生物增强活性炭优势菌群稳定及竞争特征研究
摘要
水源污染和可利用水资源的日益短缺问题为保障供水安全提出了挑战。生物增强活性炭技术(Bio-enhanced Activated Carbon,BEAC)将筛选出的优势菌群固定在活性炭载体上,用于饮用水深度处理,可以增强对有机物的降解效能、提高降解速率,因此受到广泛关注。目前对于BEAC工艺的研究重点在于通过选择适宜的活性炭、调控工艺运行,利于优势菌群生长、防止外入菌群生长、保持高生物活性和生物量,确保工艺稳定运行。然而以上研究均未涉及优势菌的菌群结构动态变化特点,优势菌间的相互关系,以及优势菌群对生态位的竞争关系。优势菌群是BEAC工艺的核心,研究优势菌群的稳定特征及竞争关系,对于保障BEAC工艺稳定运行具有重要意义。
为了减少由DNA提取不当导致的DGGE分析结果偏差,本文首先进行了3种不同活性炭上菌群DNA提取方法的研究,通过测定DNA量、DNA纯度和DGGE图谱,结果表明采用方法C(20 kHz、输入能量为40 W的超声波预处理120 s,用1% CTAB、100μL 10 mg/L蛋白酶K和1.5 mL 20% SDS对菌体细胞裂解)提取的基因组总DNA经DGGE分析后,可获得完整的DGGE菌群图谱,为最佳DNA提取方案。针对难以应用FISH检测活性炭上菌群的问题,采用超声振荡分离活性炭上菌群后,在不同条件下进行杂交试验,通过测定菌群数量,发现FISH方法??可获得最多的菌群数量,其具体步骤为:超声振荡120 s预处理后,样品用8μL含有50 ng探针的杂交缓冲液,在40°C恒温下杂交3 h,之后用不含探针的杂交缓冲液在42°C条件下洗脱20 min。
应用上述最佳DNA提取方法和FISH检测方法,对普通生物活性炭(BAC)滤池进行菌群监测,确定优势菌的菌属,并进行分离,获得5株优势菌(SRO 2,SRO 11,SRO19,SRO 20和SRO 30)。通过优势菌对TOC降解效果研究,以及测定世代时间和脱氢酶活性,发现筛选出的优势菌与其他细菌相比,同时具备较高的TOC降解能力、快速生长能力和高脱氢酶活性。
将筛选的5株优势菌混合后固定于活性炭上,建立BEAC工艺体系。采用最佳DNA提取,及DGGE、ATP、SEM和FISH等技术对比分析了BEAC和普通BAC工艺中菌群动态变化,结果发现,优势菌具有代谢能力强、生物活性高和生长速度快的优点,在BEAC工艺运行期间,上、中、下层菌群生物活性比普通BAC高(BEAC工艺运行至90 d时,菌群生物活性已超过1000 ng ATP/g炭,运行至180 d时,上、中、下层菌群生物活性均超过1500 ng ATP/g炭,而BAC工艺运行期间,最高生物活性仅为1023.5 ng ATP/g炭);BEAC工艺菌群生物活性增长速度较普通BAC快(BEAC工艺菌群活性平均增长速率为7.70[ng ATP·g~(-1)炭·d~(-1)],而普通BAC工艺仅为5.12[ng ATP·g~(-1)炭·d~(-1)]),BEAC工艺对TOC的平均去除率为76.24%,而普通BAC仅为58.97%。BEAC工艺中优势菌群沿活性炭柱上、中、下层的分布相对均一,各层优势菌群均能充分发挥作用,提高了BEAC工艺对有机物的去除效率,优势菌群对提高污染物降解效果和工艺稳定运行发挥了重要作用。
优势菌SRO 30在BEAC工艺中的比例最高,在运行至180 d时,占生境中总生物量的50%左右;其次是SRO 19,占总生物量的25%以上。BEAC工艺运行期间,优势菌群总量呈增加趋势,但优势菌SRO 11数量逐渐减少。在对BEAC工艺优势菌群竞争关系研究时发现,种群竞争与最低资源(有机物)需求(R~*)有关,由于BEAC工艺中可利用的资源(有机物)低于优势菌SRO 11的最低资源(有机物)需求(R~*),导致180 d后SRO 11从工艺体系中消失。
假单胞菌属的SRO 2、SRO 11、SRO 19和SRO 20之间以利用性竞争为主,SRO 30与4株假单胞菌(SRO 2、SRO 11、SRO 19和SRO 20)之间以干扰性竞争为主。应用Lotka-Volterra方程计算了BEAC工艺中假单胞菌属(SRO 2,SRO 19和SRO 20)和枯草芽孢杆菌SRO 30的负荷量K_1和K_2,竞争系数α和β,结果表明,在一定条件下,BEAC滤池上、中、下层的假单胞菌属竞争系数α均小于K_1/K_2,枯草芽孢杆菌竞争系数β均小于K_2/K_1,枯草芽孢杆菌(SRO 30)和假单胞菌属(SRO 2,SRO 19和SRO 20)可以共存于BEAC反应体系中,最终形成稳定体系。
通过以上研究,建立了一种活性炭微生物DNA提取和FISH检测方法,为活性炭上菌群分析提供了可靠的技术手段;发现了枯草芽孢杆菌Bacillus subtilis SRO 30和穿孔假单胞菌Pseudomonas pertucinogena SRO 19在BEAC工艺中的显著优势地位;通过数学模型预测了优势菌稳定特性,发现假单胞菌SRO 2、SRO 19和SRO 20和枯草芽孢杆菌SRO 30可以形成稳定体系。
The insurance of water supply security has in face of challenge because of water shortage and pollution. Bio-enhanced activated carbon (BEAC) is a kind of advanced drinking water treatment process, which fixed the isolated bacteria on activated carbon. The use of BEAC has become a matter of concern to many researchers, because it can improve organic pollutants removal efficiency. At present, the study of BEAC focuses on activated carbon properties in use and proper parameters of the process. The purpose of these studies are keep the dominant bacteria and BEAC process stable by means of improving the growth of dominant bacteria, preventing external bacteria from invading, keeping dominant bacterial quantity and activity. However, these studies did not concern about the importance of dominant bacteria, including the dynamics of dominant bacteria and relationship between these bacteria.
