颗粒化EBPR系统混合碳源分子水平作用机制及模型构建研究
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摘要
本研究接种污水处理厂回流污泥,以乙酸和丙酸为交替碳源进行驯化,成功实现聚磷菌(PAOs)富集。以荧光原位杂交(FISH)技术检测证实获得的除磷污泥中聚磷菌占全菌比例达75±5%。试验进而以富含PAOs的除磷污泥为基础,在乙酸和丙酸为混合碳源的条件下培养颗粒化强化生物除磷(EBPR)系统。考察碳源中乙酸／丙酸比例的系列变化对颗粒化EBPR系统的作用机制。并以PCR-DGGE、16S rDNA克隆转化等分子生物学技术,探明系列混合碳源条件对颗粒化EBPR系统微生物群落结构的演化影响规律。试验运行约90 d,结果表明混合碳源中乙酸／丙酸比例的系列变化对系统颗粒粒径生长速率、污泥沉降性能、形态结构、系统除磷效率等方面的影响都存在着非常明显的规律性。污泥颗粒化进程及系列混合碳源条件变化对颗粒化EBPR系统微生物群落结构产生了较大的筛选作用。
随着乙酸在混合碳源中的比例增加,系统颗粒粒径生长速率明显降低,颗粒体积平均粒径明显减小。至试验运行末期,混合碳源中乙酸比例为0%、25%、50%、75%与100%的系统形成的成熟颗粒污泥体积平均粒径分别为745.08μm、680.99μm、642.38μm、599.41μm与550.64μm,污泥容积指数指数(SVI)分别为75 mL/g、60 mL/g、50 mL/g、40 mL/g与30 mL/g。碳源中丙酸比例较高的系统所形成的成熟颗粒颜色近于白色,外表面更光滑,为椭球形。碳源中乙酸比例较高的系统成熟颗粒颜色近于米黄色,外表面不如前者光滑,但颗粒结构明显更致密,为圆球形。在相同的磷处理负荷下,各系统除磷效率之间存在显著性差异。碳源中乙酸比例为0%、25%、50%及100%的系统平均除磷效率分别为97.0%、85.9%,77.4%,56.5%及31.5%。各试验系统平均除磷效率与系统成熟颗粒平均粒径大小之间存在一定的正相关性。
试验初期,各系统接种的除磷污泥中共检测到22个菌种。PAOs在系统中占优势,主要包括Acinetobacter、Uncultured alpha proteobacterium、Uncultured Chlorobi bacterium、Uncultured bacterium以及Uncultured Rhodocyclaceae bacterium。系统中存在的聚糖菌(GAOs)主要为Candidatus Competibacter phosphatis。此外,系统中也检测到部分反硝化菌群Denitrifying bacterium。随着试验的进行,污泥颗粒化进程对系统群落结构产生了较大的筛选作用。试验运行30 d后,碳源中乙酸比例为0%、25%、50%、75%及100%的系统群落结构与试验初始状态的相似性分别为48.5%、43.0%、47.6%、41.4%及43.5%。原本在系统中占优势的一类聚磷菌Uncultured bacterium被迅速淘汰；聚磷菌Uncultured Rhodocyclaceae bacterium、部分聚糖菌Candidatus Competibacter phosphatis以及部分反硝化菌群Denitrifying bacterium也分别被淘汰。而聚磷菌Acinetobacter、Uncultured alpha proteobacterium以及’Uncultured Chlorobi bacterium在各系统中仍占优势。试验运行90 d,各系统群落结构与试验初始状态的相似性进一步下降至40%以下。在各运行稳定的颗粒化EBPR系统中适应生存的菌种主要包括Candidatus Competibacter phosphatis、Uncultured alpha proteobacterium、Uncultured Chlorobi bacterium、Uncultured bacterium以及Denitrifying bacterium。从整体上看,成熟的颗粒化EBPR系统相比普通的非颗粒化EBPR系统,微生物群落结构更简单。
不同混合碳源条件对颗粒化EBPR系统群落结构的影响主要表现为Candidatus Competibacter phosphatis (GAOs)与Uncultured Chlorobi bacterium (PAOs)菌种数量的差异。当试验系统碳源中存在乙酸时,Candidatus Competibacter phosphatis能在系统中生长；而当试验系统碳源中只存在丙酸而不存在乙酸时,Candidatus Competibacter phosphatis被淘汰。系统中Candidatus Competibacter phosphatis的数量与系统碳源中乙酸的比例之间存在一定的正相关性。系统混合碳源中乙酸比例的提高造成聚糖菌Candidatus Competibacter phosphatis的增长,是系统除磷效率下降的主要原因。在碳源中丙酸比例较高的系统中,特别是100%丙酸碳源的系统中,聚磷菌Uncultured Chlorobi bacterium有所增长,而在丙酸比例较低的系统中未发现此现象。Uncultured Chlorobi bacterium的增长可能也是碳源中丙酸比例相对较高的系统除磷效率相对较好的原因之一。而不同混合碳源条件对成熟颗粒化EBPR系统中的Uncultured bacterium与Denitrifying bacterium生长没有太大影响,Uncultured alpha proteobacterium及部分Uncultured Chlorobi bacterium在各成熟颗粒化EBPR系统中也都能得到较好的生长。
此外,本研究参考全耦合活性污泥3号模型中的生物除磷反应方程,以及SBR工艺好氧颗粒污泥广义模型中的物质传输扩散方程,将两者结合提出了颗粒化EBPR系统模型的初步构建。
