诺氟沙星的土壤环境行为及生态效应研究
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摘要
抗生素被广泛的用于治疗、预防人类和动物的疾病,但是抗生素进入动物或人体体内,并不被完全吸收,而是以原药或代谢产物的形式排到体外,进入土壤、水体等环境中。而后通过一系列的迁移和转化过程,在环境中广泛分布,进而对环境产生一定的影响。同时抗生素还可以与环境中已有污染物共同作用形成复合污染,对生态环境造成影响。铜是一种重要的工业原料和常用的饲料添加剂,它通过有机肥和污泥农用最终分布在农田土壤中。而且,铜离子能够与抗生素类药物分子中的功能基团结合形成络合物。因此,研究抗生素的环境行为和效应及重金属铜对其的影响有重要的意义。本研究选取诺氟沙星(Norfloxacin, Nor)为代表性污染物,利用HPLC、Biolog、 PCR-DGGE等现代检测和分析技术,研究了抗生素在典型农田土壤环境中的吸附、降解行为,及药物单独和与重金属复合污染条件下,土壤酶活性、土壤微生物群落结构的变化。结合对诺氟沙星植物毒性的研究,探讨抗生素的环境毒理学特性和潜在的环境风险。主要研究结果如下：
诺氟沙星在黑土、潮土和红壤中的吸附特性存在差异。随溶液中诺氟沙星浓度的增加,诺氟沙星在三种土壤和相应的去有机质土壤中的吸附量逐渐增加,但是去有机质土壤中的吸附量要高于相应的母体土壤,这说明土壤有机质的存在降低了诺氟沙星在土壤中的吸附量。不同模型对试验数据拟合结果表明,Freundlich方程能够对诺氟沙星在六种土壤(三种原状土壤,三种去有机质土壤)上的吸附特性进行较好的拟合,而Langmuir方程只能拟合在黑土,红壤及它们对应的去有机质土壤中的结果,完全不能用于对潮土的研究结果。该结果说明Freundlich方程比Langmuir方程更适用于土壤中诺氟沙星在吸附特性的研究。在本研究条件下,土壤溶液pH值对诺氟沙星吸附有较大影响,在黑土、潮土和红壤三种供试土壤中最大的吸附量分别出现各自土壤溶液pH为6.1—6.9,6.6—7.4和5.7—6.0的区间内,而且红壤中的KD和Koc值均高于其它两种土壤的。铜离子对诺氟沙星在土壤中的吸附表现出一定的浓度效应,随着添加铜离子浓度的增加,三种土壤中诺氟沙星吸附量、KD和Koc值均表现为先升高,在100mg·L-1达到峰值,而后下降的变化趋势。
随着培养时间的增加,无论是在灭菌土壤还是在非灭菌土壤中,诺氟沙星残留浓度均降低,但是其降解特性存在土壤差异。用一级动力学方程拟合结果很好的描述了土壤中诺氟沙星的降解过程,所得结果表明,在本实验条件下,潮土中降解较快,其次为红壤,黑土中降解最慢。
在整个培养期间,污染物对土壤酶活性的影响与污染物浓度和处理时间相关,而且土壤之间存在差异。添加污染物后,土壤过氧化氢酶活性无显著变化,而对土壤脲酶、磷酸酶、蔗糖酶活性则产生显著影响。土壤中添加铜后,土壤酶活性受到抑制,特别是在培养后期表现出明显的浓度效应。就土壤磷酸酶而言,Norl处理对黑土中磷酸酶活性表现为激活作用,尤其是在培养28天后,该处理酶活比对照处理酶活增加了47%,而在潮土中与对照处理相比,Norl处理在培养前14天,土壤磷酸酶活性升高,而在培养28天后则对土壤酶活性表现为抑制作用。复合污染显著的抑制了黑土土壤磷酸酶活性,然而在潮土中,仅在培养前28天,复合污染处理对磷酸酶活性的影响显著高于污染物单独处理的。对于土壤脲酶和蔗糖酶的影响也表现出了相同的土壤差异的趋势。
诺氟沙星单独及与铜复合处理均影响土壤微生物群落代谢功能多样性,且与污染物浓度和处理时间有很大的关系。AWCD值变化与处理强度和处理时间有关,而且与处理时间关系更加密切,污染处理28天后,各处理间差异达最大,Nor1O+Cu100显著低于其它处理。诺氟沙星单独处理对土壤微生物群落多样性和均一性、优势种微生物的影响要比复合污染处理的显著。对于不同碳源的利用,也表现出处理和时间的差异。在整个处理过程中,各处理土壤微生物对聚合物类底物的利用强度均比对其它类型高。在暴露处理7天和28天后,Nor10+Cu50, Nor10+Cu100两个复合污染处理土壤微生物对胺类和氨基酸类培养基利用强度较高；另外,暴露处理14和28天后,土壤微生物则对氨基酸类,醣类和羧酸类培养基利用强度也较高。聚类分析结果也表明不同暴露时间的处理之间碳源利用存在一定的差异,暴露3天后的处理被归为同一簇,而暴露28天后的处理则在不同簇出现。
基于16S rDNA PCR-DGGE方法的研究结果表明,诺氟沙星单独或与铜复合污染均对土壤微生物遗传多样性产生影响。污染物处理28天,不同浓度诺氟沙星单独污染和与铜复合污染对土壤微生物区系基因条带的影响有一定的差别。与对照相比,诺氟沙星单独污染对土壤DNA条带数影响较小,诺氟沙星与铜复合污染则显著减少了DNA条带数,尤其是在较高铜浓度的处理中,但是聚类分析结果表明Nor10+Cu100处理与对照被归为同一簇。污染物处理56天后,与对照相比,诺氟沙星单独处理和Nor10+Cu50处理土壤微生物条带增多,Nor10+Cu100处理样品条带数虽少于对照处理,但是却多于同处理第28天时的样品。比较添加污染物处理不同时期与对照处理条带的相似性可以发现,所有处理在处理56天时的相似性均高于处理28天时的。这些结果说明随着处理时间的增加,土壤微生物对污染物胁迫产生了一定的耐性,样品微生物种群丰富度增加,在图谱上即表现为条带数增加,相似性升高。
诺氟沙星对小麦幼苗生长和生理过程的影响存在显著的剂量效应。在1.0mg·L-1时,诺氟沙星促进了小麦幼苗地上部和根系生长。当培养液中浓度达到10mg·L-1时,幼苗生物量和叶片叶绿素含量显著低于对照处理。小麦幼苗丙二醛含量均随着诺氟沙星处理水平的升高而明显增加,而且根系中MDA含量显著地高于地上部。诺氟沙星胁迫同样使小麦幼苗叶片和根系超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)和抗坏血酸过氧化物酶(APX)酶活性发生变化。在地上部,与对照处理相比,诺氟沙星胁迫条件下,地上部SOD酶和POD酶活性显著升高,CAT酶活性仅在10.0和20.0mg·L-1两个处理下有显著的促进作用,而APX酶活性则在1.0和5.0mg·L-1两个处理浓度下有所下降,其它添加处理酶活均显著升高。而在根系中,与对照处理相比,诺氟沙星胁迫使SOD酶活性显著降低,CAT酶活性的变化则为先降低而后升高。另外,添加诺氟沙星显著的抑制了小麦幼苗根系POD酶活,但是促进了APX酶活性。
