芳氧苯氧丙酸类和氯乙酰胺类除草剂降解菌的分离鉴定、降解基因克隆表达及其代谢途径研究
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摘要
除草剂大量使用造成的环境污染问题日益受到人们的关注,解决污染的手段与方法不断地被提出和更新,其中利用微生物降解除草剂已成为近年来的研究热点。芳氧苯氧丙酸类和氯乙酰胺类除草剂是我国除草甘膦以外,使用最多的两大类除草剂,并且这两类除草剂的品种多,使用量大；在解决了农田杂草危害的同时,也给环境带来了巨大的污染。本研究以精嗯唑禾草灵和乙草胺为底物,对芳氧苯氧丙酸类和氯乙酰胺类除草剂降解微生物进行了分离鉴定,以期为这两大类除草剂污染的生物修复及其抗除草剂转基因作物的研究提供理论基础和微生物资源。
一、精噁唑禾草灵降解菌的分离鉴定及降解特性研究
从长期受农药污染的土壤中分离出能够快速降解精嗯唑禾草灵的菌株T1,根据表型特征、生理生化特性及16S rDNA系统发育分析,将其初步鉴定为红球菌属(Rhodococcus sp.)。菌株T1最适生长温度和pH分别25℃和7.0,好氧性生长,当NaCl浓度小于3%时,菌体生长良好。
经HPLC/MS鉴定菌株T1代谢精嗯唑禾草灵的产物为精噁唑禾草灵酸,即该菌株降解精噁唑禾草灵是依靠断裂其酯键来实现的,但是它不能进一步降解精嗯唑禾草灵酸。菌株T1可以精嗯唑禾草灵降解过程中产生的乙醇进行生长,其降解精噁唑禾草灵的最适温度和pH分别为30℃和8.0。在该条件下,5%接种量的菌株T1在24h内可将100mg·L-1的精噁唑禾草灵降解94%以上,且该菌株还能降解包括精喹禾灵、氰氟草酯、炔草酯、高效氟吡甲禾灵等在内的多种芳氧苯氧丙酸类除草剂,降解谱很广。
二、精嗯唑禾草灵水解基因feh的克隆表达及酶学性质研究
以三丁酸甘油酯为底物,利用鸟枪法从Rhobococcus sp. T1基因组文库中筛选到了两个含有酯酶基因的阳性克隆；其中阳性克隆pTT2具有精噁唑禾草灵水解活性。利用重叠延伸PCR技术从阳性克隆pTT2中成功扩增出了精噁唑禾草灵水解酶基因feh,并成功构建了该基因的表达载体pET29a-feh; feh可以在E.coli BL21(DE3)中很好地表达,重组酶Feh经Ni-NTA柱纯化后获得纯酶。
精嗯唑禾草灵水解酶Feh的最适反应温度为50℃,最适反应pH值为9.5,具有较好的pH稳定性和热稳定性。Feh可以水解对硝基苯酯类和三酰甘油酯类物质,但水解能力受碳链长度影响。Feh以精噁唑禾草灵为底物时的Km和Vmax分别为0.37mmol·L-1和1.08mmol-min-1·mg-1.