In this study, three different DNA extraction protocols were used for evaluating the DNA extraction efficiency of bacteria attached on activated carbon by DGGE analysis. The results showed that method C with 120 seconds ultra-sonication under 20 kHz and 40 W input energy could produce the intact bacterial profiles, which was the most proper DNA extraction protocol. After comparison with three different fluorescence in situ hybridization methods, the results showed that 120 s ultra-sonication pretreatment was benefit for bacteria separated from activated carbon. The most proper hybridization condition was that the samples were hybridized for 3 hours in 8μL hybridization solutions with 50 ng probes at 40°C, then eluted in hybridization for 20 minutes at 42°C.
With the most proper DNA extraction and FISH methods described above, bacterial community of BAC filter operated for 300 days was tested, and dominant bacteria were isolated and screened. It was found that five bacteria (SRO 2, SRO 11, SRO 19, SRO 20 and SRO 30) biodegraded TOC with higher efficiency, grew faster, and activated with higher dehydrigebase activity.
Five bacteria were combined and fixed on activated carbon to form BEAC process. Parameters of BEAC process were tested by comparing with BAC, including water qualities of influent and effluent water, bacterial activity, structure and dynamics with DNA extraction, ATP, SEM and FISH analysis. The results showed that dominant bacterial played an important role in improving pollutants removal efficiency and stable operation of process. Bacterial activity of BEAC filter was higher than that of BAC filter. On the 90~(th) day of BEAC process, bacterial activity was above 1000 ngATP/g carbon. On the 180~(th) day of BEAC process, bacterial activity was above 1500 ngATP/g carboon, while the highest bacterial activity was 1023.5 ngATP/g carbon during BAC process. Bacterial activity of the BEAC filter increased faster than BAC because of the dominant bacteria attached on BEAC with the properties of strong metabolism, high biological activity and low growth time. The average increase rate of bacterial activity in BEAC filter was 7.70 ngATP·g~(-1)carbon·d~(-1), while it was 5.12 ngATP·g~(-1)carbon·d~(-1) in BAC filter. The higher biological activity of BEAC improved pollutants removal efficiency. During BEAC operation, average TOC removal efficiency of BEAC was 76.24%, while its of BAC was only 58.97%. Bacterial distribution along the BEAC filter was relatively more uniform than BAC, which made bacteria on each layer of the filter act and promoted the removal efficiency.