Pharmaceutical antibiotics are widely used in human and veterinary medicine to treat and prevent infectious bacterial diseases. The pharmaceutical antibiotics are excreted in urine or faeces from the treated body, either unaltered or as metabolites, some of which are still bioactive. Entering into the environment, antibiotics are distributed in various environments through a number of physical and chemical processes, and then influenced the environmental progress. The effects of drugs on environment depend on the dose and form of antibiotics, which were decided by their behavior in environment. The behaviors of antibiotics are related to various environmental properties and pharmaceutical properties in environment. Copper, an industrial product and food additive, has been widely found in manure and sludge, and might finally enter into the agricultural environment. It can bond to some antibiotic molecules through some functional groups. Therefore, norfloxacin (Nor), a fluoroquinolone and most commonly used in animal industries worldwide, was selected in the present study to investigate the sorption and degradation behavior of norfloxacin in various soils, the effect of norfloxacin on soil enzymes, soil microbial community and wheat seedlings using HPLC, BIOLOG and PCR-DGGE. At the same time, the effect of copper on the behavior and ecotoxicity of norfloxacin were studied in the present study. The main results were presented as follows:
Sorption experiments were studied by the batch equibibrium method and sorption of norfloxacin was different in different soils. With increasing of norfloxacin concentrations, sorption amount of norfloxacin increased in both the bulk soils and their SOM-removed soils, but the sorption capacity in SOM-removed soils was higher than that of their corresponding bulk soils, indicating that the process of norfloxacin sorption in soil was influenced by the soil properties including SOM. The sorption data in bulk soils and SOM-removed soils were fitted to Freundlich and Langmuir models. The correlation coefficients suggested that the experimental data fitted better to Freundlich equation than to Langmuir equation. Furthermore, the data from soil F and SOM-removed F could not be described by Langmuir equation. The norfloxacin sorption amount decreased in soil B and soil F, whereas it increased in soil R as solution pH increased. The maximum KD and KOC were achieved in soil R when the equilibrium solution pH was6. And the Nor sorption was also influenced by the exogenous Cu2+ions, which depended on the soil types and Cu2+concentrations. With increasing of Cu2+concentrations in solution, generally, sorption amount, KD and Koc for norfloxacin in soils increased and were up to a peak at100mg·L-1Cu2+, and then the sorption amount decreased regardless of norfloxacin levels.