In order to obtain highly enriched cultures of phosphate-accumulating organisms (PAOs), activated sludge from a local sewage treatment plant was cultivated in lab-scale sequencing batch reactors (SBR) with alternating acetate and propionate as carbon sources. After operating for over a month, the anaerobic phosphorus release rate, as well as aerobic phosphorus uptake rate, of the system were increased from 0.60-0.75 mg/(g·h) to over 8.5 mg/(g·h). Meanwhile, the phosphorus removal efficiency of the system was increased from 10% to over 90%. As assessed by fluorescence in situ hybridization (FISH) quantification, the proportion of PAOs in all bacteria was 75±5%, which indicated that the highly enrichment of PAOs was achieved.
Then, the activated sludge highly enriched of PAOs were seeded, and cultivated with mixed carbon sources of different acetate to propionate ratio, to obtain granule-based enhanced biological phosphorus removal (EBPR) system. The long-term effects of acetate to propionate ratio on the granulation progress of EBPR system were investigated. Furthermore, the microbial community of the granule-based EBPR systems cultivated with mixed carbon sources of different acetate to propionate ratio were investigated by molecular biotechnology such as polymerase chain reaction (PCR), denaturing gradient gel electrophoresis (DGGE), 16S rDNA clone, and etc.
After operating for 90 d, the results showed that the effects of acetate to propionate ratio on the granule growth rate, granule sludge characteristics and morphology, and phosphorus removal efficiency were all of regularity. The granule growth rate was obviously decreased as the acetate to propionate ratio increased, which leading to a gradient of granule volume average particle diameter of different granule-based EBPR systems. The volume average particle diameters of the mature granules were 745.08μm,680.99μm,642.38μm,599.41μm, and 550.64μm in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively. And the sludge volume index (SVI) of the mature granules were 75 mL/g,60 mL/g,50 mL/g,40 mL/g and 30 mL/g in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively. The granules of all systems were formed after operation for 20 d, and grew mature after 90 d. In the systems with relatively higher proportion of propionate in the mixed carbon sources than acetate, the mature granules were tended to have white color, smooth surface and ellipsoidal shape. In the systems with relatively lower proportion of propionate in the mixed carbon sources than acetate, the mature granules were tended to have buff color, dense microstructure and spheroidal shape. The phosphorus removal of granule-based EBPR system showed significant differences among the treatments, which were 97.0%,85.9%,77.4%,56.5% and 31.5% in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively.