三、乙草胺降解菌的分离鉴定
利用以乙草胺为唯一碳源的无机盐培养基,从乙草胺农药厂的活性污泥中获得了能够将乙草胺完全降解的富集液T3和T4,二者均能在6d内将100mg·L-1的乙草胺完全降解。富集液T3和T4均能降解丁草胺,并使敌稗部分降解产生新的产物,但二者均不能降解吡草胺和丙草胺。利用RFLP和平板培养发现富集液T3和T4中具有丰富的细菌多样性。
从富集液T3和T4中逐步分离到了能够将乙草胺完全降解的菌株T3-1、T3-6、T4-7和MEA3-1；它们联合降解乙草胺的途径为：菌株T3-1将乙草胺降解为2’-甲基-6’-乙基-2-氯乙酰苯胺(CMEPA),菌株T3-6和菌株T4-7再将CMEPA降解为2,6-甲乙基苯胺(MEA),而菌株MEA3-1则可将MEA完全降解。利用生理生化特性和16S rDNA序列分析将菌株T3-1、T3-6和MEA3-1分别鉴定为Rhodococcus sp.、Delftia sp和Sphingobium sp.。
四、乙草胺降解菌的降解特性研究及其对乙草胺的联合降解
菌株T3-1可以乙草胺降解产物为碳源生长,其降解乙草胺的最适温度和pH分别为37℃和7.0。在该条件下,5%接种量的T3-1在14h对200mg·L-1乙草胺的降解率为95.5%。菌株T3-1还可以降解丁草胺,但不能降解丙草胺、异丙草胺和吡草胺。菌株T3-6不能以CMEPA降解产物MEA或氯乙酸为唯一碳源生长,其降解CMEAP的最适温度和pH分别为30℃和7.0；在该条件下,5%接种量的T3-6在10h内即可将500mg·L-1的CMEPA完全转化为MEA和氯乙酸。菌株T3-6还可以降解苯胺和邻苯二酚,并使对苯二酚发生部分转化。菌株MEA3-1可以MEA为唯一碳源进行生长,使MEA完全矿化,其降解MEA的最适温度和pH分别为30℃和7.0。在该条件下,5%接种量的MEA3-1在10h内即可将50mg·L-1的MEA降解完。菌株MEA3-1不能降解苯胺,并且只能使邻苯二酚和对苯二酚发生部分转化。
在30℃和pH7.0条件下,菌株T3-1、T3-6和MEA3-1的共同作用可以在24h内将200mg·L-1的乙草胺完全矿化80%以上,其降解效果远远高于已报道的菌株。因此,由这三个菌株制成复合培养物,对乙草胺和丁草胺污染的生物修复有很大的应用价值。
The environmental pollution of extensive use of herbicides caused the people's growing concern. The means and methods to solve pollution continue to be made and updated, and microbial degradation of herbicides has become a research hotspot in recent years. Aryloxyphenoxypropionate herbicides and chloroacetanilide herbicides are two of the most used herbicides except glyphosate in China. The variety and extensive use of these two herbicides caused the serve environmental pollution as well as processed the weeds in the farmland. In this study, the fenoxaprop-ethyl and acetochlor were used as substrates and the aryloxyphenoxypropionate herbicides and chloroacetanilide herbicides degrading-microorganisms were isolated and identified in order to provide the theoretical basis and microbial resources for bioremendiation and research of herbicide resistant transgenic crops of these two type herbicides.
1. Isolation, identification and degradation characteristics of fenoxaprop-ethyl (FE)-degrading bacteria
An efficient FE-degrading strain T1was isolated from the long-term pesticide-contaminated soil and identified as Rhodococcus sp. based on morphology, physiological and biochemical characteristics as well as16S rDNA sequence analysis. The optimum growth temperature and pH of strain T1were25℃and7.0, aerobic growth and the cell growth is good when the NaCl concentration is less than3%.
The metabolite fenoxaprop acid (FA) was identified by HPLC/MS analysis and this strain converted FE by cleavage of the ester bond, but it could not further degrade FA. Strain T1could utilize ethanol for growth, which produced during the degradation of FE and the optimum temperature and pH were30℃and8.0for FE degradation, respectively. Under these conditions, strain T1could degrade94%of100mg-L"1FE within24h at5% inoculation and it could also efficiently degrade a variety of aryloxyphenoxy propionate herbicides including quizalofop-p-ethyl, cyhalofop-butyl, clodinafop-propargyl and haloxyfop-R-methyl, the degradation spectrum is very wide.
2. Cloning, expression and enzymatic properties of fenoxaprop-ethyl hydrolase gene feh
Two positive clones containing the esterase gene were screened from the genomic library of Rhobococcus sp. T1, which constructed by shotgun and used trbutyrin as the substrate, and clones pTT2with fenoxaprop-ethyl hydrolysis activity. FE hydrolase gene feh was successfully amplified from the positive clone pTT2by using overlap extension PCR, and the gene expression vector pET29a-feh was successfully constructed. The feh could express in E.coli BL21(DE3) largely and the pure recombinant enzyme Feh was obtained after purifying by Ni-NTA.
The optimum temperature and pH value of Feh is50℃and9.5, which with a good pH stability and thermal stability. Feh could also hydrolyze the p-nitrophenyl esters and the triglyceride substances, but the hydrolysis was affected by carbon chain length. Km and Vmax of Feh for fenoxaprop were0.37mmol·L-1and1.08mmol·min-1·mg-1, respectively.