Results from the experiments with the degradation of norfloxacin suggested that with increasing of incubation time, the residue norfloxacin concentrations were decreased in all samples, while differences were found among soil types. The norfloxacin degradation data were fitted to the first dynamic equation, showing that the norfloxacin degradation was in the order of soil F> soil R> soil B.
No significant differences in hydrogen peroxidase activity were found among soil samples treated with pollutants, however the activities of urease, phosphatase and sucrase were significantly influenced compared with the control. The effects of pollutants on activities of soil enzymes depended on the concentration of pollutant and exposure time, which also based on the soil types. The activities of soil enzymes were inhibited by adding copper, especially in the later incubation time. In Norl treatments, activities of soil phosphatase increased in every treatment in both soils except of the treatment exposure28-day in fluvo-aquic soil compared with respective control. After treating28-day, the activity of soil phosphatase was inhibited in fluvo-aquic soil, while it increased47%in black soil. The effects of cocontamination of norfloxacin and copper on soil enzymes were similar to the effect of norfloxacin alone, but the effect of cocontamination on soil enzymes was stronger and time-dependent.
Biolog was employed to assess the effect of norfloxacin alone and cocotamination with copper on soil microbial functional community. With increasing of exposure time, the differences in average well colour development (AWCD) values were enlarged among treatments, and the AWCD value in28-day Nor10+Cu100treatment was significantly lower than that in any other treatment. Comparing diversity variation of soil microbial communities under different treatments revealed that the richness and evenness of microbial communities in norfloxacin alone treatments were different from that in co-contamination treatments. Higher percent of polymers was utilized by soil microbe in the whole incubation period compared with other substrates, and more amino acids, carbohydrates and carboxylic substrates were utilized in the soil treated with pollutant after14and28days.Cluster analyses also indicated that samples from different treatments could be distinguished from each other, and that exposure3-day treatments sorted into the same cluster, while28-day treatments distributed in different cluster.
The polymerase chain reaction (PCR) based on16S rDNA, followed by denaturing gradient gel electrophoresis (DGGE), was used to describe the effect of norfloxacin alone and cocotamination with copper on soil microbial community. Microbial diversity in soil treated with pollutants changed with time. At28day, no significant changes were found in the number of bands in soil samples treated with norfloxacin alone. Diversity of microorganisms in soil obviously decreased under the stress of norfloxacin and copper compared to control, but the Nor10+Cu100were sorted into the same clusters. At56day, the number of bands increased in norfloxacin alone treatments and Nor10+Cu50compared to control.The number of bands in Nor10+Cu100was lower than that in control, while it was higher than that in the samples treated with Nor10+Cu100for28day. Comparison of the bacterial DGGE profiles among treatments revealed that the similarity of bacterial communities in the56-day treatments was significantly higher than that in the28-day treatments. The changes in microbial community function and composition in response to the pollutants could be attributed to increased tolerance of soil microbe to pollutants (norfloxacin, norfloxacin+copper) in soil system.