At the beginning of the treatment, variety of microbial community was detected in the seeding sludge, which were about 22 different bacteriums. PAOs was dominant in the seeding sludge, including Acinetobacter, Uncultured alpha proteobacterium, Uncultured Chlorobi bacterium, Uncultured bacterium and Uncultured Rhodocyclaceae bacterium. GAOs existed in the seeding sludge was Candidatus Competibacter phosphatis. Besides that, there was certain Denitrifying bacterium in the seeding sludge as well.
After operating for 30 d, the similarities of the microbial community were 48.5%,43.0%,47.6%,41.4% and 43.5% in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively, comparing to the beginning of the treatment. It indicated that the microbial community structures of all systems were changed significantly during the sludge granulation progress. Uncultured bacterium, a type of PAOs that was dominant in the seeding sludge, was disappeared in all systems. Uncultured Rhodocyclaceae bacterium, some Candidatus Competibacter phosphatis and some Denitrifying bacterium were disappeared as well, while Acinetobacter, Uncultured alpha proteobacterium and Uncultured Chlorobi bacterium were still dominant in the systems.
The granules grew mature in all systems after operating for three months; meanwhile the microbial community structure reached stable state. In the mature granule-based EBPR system, the microbial community was mainly including Candidatus Competibacter phosphatis, Uncultured alpha proteobacterium, Uncultured Chlorobi bacterium, Uncultured bacterium and Denitrifying bacterium. Comparing to the non-granule-based EBPR system, the microbial community of granule-based EBPR system was relatively pure.
The differences among the granule-based EBPR system cultivated with mixed carbon sources of different acetate to propionate ratio, was mainly manifested as the quantity variance of Candidatus Competibacter phosphatis (GAOs) and Uncultured Chlorobi bacterium (PAOs). As the proportion of acetate in the mixed carbon sources increased, Candidatus Competibacter phosphatis proliferated in the system, which was responsible for the decline of phosphorus removal efficiency. Moreover, Uncultured Chlorobi bacterium was found to proliferate in the system with higher proportion of propionate in the mixed carbon sources, which should be another reason of the higher phosphorus removal efficiency in those systems.
Additionally, the fully coupled activated sludge model No.3 (FCASM3) and a generalized model for aerobic granule-based SBR were adjusted and employed for describing the biological reaction and mass transfer progress in the granule-based EBPR system, respectively. Based on those two models, a preliminary model for granule-based EBPR system was conceived.
随着乙酸在混合碳源中的比例增加,系统颗粒粒径生长速率明显降低,颗粒体积平均粒径明显减小。至试验运行末期,混合碳源中乙酸比例为0%、25%、50%、75%与100%的系统形成的成熟颗粒污泥体积平均粒径分别为745.08μm、680.99μm、642.38μm、599.41μm与550.64μm,污泥容积指数指数(SVI)分别为75 mL/g、60 mL/g、50 mL/g、40 mL/g与30 mL/g。碳源中丙酸比例较高的系统所形成的成熟颗粒颜色近于白色,外表面更光滑,为椭球形。碳源中乙酸比例较高的系统成熟颗粒颜色近于米黄色,外表面不如前者光滑,但颗粒结构明显更致密,为圆球形。在相同的磷处理负荷下,各系统除磷效率之间存在显著性差异。碳源中乙酸比例为0%、25%、50%及100%的系统平均除磷效率分别为97.0%、85.9%,77.4%,56.5%及31.5%。各试验系统平均除磷效率与系统成熟颗粒平均粒径大小之间存在一定的正相关性。
