid, PCA)和2-羟基吩嗪(2-hydroxyphenazine,2-OH-PHZ)等抗生素对植物病原真菌等具有有效抑制作用。本课题主要针对GP72中PCA的次级代谢产物2-OH-PHZ的合成机理,首次研究了它的生物合成途径中关键基因phzO的功能。PhzO蛋白酶是一种大小约55KDa的单脱氧酶,能够将PCA转化为2-OH-PHZ。同时,本课题还研究了phzE基因和PCA浓度对2-OH-PHZ生物合成的影响。
首先,运用PCR技术从GP72菌株中克隆phzO基因,并通过重组和转化技术构建了该基因的过表达菌株。与野生菌株相比,phzO过表达菌株(GP72-pME6032-phzO)和对照组菌株(GP72-pME6032)生长稍次于野生型菌株。利用HPLC技术进一步监测phzO基因过表达对PCA和2-OH-PHZ合成的影响,发现phzO基因在GP72菌株中过表达未能显著提高2-OH-PHZ产量,而PCA产量较野生型菌株显著降低。论文进一步通过在DH5α中异源表达phzO,加入外源PCA分子作为反应底物,借助HPLC对反应产物进行检测,发现PhzO蛋白酶能够将底物PCA分子转化为2-OH-PHZ产物,从而证实了phzO的功能。
其次,将克隆成功的phzO基因重组到假单胞菌M18菌株中,通过发酵实验验证M18菌株中phzE基因对2-OH-PHZ生物合成的影响。HPLC结果显示phzE基因对2-OH-PHZ的生物合成影响甚微。同时,实验证明了phzO基因跟PCA浓度对2-OH-PHZ的生物合成具有协同作用。
本课题结果初步断定PCA转化为2-OH-PHZ的途径为一步反应,可由phzO单个基因完成。2-OH-PHZ的产量受前体PCA浓度及PhzO蛋白酶量协同影响。
Pseudomonas chlororaphis GP72, isolated from the rhizosphere of sweet melon, is an effective root-colonizing bio-control agent against soil-borne pathogenic fungi on crop plants. This capability is primarily depending on its ability to produce two different types of antibiotics, phenazine-1-carboxylic acid (PCA) and 2-hydroxyphenazine (2-OH-PHZ). The phenazine biosynthetic genes in strain GP72 differ from those in other known pseudomonads by the presence of phzO, encoding a 55-kDa aromatic monooxygenase responsible for the hydroxylation of PCA to produce 2-OH- PCA. In this study, we focused on elucidating the reaction mechanism of how PCA converted to 2-OH-PHZ in strain GP72. Meanwhile, we have also investigated the effect of phzE and PCA concentration on the 2-OH-PHZ production.
Firstly, phzO gene was cloned from GP72 by PCR and phzO-over-expression strain was constructed by transformation. The phzO-over-expression strain (GP72-pME6032-phzO) and the controlled strain (GP72-pME6032) displayed worse cell growth as compared to the wild-type strain GP72. Thus, we continued to examine its influence on the PCA and 2-OH-PHZ production by HPLC. The results showed that over expression of phzO failed to remarkably increase 2-OH-PHZ amount. On the contratry, GP72-pME6032-phzO strain and GP72-pME6032 strain showed much lower PCA production than that of wild GP72 strain. In order to examine the function of converting PCA to 2-OH-PHZ of phzO, it was expressed in Escherichia coli DH5αand PCA was added to the culture. Results showed that expression of phzO in Escherichia coli DH5αcould convert free PCA to 2-OH-PHZ directly by HPLC analyses, though the conversion rate is relatively low.
Secondly, we transformed the successfully-cloned phzO gene into Pseudomonas sp. M18, a PCA-producing strain. Based on the HPLC analysis, the fermentation results showed that phzE gene had little influence on the production of 2-OH-PHZ in strain GP72. It was also turned out that the PCA concentration and the amount of PhzO protein both have effect on the output of 2-OH-PHZ.
Based on this study, we concluded that the conversion of PCA to 2-OH-PHZ was a one-step reaction, involved only one gene, phzO. Also, we found out that the output of 2-OH-PHZ was dermined both by PCA concentration and the amount of PhzO protein.
首先,运用PCR技术从GP72菌株中克隆phzO基因,并通过重组和转化技术构建了该基因的过表达菌株。与野生菌株相比,phzO过表达菌株(GP72-pME6032-phzO)和对照组菌株(GP72-pME6032)生长稍次于野生型菌株。利用HPLC技术进一步监测phzO基因过表达对PCA和2-OH-PHZ合成的影响,发现phzO基因在GP72菌株中过表达未能显著提高2-OH-PHZ产量,而PCA产量较野生型菌株显著降低。论文进一步通过在DH5α中异源表达phzO,加入外源PCA分子作为反应底物,借助HPLC对反应产物进行检测,发现PhzO蛋白酶能够将底物PCA分子转化为2-OH-PHZ产物,从而证实了phzO的功能。
其次,将克隆成功的phzO基因重组到假单胞菌M18菌株中,通过发酵实验验证M18菌株中phzE基因对2-OH-PHZ生物合成的影响。HPLC结果显示phzE基因对2-OH-PHZ的生物合成影响甚微。同时,实验证明了phzO基因跟PCA浓度对2-OH-PHZ的生物合成具有协同作用。
本课题结果初步断定PCA转化为2-OH-PHZ的途径为一步反应,可由phzO单个基因完成。2-OH-PHZ的产量受前体PCA浓度及PhzO蛋白酶量协同影响。
Pseudomonas chlororaphis GP72, isolated from the rhizosphere of sweet melon, is an effective root-colonizing bio-control agent against soil-borne pathogenic fungi on crop plants. This capability is primarily depending on its ability to produce two different types of antibiotics, phenazine-1-carboxylic acid (PCA) and 2-hydroxyphenazine (2-OH-PHZ). The phenazine biosynthetic genes in strain GP72 differ from those in other known pseudomonads by the presence of phzO, encoding a 55-kDa aromatic monooxygenase responsible for the hydroxylation of PCA to produce 2-OH- PCA. In this study, we focused on elucidating the reaction mechanism of how PCA converted to 2-OH-PHZ in strain GP72. Meanwhile, we have also investigated the effect of phzE and PCA concentration on the 2-OH-PHZ production.