3. Isolation and identification of acetochlor-degrading bacteria
The acetochlor-degrading enrichment culture T3and T4were obtained from the activated sludge of acetochlor plant by using acetochlor as sole carbon source in the mineral medium, which could completely degrade100mg-L"1acetochlor in six days. T3and T4could also degrade butachlor and transform propanil to new product, but they could not degrade metazachlor and pretilachlor. T3and T4were found with the rich diversity of bacteria by RFLP and plate culture medium.
Acetochlor-degrading bacteria strain T3-1, T3-6, T4-7and MEA3-1were progressively isolated from the enrichment culture T3and T4, which could degrade acetochlor completely by the cooperation among them. The metabolic pathway of acetochlor by these bacteria is strain T3-1transforming acetochlor to2'-methyl-6'-ethyl-2-chloroacetanilide (CMEPA), and then CMEPA would be degrade to2-methyl-6-ethyl aniline (MEA) by strain T3-6and T4-7, and at last MEA could be completely degraded by strain MEA3-1. Strain T3-1, T3-6and MEA3-1were identified as Rhodococcus sp., Delftia sp. and Sphingobium sp. based on morphology, physiological and biochemical characteristics as well as16S rDNA sequence analysis, respectively.
4. Degradation characteristics of acetochlor-degrading bacteria and the combined degradation of acetochlor
Strain T3-1could use the metabolites of acetochlor as sole carbon source for growth. The optimum temperature and pH were37℃and7.0for acetochlor degradation, respectively. Under these conditions, strain T3-1could degrade95.5%of200mg-L-1acetochlor within14h at5%inoculation. Strain T3-1could also efficiently degrade butachlor, but could not degrade pretilachlor, propisochlor and metazachlor. Strain T3-6could not use MEA or chloroacetic acid as sole carbon source for growth, which generated from CMEPA degradation and the optimum temperature and pH were30℃and7.0for CMEPA degradation, respectively. Under these conditions, strain T3-6could completely transform500mg-L"1CMEPA to MEA and chloroacetic acid within10h at5%inoculation. Strain T3-6could also degrade aniline and catechol, and convert hydroquinone partially. Strain MEA3-1could use MEA as sole carbon source for growth and minerilize MEA completely. The optimum temperature and pH were also30℃and7.0for MEA degradation, respectively. Under these conditions, strain MEA3-1could completely degrade50mg·L-1MEA within10h at5%inoculation. Strain MEA3-1could not degrade aniline, and catechol and hydroquinone could only partially transformed by it.
At temperature of30℃and pH7.0, more than80%of200mg-L-1acetochlor could be completely mineralize by the cooperation of strain T3-1, T3-6and MEA3-1within24h and the degradation rate is much higher than other reported strains. Therefore, the complex culture made by the three strains has great potential utility for the bioremediation of acetochlor and butachlor contaminated environment.
一、精噁唑禾草灵降解菌的分离鉴定及降解特性研究
从长期受农药污染的土壤中分离出能够快速降解精嗯唑禾草灵的菌株T1,根据表型特征、生理生化特性及16S rDNA系统发育分析,将其初步鉴定为红球菌属(Rhodococcus sp.)。菌株T1最适生长温度和pH分别25℃和7.0,好氧性生长,当NaCl浓度小于3%时,菌体生长良好。
经HPLC/MS鉴定菌株T1代谢精嗯唑禾草灵的产物为精噁唑禾草灵酸,即该菌株降解精噁唑禾草灵是依靠断裂其酯键来实现的,但是它不能进一步降解精嗯唑禾草灵酸。菌株T1可以精嗯唑禾草灵降解过程中产生的乙醇进行生长,其降解精噁唑禾草灵的最适温度和pH分别为30℃和8.0。在该条件下,5%接种量的菌株T1在24h内可将100mg·L-1的精噁唑禾草灵降解94%以上,且该菌株还能降解包括精喹禾灵、氰氟草酯、炔草酯、高效氟吡甲禾灵等在内的多种芳氧苯氧丙酸类除草剂,降解谱很广。
二、精嗯唑禾草灵水解基因feh的克隆表达及酶学性质研究
以三丁酸甘油酯为底物,利用鸟枪法从Rhobococcus sp. T1基因组文库中筛选到了两个含有酯酶基因的阳性克隆；其中阳性克隆pTT2具有精噁唑禾草灵水解活性。利用重叠延伸PCR技术从阳性克隆pTT2中成功扩增出了精噁唑禾草灵水解酶基因feh,并成功构建了该基因的表达载体pET29a-feh; feh可以在E.coli BL21(DE3)中很好地表达,重组酶Feh经Ni-NTA柱纯化后获得纯酶。
精嗯唑禾草灵水解酶Feh的最适反应温度为50℃,最适反应pH值为9.5,具有较好的pH稳定性和热稳定性。Feh可以水解对硝基苯酯类和三酰甘油酯类物质,但水解能力受碳链长度影响。Feh以精噁唑禾草灵为底物时的Km和Vmax分别为0.37mmol·L-1和1.08mmol-min-1·mg-1.