试验初期,各系统接种的除磷污泥中共检测到22个菌种。PAOs在系统中占优势,主要包括Acinetobacter、Uncultured alpha proteobacterium、Uncultured Chlorobi bacterium、Uncultured bacterium以及Uncultured Rhodocyclaceae bacterium。系统中存在的聚糖菌(GAOs)主要为Candidatus Competibacter phosphatis。此外,系统中也检测到部分反硝化菌群Denitrifying bacterium。随着试验的进行,污泥颗粒化进程对系统群落结构产生了较大的筛选作用。试验运行30 d后,碳源中乙酸比例为0%、25%、50%、75%及100%的系统群落结构与试验初始状态的相似性分别为48.5%、43.0%、47.6%、41.4%及43.5%。原本在系统中占优势的一类聚磷菌Uncultured bacterium被迅速淘汰；聚磷菌Uncultured Rhodocyclaceae bacterium、部分聚糖菌Candidatus Competibacter phosphatis以及部分反硝化菌群Denitrifying bacterium也分别被淘汰。而聚磷菌Acinetobacter、Uncultured alpha proteobacterium以及’Uncultured Chlorobi bacterium在各系统中仍占优势。试验运行90 d,各系统群落结构与试验初始状态的相似性进一步下降至40%以下。在各运行稳定的颗粒化EBPR系统中适应生存的菌种主要包括Candidatus Competibacter phosphatis、Uncultured alpha proteobacterium、Uncultured Chlorobi bacterium、Uncultured bacterium以及Denitrifying bacterium。从整体上看,成熟的颗粒化EBPR系统相比普通的非颗粒化EBPR系统,微生物群落结构更简单。
不同混合碳源条件对颗粒化EBPR系统群落结构的影响主要表现为Candidatus Competibacter phosphatis (GAOs)与Uncultured Chlorobi bacterium (PAOs)菌种数量的差异。当试验系统碳源中存在乙酸时,Candidatus Competibacter phosphatis能在系统中生长；而当试验系统碳源中只存在丙酸而不存在乙酸时,Candidatus Competibacter phosphatis被淘汰。系统中Candidatus Competibacter phosphatis的数量与系统碳源中乙酸的比例之间存在一定的正相关性。系统混合碳源中乙酸比例的提高造成聚糖菌Candidatus Competibacter phosphatis的增长,是系统除磷效率下降的主要原因。在碳源中丙酸比例较高的系统中,特别是100%丙酸碳源的系统中,聚磷菌Uncultured Chlorobi bacterium有所增长,而在丙酸比例较低的系统中未发现此现象。Uncultured Chlorobi bacterium的增长可能也是碳源中丙酸比例相对较高的系统除磷效率相对较好的原因之一。而不同混合碳源条件对成熟颗粒化EBPR系统中的Uncultured bacterium与Denitrifying bacterium生长没有太大影响,Uncultured alpha proteobacterium及部分Uncultured Chlorobi bacterium在各成熟颗粒化EBPR系统中也都能得到较好的生长。
此外,本研究参考全耦合活性污泥3号模型中的生物除磷反应方程,以及SBR工艺好氧颗粒污泥广义模型中的物质传输扩散方程,将两者结合提出了颗粒化EBPR系统模型的初步构建。
In order to obtain highly enriched cultures of phosphate-accumulating organisms (PAOs), activated sludge from a local sewage treatment plant was cultivated in lab-scale sequencing batch reactors (SBR) with alternating acetate and propionate as carbon sources. After operating for over a month, the anaerobic phosphorus release rate, as well as aerobic phosphorus uptake rate, of the system were increased from 0.60-0.75 mg/(g·h) to over 8.5 mg/(g·h). Meanwhile, the phosphorus removal efficiency of the system was increased from 10% to over 90%. As assessed by fluorescence in situ hybridization (FISH) quantification, the proportion of PAOs in all bacteria was 75±5%, which indicated that the highly enrichment of PAOs was achieved.
Then, the activated sludge highly enriched of PAOs were seeded, and cultivated with mixed carbon sources of different acetate to propionate ratio, to obtain granule-based enhanced biological phosphorus removal (EBPR) system. The long-term effects of acetate to propionate ratio on the granulation progress of EBPR system were investigated. Furthermore, the microbial community of the granule-based EBPR systems cultivated with mixed carbon sources of different acetate to propionate ratio were investigated by molecular biotechnology such as polymerase chain reaction (PCR), denaturing gradient gel electrophoresis (DGGE), 16S rDNA clone, and etc.