On the 180~(th) day of BEAC operation, the amount of SRO 30 was the highest, with 50% of total biomass. The ammount of SRO 19 was 25% of total biomass. The amount of dominant bacteria was increased during BEAC process, while the amount of SRO 11 was decreased and eliminated on the 180~(th) day of operation. The study of competition between dominant bacteria on BEAC showed that the competition was related to the minus required resource (R~*). The organic resouce in influent was lower than R~* of SRO 11, so it limited the growth of SRO 11.
The competition between SRO 2, SRO 11, SRO 19 and SRO 20 of Pseudomonas genus is exploitative. The competition between SRO 2, SRO 11, SRO 19, SRO 20 and SRO 30 is interfering. The environmental load of Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 are represented by K_1 and K_2 respectively. The competition coefficient of Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 are represented byαandβ. K_1, K_2,αandβwere calculated by using Lotka-Volterra fomula. The results showed thatαwas lower than K_1/K_2, andβwas lower than K_2/K_1, which indicated that Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 could coexist in the BEAC process.
After the above study, the most proper DNA extraction and FISH methods were constructed, which could provide the reliable technologies for future study of bacteria on BAC or BEAC. It has been confirmed by this research that Bacillus subtilis SRO 30 and Pseudomonas pertucinogena SRO 19 obviously dominant achieved in BEAC ecosystem. The reasons about the dominance of BEAC were pointed out. The choice of dominant bacteria is important for BEAC process stability, during which should avoid overlapping resource, choose bacteria with low minus required resource (R~*), minimize internal competition. The forecasting results of dominant bacterial competition by using methametic fomula showed that Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 could coexist in the BEAC process, forming stable bacteria community.
为了减少由DNA提取不当导致的DGGE分析结果偏差,本文首先进行了3种不同活性炭上菌群DNA提取方法的研究,通过测定DNA量、DNA纯度和DGGE图谱,结果表明采用方法C(20 kHz、输入能量为40 W的超声波预处理120 s,用1% CTAB、100μL 10 mg/L蛋白酶K和1.5 mL 20% SDS对菌体细胞裂解)提取的基因组总DNA经DGGE分析后,可获得完整的DGGE菌群图谱,为最佳DNA提取方案。针对难以应用FISH检测活性炭上菌群的问题,采用超声振荡分离活性炭上菌群后,在不同条件下进行杂交试验,通过测定菌群数量,发现FISH方法??可获得最多的菌群数量,其具体步骤为:超声振荡120 s预处理后,样品用8μL含有50 ng探针的杂交缓冲液,在40°C恒温下杂交3 h,之后用不含探针的杂交缓冲液在42°C条件下洗脱20 min。
应用上述最佳DNA提取方法和FISH检测方法,对普通生物活性炭(BAC)滤池进行菌群监测,确定优势菌的菌属,并进行分离,获得5株优势菌(SRO 2,SRO 11,SRO19,SRO 20和SRO 30)。通过优势菌对TOC降解效果研究,以及测定世代时间和脱氢酶活性,发现筛选出的优势菌与其他细菌相比,同时具备较高的TOC降解能力、快速生长能力和高脱氢酶活性。