The wheat seedlings growth and chlorophyll content were promoted by the treatment with norfloxacin at the rate of1.0mg·L-1, while they were significantly reduced at the rate of10mg·L-1compared with the control. The content of MDA was enhanced under norfloxacin stress and, was significantly higher in roots than in shoots. The activities of superoxide dismutase (SOD), catalase (CAT), guaiacol peroxidase (POD) and ascorbate peroxidase (APX) in wheat seedlings were changed by the treatment with norfloxacin. In shoots, compared with the control treatment, the activities of SOD and POD significantly increased, but the activities of CAT and APX significantly increased only at some norfloxacin concentration in shoots. In roots, the activities of SOD and POD obviously decreased, while the activity of APX remarkably increased by adding norfloxacin.
诺氟沙星在黑土、潮土和红壤中的吸附特性存在差异。随溶液中诺氟沙星浓度的增加,诺氟沙星在三种土壤和相应的去有机质土壤中的吸附量逐渐增加,但是去有机质土壤中的吸附量要高于相应的母体土壤,这说明土壤有机质的存在降低了诺氟沙星在土壤中的吸附量。不同模型对试验数据拟合结果表明,Freundlich方程能够对诺氟沙星在六种土壤(三种原状土壤,三种去有机质土壤)上的吸附特性进行较好的拟合,而Langmuir方程只能拟合在黑土,红壤及它们对应的去有机质土壤中的结果,完全不能用于对潮土的研究结果。该结果说明Freundlich方程比Langmuir方程更适用于土壤中诺氟沙星在吸附特性的研究。在本研究条件下,土壤溶液pH值对诺氟沙星吸附有较大影响,在黑土、潮土和红壤三种供试土壤中最大的吸附量分别出现各自土壤溶液pH为6.1—6.9,6.6—7.4和5.7—6.0的区间内,而且红壤中的KD和Koc值均高于其它两种土壤的。铜离子对诺氟沙星在土壤中的吸附表现出一定的浓度效应,随着添加铜离子浓度的增加,三种土壤中诺氟沙星吸附量、KD和Koc值均表现为先升高,在100mg·L-1达到峰值,而后下降的变化趋势。
随着培养时间的增加,无论是在灭菌土壤还是在非灭菌土壤中,诺氟沙星残留浓度均降低,但是其降解特性存在土壤差异。用一级动力学方程拟合结果很好的描述了土壤中诺氟沙星的降解过程,所得结果表明,在本实验条件下,潮土中降解较快,其次为红壤,黑土中降解最慢。
在整个培养期间,污染物对土壤酶活性的影响与污染物浓度和处理时间相关,而且土壤之间存在差异。添加污染物后,土壤过氧化氢酶活性无显著变化,而对土壤脲酶、磷酸酶、蔗糖酶活性则产生显著影响。土壤中添加铜后,土壤酶活性受到抑制,特别是在培养后期表现出明显的浓度效应。就土壤磷酸酶而言,Norl处理对黑土中磷酸酶活性表现为激活作用,尤其是在培养28天后,该处理酶活比对照处理酶活增加了47%,而在潮土中与对照处理相比,Norl处理在培养前14天,土壤磷酸酶活性升高,而在培养28天后则对土壤酶活性表现为抑制作用。复合污染显著的抑制了黑土土壤磷酸酶活性,然而在潮土中,仅在培养前28天,复合污染处理对磷酸酶活性的影响显著高于污染物单独处理的。对于土壤脲酶和蔗糖酶的影响也表现出了相同的土壤差异的趋势。
诺氟沙星单独及与铜复合处理均影响土壤微生物群落代谢功能多样性,且与污染物浓度和处理时间有很大的关系。AWCD值变化与处理强度和处理时间有关,而且与处理时间关系更加密切,污染处理28天后,各处理间差异达最大,Nor1O+Cu100显著低于其它处理。诺氟沙星单独处理对土壤微生物群落多样性和均一性、优势种微生物的影响要比复合污染处理的显著。对于不同碳源的利用,也表现出处理和时间的差异。在整个处理过程中,各处理土壤微生物对聚合物类底物的利用强度均比对其它类型高。在暴露处理7天和28天后,Nor10+Cu50, Nor10+Cu100两个复合污染处理土壤微生物对胺类和氨基酸类培养基利用强度较高；另外,暴露处理14和28天后,土壤微生物则对氨基酸类,醣类和羧酸类培养基利用强度也较高。聚类分析结果也表明不同暴露时间的处理之间碳源利用存在一定的差异,暴露3天后的处理被归为同一簇,而暴露28天后的处理则在不同簇出现。
基于16S rDNA PCR-DGGE方法的研究结果表明,诺氟沙星单独或与铜复合污染均对土壤微生物遗传多样性产生影响。污染物处理28天,不同浓度诺氟沙星单独污染和与铜复合污染对土壤微生物区系基因条带的影响有一定的差别。与对照相比,诺氟沙星单独污染对土壤DNA条带数影响较小,诺氟沙星与铜复合污染则显著减少了DNA条带数,尤其是在较高铜浓度的处理中,但是聚类分析结果表明Nor10+Cu100处理与对照被归为同一簇。污染物处理56天后,与对照相比,诺氟沙星单独处理和Nor10+Cu50处理土壤微生物条带增多,Nor10+Cu100处理样品条带数虽少于对照处理,但是却多于同处理第28天时的样品。比较添加污染物处理不同时期与对照处理条带的相似性可以发现,所有处理在处理56天时的相似性均高于处理28天时的。这些结果说明随着处理时间的增加,土壤微生物对污染物胁迫产生了一定的耐性,样品微生物种群丰富度增加,在图谱上即表现为条带数增加,相似性升高。
诺氟沙星对小麦幼苗生长和生理过程的影响存在显著的剂量效应。在1.0mg·L-1时,诺氟沙星促进了小麦幼苗地上部和根系生长。当培养液中浓度达到10mg·L-1时,幼苗生物量和叶片叶绿素含量显著低于对照处理。小麦幼苗丙二醛含量均随着诺氟沙星处理水平的升高而明显增加,而且根系中MDA含量显著地高于地上部。诺氟沙星胁迫同样使小麦幼苗叶片和根系超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)和抗坏血酸过氧化物酶(APX)酶活性发生变化。在地上部,与对照处理相比,诺氟沙星胁迫条件下,地上部SOD酶和POD酶活性显著升高,CAT酶活性仅在10.0和20.0mg·L-1两个处理下有显著的促进作用,而APX酶活性则在1.0和5.0mg·L-1两个处理浓度下有所下降,其它添加处理酶活均显著升高。而在根系中,与对照处理相比,诺氟沙星胁迫使SOD酶活性显著降低,CAT酶活性的变化则为先降低而后升高。另外,添加诺氟沙星显著的抑制了小麦幼苗根系POD酶活,但是促进了APX酶活性。