After operating for 90 d, the results showed that the effects of acetate to propionate ratio on the granule growth rate, granule sludge characteristics and morphology, and phosphorus removal efficiency were all of regularity. The granule growth rate was obviously decreased as the acetate to propionate ratio increased, which leading to a gradient of granule volume average particle diameter of different granule-based EBPR systems. The volume average particle diameters of the mature granules were 745.08μm,680.99μm,642.38μm,599.41μm, and 550.64μm in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively. And the sludge volume index (SVI) of the mature granules were 75 mL/g,60 mL/g,50 mL/g,40 mL/g and 30 mL/g in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively. The granules of all systems were formed after operation for 20 d, and grew mature after 90 d. In the systems with relatively higher proportion of propionate in the mixed carbon sources than acetate, the mature granules were tended to have white color, smooth surface and ellipsoidal shape. In the systems with relatively lower proportion of propionate in the mixed carbon sources than acetate, the mature granules were tended to have buff color, dense microstructure and spheroidal shape. The phosphorus removal of granule-based EBPR system showed significant differences among the treatments, which were 97.0%,85.9%,77.4%,56.5% and 31.5% in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively.
At the beginning of the treatment, variety of microbial community was detected in the seeding sludge, which were about 22 different bacteriums. PAOs was dominant in the seeding sludge, including Acinetobacter, Uncultured alpha proteobacterium, Uncultured Chlorobi bacterium, Uncultured bacterium and Uncultured Rhodocyclaceae bacterium. GAOs existed in the seeding sludge was Candidatus Competibacter phosphatis. Besides that, there was certain Denitrifying bacterium in the seeding sludge as well.
After operating for 30 d, the similarities of the microbial community were 48.5%,43.0%,47.6%,41.4% and 43.5% in the treatment of 0%,25%,50%,75% and 100% acetate in mixed carbon sources, respectively, comparing to the beginning of the treatment. It indicated that the microbial community structures of all systems were changed significantly during the sludge granulation progress. Uncultured bacterium, a type of PAOs that was dominant in the seeding sludge, was disappeared in all systems. Uncultured Rhodocyclaceae bacterium, some Candidatus Competibacter phosphatis and some Denitrifying bacterium were disappeared as well, while Acinetobacter, Uncultured alpha proteobacterium and Uncultured Chlorobi bacterium were still dominant in the systems.
The granules grew mature in all systems after operating for three months; meanwhile the microbial community structure reached stable state. In the mature granule-based EBPR system, the microbial community was mainly including Candidatus Competibacter phosphatis, Uncultured alpha proteobacterium, Uncultured Chlorobi bacterium, Uncultured bacterium and Denitrifying bacterium. Comparing to the non-granule-based EBPR system, the microbial community of granule-based EBPR system was relatively pure.
The differences among the granule-based EBPR system cultivated with mixed carbon sources of different acetate to propionate ratio, was mainly manifested as the quantity variance of Candidatus Competibacter phosphatis (GAOs) and Uncultured Chlorobi bacterium (PAOs). As the proportion of acetate in the mixed carbon sources increased, Candidatus Competibacter phosphatis proliferated in the system, which was responsible for the decline of phosphorus removal efficiency. Moreover, Uncultured Chlorobi bacterium was found to proliferate in the system with higher proportion of propionate in the mixed carbon sources, which should be another reason of the higher phosphorus removal efficiency in those systems.
Additionally, the fully coupled activated sludge model No.3 (FCASM3) and a generalized model for aerobic granule-based SBR were adjusted and employed for describing the biological reaction and mass transfer progress in the granule-based EBPR system, respectively. Based on those two models, a preliminary model for granule-based EBPR system was conceived.
引文
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