三、乙草胺降解菌的分离鉴定
利用以乙草胺为唯一碳源的无机盐培养基,从乙草胺农药厂的活性污泥中获得了能够将乙草胺完全降解的富集液T3和T4,二者均能在6d内将100mg·L-1的乙草胺完全降解。富集液T3和T4均能降解丁草胺,并使敌稗部分降解产生新的产物,但二者均不能降解吡草胺和丙草胺。利用RFLP和平板培养发现富集液T3和T4中具有丰富的细菌多样性。
从富集液T3和T4中逐步分离到了能够将乙草胺完全降解的菌株T3-1、T3-6、T4-7和MEA3-1；它们联合降解乙草胺的途径为：菌株T3-1将乙草胺降解为2’-甲基-6’-乙基-2-氯乙酰苯胺(CMEPA),菌株T3-6和菌株T4-7再将CMEPA降解为2,6-甲乙基苯胺(MEA),而菌株MEA3-1则可将MEA完全降解。利用生理生化特性和16S rDNA序列分析将菌株T3-1、T3-6和MEA3-1分别鉴定为Rhodococcus sp.、Delftia sp和Sphingobium sp.。
四、乙草胺降解菌的降解特性研究及其对乙草胺的联合降解
菌株T3-1可以乙草胺降解产物为碳源生长,其降解乙草胺的最适温度和pH分别为37℃和7.0。在该条件下,5%接种量的T3-1在14h对200mg·L-1乙草胺的降解率为95.5%。菌株T3-1还可以降解丁草胺,但不能降解丙草胺、异丙草胺和吡草胺。菌株T3-6不能以CMEPA降解产物MEA或氯乙酸为唯一碳源生长,其降解CMEAP的最适温度和pH分别为30℃和7.0；在该条件下,5%接种量的T3-6在10h内即可将500mg·L-1的CMEPA完全转化为MEA和氯乙酸。菌株T3-6还可以降解苯胺和邻苯二酚,并使对苯二酚发生部分转化。菌株MEA3-1可以MEA为唯一碳源进行生长,使MEA完全矿化,其降解MEA的最适温度和pH分别为30℃和7.0。在该条件下,5%接种量的MEA3-1在10h内即可将50mg·L-1的MEA降解完。菌株MEA3-1不能降解苯胺,并且只能使邻苯二酚和对苯二酚发生部分转化。
在30℃和pH7.0条件下,菌株T3-1、T3-6和MEA3-1的共同作用可以在24h内将200mg·L-1的乙草胺完全矿化80%以上,其降解效果远远高于已报道的菌株。因此,由这三个菌株制成复合培养物,对乙草胺和丁草胺污染的生物修复有很大的应用价值。
The environmental pollution of extensive use of herbicides caused the people's growing concern. The means and methods to solve pollution continue to be made and updated, and microbial degradation of herbicides has become a research hotspot in recent years. Aryloxyphenoxypropionate herbicides and chloroacetanilide herbicides are two of the most used herbicides except glyphosate in China. The variety and extensive use of these two herbicides caused the serve environmental pollution as well as processed the weeds in the farmland. In this study, the fenoxaprop-ethyl and acetochlor were used as substrates and the aryloxyphenoxypropionate herbicides and chloroacetanilide herbicides degrading-microorganisms were isolated and identified in order to provide the theoretical basis and microbial resources for bioremendiation and research of herbicide resistant transgenic crops of these two type herbicides.
1. Isolation, identification and degradation characteristics of fenoxaprop-ethyl (FE)-degrading bacteria
An efficient FE-degrading strain T1was isolated from the long-term pesticide-contaminated soil and identified as Rhodococcus sp. based on morphology, physiological and biochemical characteristics as well as16S rDNA sequence analysis. The optimum growth temperature and pH of strain T1were25℃and7.0, aerobic growth and the cell growth is good when the NaCl concentration is less than3%.