Pharmaceutical antibiotics are widely used in human and veterinary medicine to treat and prevent infectious bacterial diseases. The pharmaceutical antibiotics are excreted in urine or faeces from the treated body, either unaltered or as metabolites, some of which are still bioactive. Entering into the environment, antibiotics are distributed in various environments through a number of physical and chemical processes, and then influenced the environmental progress. The effects of drugs on environment depend on the dose and form of antibiotics, which were decided by their behavior in environment. The behaviors of antibiotics are related to various environmental properties and pharmaceutical properties in environment. Copper, an industrial product and food additive, has been widely found in manure and sludge, and might finally enter into the agricultural environment. It can bond to some antibiotic molecules through some functional groups. Therefore, norfloxacin (Nor), a fluoroquinolone and most commonly used in animal industries worldwide, was selected in the present study to investigate the sorption and degradation behavior of norfloxacin in various soils, the effect of norfloxacin on soil enzymes, soil microbial community and wheat seedlings using HPLC, BIOLOG and PCR-DGGE. At the same time, the effect of copper on the behavior and ecotoxicity of norfloxacin were studied in the present study. The main results were presented as follows:
Sorption experiments were studied by the batch equibibrium method and sorption of norfloxacin was different in different soils. With increasing of norfloxacin concentrations, sorption amount of norfloxacin increased in both the bulk soils and their SOM-removed soils, but the sorption capacity in SOM-removed soils was higher than that of their corresponding bulk soils, indicating that the process of norfloxacin sorption in soil was influenced by the soil properties including SOM. The sorption data in bulk soils and SOM-removed soils were fitted to Freundlich and Langmuir models. The correlation coefficients suggested that the experimental data fitted better to Freundlich equation than to Langmuir equation. Furthermore, the data from soil F and SOM-removed F could not be described by Langmuir equation. The norfloxacin sorption amount decreased in soil B and soil F, whereas it increased in soil R as solution pH increased. The maximum KD and KOC were achieved in soil R when the equilibrium solution pH was6. And the Nor sorption was also influenced by the exogenous Cu2+ions, which depended on the soil types and Cu2+concentrations. With increasing of Cu2+concentrations in solution, generally, sorption amount, KD and Koc for norfloxacin in soils increased and were up to a peak at100mg·L-1Cu2+, and then the sorption amount decreased regardless of norfloxacin levels.