Firstly, phzO gene was cloned from GP72 by PCR and phzO-over-expression strain was constructed by transformation. The phzO-over-expression strain (GP72-pME6032-phzO) and the controlled strain (GP72-pME6032) displayed worse cell growth as compared to the wild-type strain GP72. Thus, we continued to examine its influence on the PCA and 2-OH-PHZ production by HPLC. The results showed that over expression of phzO failed to remarkably increase 2-OH-PHZ amount. On the contratry, GP72-pME6032-phzO strain and GP72-pME6032 strain showed much lower PCA production than that of wild GP72 strain. In order to examine the function of converting PCA to 2-OH-PHZ of phzO, it was expressed in Escherichia coli DH5αand PCA was added to the culture. Results showed that expression of phzO in Escherichia coli DH5αcould convert free PCA to 2-OH-PHZ directly by HPLC analyses, though the conversion rate is relatively low.
Secondly, we transformed the successfully-cloned phzO gene into Pseudomonas sp. M18, a PCA-producing strain. Based on the HPLC analysis, the fermentation results showed that phzE gene had little influence on the production of 2-OH-PHZ in strain GP72. It was also turned out that the PCA concentration and the amount of PhzO protein both have effect on the output of 2-OH-PHZ.
Based on this study, we concluded that the conversion of PCA to 2-OH-PHZ was a one-step reaction, involved only one gene, phzO. Also, we found out that the output of 2-OH-PHZ was dermined both by PCA concentration and the amount of PhzO protein.
引文
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[2] Noel D. Uri. Development and Use of Biopesticides. Implications of Government Policy and Consumers’Preferences, Technological Forecasting and Social Change, 1998, (59): 291-304.
[3]张兴,马志卿,李广译.农药管理与科学. 2002,23(1): 34-36.
[4]顾宝根,姜辉.微生物农药及产业化.科学出版社,2000,13-18.
[5]张锡贞,张红雨.生物农药的应用与研发现状.山东理工大学学报, 2004,18(1): 96-100.
[6] Emilio M. Development registration and commercialization of microbial pesticides for plant protection. Int Microbiol, 2003, (6): 245-252.
[7] Mannion A. M. Agriculture, environment and biotechnology. Agriculture, Ecosystems and Environments, 1995, (53):31-45.
[8]陈忠孝,胡国文.印楝质杀虫剂的研究及前景.世界农业,1992,(7):30.
[9]成为宁,李修炼.植物性杀虫剂防止小麦吸浆虫研究.西北农业学报,2000,9(2):41-44.
[10]李小东,赵善欢.印楝对昆虫的毒理作用机制.华南农业大学学报,1996,17(1):118-122.
[11] Kotkar H.M., Mendki P.S., Sadan S.V., Jha S.R., Upasani S.M., Maheshwari V.L. Antimicrobial and pesticidal activity of partially purified flavonoids of Annona squamosa. Pest Manag Sci. 2002, 58(1):33-39.
[12] Karl J., Kramer, Subbaratnnam M., Molecular Biology and Potential Use as Biopesticides. Insect Biochem. Molec. Biol., 1997, 27(11): 887-900.
[13]吴文君,高希武.生物农药及其应用.化学工业出版社,2004.
[14] Leonard G., Copping, Julius J.M. Biopesticides. A review of their action, applications and efficacy. Pest Manag Sci, 2000, 56: 651-676.
[15] Campbell R., Greaves M. P. Anatomy and community structure of the rhizosphere. The Rhizosphere, 1990: 11-34.
[16] Haas D., Defago G. Biological control of soil-borne pathogens by fluorescent pseudomonads. Nat Rev Microbiol, 2005, 3: 307-319.
[17] Turner J. M., Messenger A. J. Occurrence, biochemistry and physiology of phenazine pigment production. Advances in microbial physiology, 1986, 27: 211-275.
[18] Mavrodi D. V., Ksenzenko V. N., Bonsall R. F., Cook R. J. A seven-gene locus for synthesis of phenazine-1-carboxylic acid by Pseudomonas fluorescens 2-79. J Bacteriol, 1998, 180: 2541-2548.
[19] Pierson Iii L. S., Keppenne V. D., Wood D. W. Phenazine antibiotic biosynthesis in Pseudomonas aureofaciens 30-84 is regulated by PhzR in response to cell density. Journal of Bacteriology, 1994, 176: 3966-3974.
[20] Pessi G., Williams F., Hindle Z., Heurlier K. The global posttranscriptional regulator RsmA modulates production of virulence determinants and N-acylhomoserine lactones in Pseudomonas aeruginosa. J Bacteriol, 2001, 183: 6676-6683.
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