The metabolite fenoxaprop acid (FA) was identified by HPLC/MS analysis and this strain converted FE by cleavage of the ester bond, but it could not further degrade FA. Strain T1could utilize ethanol for growth, which produced during the degradation of FE and the optimum temperature and pH were30℃and8.0for FE degradation, respectively. Under these conditions, strain T1could degrade94%of100mg-L"1FE within24h at5% inoculation and it could also efficiently degrade a variety of aryloxyphenoxy propionate herbicides including quizalofop-p-ethyl, cyhalofop-butyl, clodinafop-propargyl and haloxyfop-R-methyl, the degradation spectrum is very wide.
2. Cloning, expression and enzymatic properties of fenoxaprop-ethyl hydrolase gene feh
Two positive clones containing the esterase gene were screened from the genomic library of Rhobococcus sp. T1, which constructed by shotgun and used trbutyrin as the substrate, and clones pTT2with fenoxaprop-ethyl hydrolysis activity. FE hydrolase gene feh was successfully amplified from the positive clone pTT2by using overlap extension PCR, and the gene expression vector pET29a-feh was successfully constructed. The feh could express in E.coli BL21(DE3) largely and the pure recombinant enzyme Feh was obtained after purifying by Ni-NTA.
The optimum temperature and pH value of Feh is50℃and9.5, which with a good pH stability and thermal stability. Feh could also hydrolyze the p-nitrophenyl esters and the triglyceride substances, but the hydrolysis was affected by carbon chain length. Km and Vmax of Feh for fenoxaprop were0.37mmol·L-1and1.08mmol·min-1·mg-1, respectively.
3. Isolation and identification of acetochlor-degrading bacteria
The acetochlor-degrading enrichment culture T3and T4were obtained from the activated sludge of acetochlor plant by using acetochlor as sole carbon source in the mineral medium, which could completely degrade100mg-L"1acetochlor in six days. T3and T4could also degrade butachlor and transform propanil to new product, but they could not degrade metazachlor and pretilachlor. T3and T4were found with the rich diversity of bacteria by RFLP and plate culture medium.
Acetochlor-degrading bacteria strain T3-1, T3-6, T4-7and MEA3-1were progressively isolated from the enrichment culture T3and T4, which could degrade acetochlor completely by the cooperation among them. The metabolic pathway of acetochlor by these bacteria is strain T3-1transforming acetochlor to2'-methyl-6'-ethyl-2-chloroacetanilide (CMEPA), and then CMEPA would be degrade to2-methyl-6-ethyl aniline (MEA) by strain T3-6and T4-7, and at last MEA could be completely degraded by strain MEA3-1. Strain T3-1, T3-6and MEA3-1were identified as Rhodococcus sp., Delftia sp. and Sphingobium sp. based on morphology, physiological and biochemical characteristics as well as16S rDNA sequence analysis, respectively.
4. Degradation characteristics of acetochlor-degrading bacteria and the combined degradation of acetochlor
Strain T3-1could use the metabolites of acetochlor as sole carbon source for growth. The optimum temperature and pH were37℃and7.0for acetochlor degradation, respectively. Under these conditions, strain T3-1could degrade95.5%of200mg-L-1acetochlor within14h at5%inoculation. Strain T3-1could also efficiently degrade butachlor, but could not degrade pretilachlor, propisochlor and metazachlor. Strain T3-6could not use MEA or chloroacetic acid as sole carbon source for growth, which generated from CMEPA degradation and the optimum temperature and pH were30℃and7.0for CMEPA degradation, respectively. Under these conditions, strain T3-6could completely transform500mg-L"1CMEPA to MEA and chloroacetic acid within10h at5%inoculation. Strain T3-6could also degrade aniline and catechol, and convert hydroquinone partially. Strain MEA3-1could use MEA as sole carbon source for growth and minerilize MEA completely. The optimum temperature and pH were also30℃and7.0for MEA degradation, respectively. Under these conditions, strain MEA3-1could completely degrade50mg·L-1MEA within10h at5%inoculation. Strain MEA3-1could not degrade aniline, and catechol and hydroquinone could only partially transformed by it.
At temperature of30℃and pH7.0, more than80%of200mg-L-1acetochlor could be completely mineralize by the cooperation of strain T3-1, T3-6and MEA3-1within24h and the degradation rate is much higher than other reported strains. Therefore, the complex culture made by the three strains has great potential utility for the bioremediation of acetochlor and butachlor contaminated environment.
引文
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