Results from the experiments with the degradation of norfloxacin suggested that with increasing of incubation time, the residue norfloxacin concentrations were decreased in all samples, while differences were found among soil types. The norfloxacin degradation data were fitted to the first dynamic equation, showing that the norfloxacin degradation was in the order of soil F> soil R> soil B.
No significant differences in hydrogen peroxidase activity were found among soil samples treated with pollutants, however the activities of urease, phosphatase and sucrase were significantly influenced compared with the control. The effects of pollutants on activities of soil enzymes depended on the concentration of pollutant and exposure time, which also based on the soil types. The activities of soil enzymes were inhibited by adding copper, especially in the later incubation time. In Norl treatments, activities of soil phosphatase increased in every treatment in both soils except of the treatment exposure28-day in fluvo-aquic soil compared with respective control. After treating28-day, the activity of soil phosphatase was inhibited in fluvo-aquic soil, while it increased47%in black soil. The effects of cocontamination of norfloxacin and copper on soil enzymes were similar to the effect of norfloxacin alone, but the effect of cocontamination on soil enzymes was stronger and time-dependent.
Biolog was employed to assess the effect of norfloxacin alone and cocotamination with copper on soil microbial functional community. With increasing of exposure time, the differences in average well colour development (AWCD) values were enlarged among treatments, and the AWCD value in28-day Nor10+Cu100treatment was significantly lower than that in any other treatment. Comparing diversity variation of soil microbial communities under different treatments revealed that the richness and evenness of microbial communities in norfloxacin alone treatments were different from that in co-contamination treatments. Higher percent of polymers was utilized by soil microbe in the whole incubation period compared with other substrates, and more amino acids, carbohydrates and carboxylic substrates were utilized in the soil treated with pollutant after14and28days.Cluster analyses also indicated that samples from different treatments could be distinguished from each other, and that exposure3-day treatments sorted into the same cluster, while28-day treatments distributed in different cluster.
The polymerase chain reaction (PCR) based on16S rDNA, followed by denaturing gradient gel electrophoresis (DGGE), was used to describe the effect of norfloxacin alone and cocotamination with copper on soil microbial community. Microbial diversity in soil treated with pollutants changed with time. At28day, no significant changes were found in the number of bands in soil samples treated with norfloxacin alone. Diversity of microorganisms in soil obviously decreased under the stress of norfloxacin and copper compared to control, but the Nor10+Cu100were sorted into the same clusters. At56day, the number of bands increased in norfloxacin alone treatments and Nor10+Cu50compared to control.The number of bands in Nor10+Cu100was lower than that in control, while it was higher than that in the samples treated with Nor10+Cu100for28day. Comparison of the bacterial DGGE profiles among treatments revealed that the similarity of bacterial communities in the56-day treatments was significantly higher than that in the28-day treatments. The changes in microbial community function and composition in response to the pollutants could be attributed to increased tolerance of soil microbe to pollutants (norfloxacin, norfloxacin+copper) in soil system.
The wheat seedlings growth and chlorophyll content were promoted by the treatment with norfloxacin at the rate of1.0mg·L-1, while they were significantly reduced at the rate of10mg·L-1compared with the control. The content of MDA was enhanced under norfloxacin stress and, was significantly higher in roots than in shoots. The activities of superoxide dismutase (SOD), catalase (CAT), guaiacol peroxidase (POD) and ascorbate peroxidase (APX) in wheat seedlings were changed by the treatment with norfloxacin. In shoots, compared with the control treatment, the activities of SOD and POD significantly increased, but the activities of CAT and APX significantly increased only at some norfloxacin concentration in shoots. In roots, the activities of SOD and POD obviously decreased, while the activity of APX remarkably increased by adding norfloxacin.
引文
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