将筛选的5株优势菌混合后固定于活性炭上,建立BEAC工艺体系。采用最佳DNA提取,及DGGE、ATP、SEM和FISH等技术对比分析了BEAC和普通BAC工艺中菌群动态变化,结果发现,优势菌具有代谢能力强、生物活性高和生长速度快的优点,在BEAC工艺运行期间,上、中、下层菌群生物活性比普通BAC高(BEAC工艺运行至90 d时,菌群生物活性已超过1000 ng ATP/g炭,运行至180 d时,上、中、下层菌群生物活性均超过1500 ng ATP/g炭,而BAC工艺运行期间,最高生物活性仅为1023.5 ng ATP/g炭);BEAC工艺菌群生物活性增长速度较普通BAC快(BEAC工艺菌群活性平均增长速率为7.70[ng ATP·g~(-1)炭·d~(-1)],而普通BAC工艺仅为5.12[ng ATP·g~(-1)炭·d~(-1)]),BEAC工艺对TOC的平均去除率为76.24%,而普通BAC仅为58.97%。BEAC工艺中优势菌群沿活性炭柱上、中、下层的分布相对均一,各层优势菌群均能充分发挥作用,提高了BEAC工艺对有机物的去除效率,优势菌群对提高污染物降解效果和工艺稳定运行发挥了重要作用。
优势菌SRO 30在BEAC工艺中的比例最高,在运行至180 d时,占生境中总生物量的50%左右;其次是SRO 19,占总生物量的25%以上。BEAC工艺运行期间,优势菌群总量呈增加趋势,但优势菌SRO 11数量逐渐减少。在对BEAC工艺优势菌群竞争关系研究时发现,种群竞争与最低资源(有机物)需求(R~*)有关,由于BEAC工艺中可利用的资源(有机物)低于优势菌SRO 11的最低资源(有机物)需求(R~*),导致180 d后SRO 11从工艺体系中消失。
假单胞菌属的SRO 2、SRO 11、SRO 19和SRO 20之间以利用性竞争为主,SRO 30与4株假单胞菌(SRO 2、SRO 11、SRO 19和SRO 20)之间以干扰性竞争为主。应用Lotka-Volterra方程计算了BEAC工艺中假单胞菌属(SRO 2,SRO 19和SRO 20)和枯草芽孢杆菌SRO 30的负荷量K_1和K_2,竞争系数α和β,结果表明,在一定条件下,BEAC滤池上、中、下层的假单胞菌属竞争系数α均小于K_1/K_2,枯草芽孢杆菌竞争系数β均小于K_2/K_1,枯草芽孢杆菌(SRO 30)和假单胞菌属(SRO 2,SRO 19和SRO 20)可以共存于BEAC反应体系中,最终形成稳定体系。
通过以上研究,建立了一种活性炭微生物DNA提取和FISH检测方法,为活性炭上菌群分析提供了可靠的技术手段;发现了枯草芽孢杆菌Bacillus subtilis SRO 30和穿孔假单胞菌Pseudomonas pertucinogena SRO 19在BEAC工艺中的显著优势地位;通过数学模型预测了优势菌稳定特性,发现假单胞菌SRO 2、SRO 19和SRO 20和枯草芽孢杆菌SRO 30可以形成稳定体系。
The insurance of water supply security has in face of challenge because of water shortage and pollution. Bio-enhanced activated carbon (BEAC) is a kind of advanced drinking water treatment process, which fixed the isolated bacteria on activated carbon. The use of BEAC has become a matter of concern to many researchers, because it can improve organic pollutants removal efficiency. At present, the study of BEAC focuses on activated carbon properties in use and proper parameters of the process. The purpose of these studies are keep the dominant bacteria and BEAC process stable by means of improving the growth of dominant bacteria, preventing external bacteria from invading, keeping dominant bacterial quantity and activity. However, these studies did not concern about the importance of dominant bacteria, including the dynamics of dominant bacteria and relationship between these bacteria.
In this study, three different DNA extraction protocols were used for evaluating the DNA extraction efficiency of bacteria attached on activated carbon by DGGE analysis. The results showed that method C with 120 seconds ultra-sonication under 20 kHz and 40 W input energy could produce the intact bacterial profiles, which was the most proper DNA extraction protocol. After comparison with three different fluorescence in situ hybridization methods, the results showed that 120 s ultra-sonication pretreatment was benefit for bacteria separated from activated carbon. The most proper hybridization condition was that the samples were hybridized for 3 hours in 8μL hybridization solutions with 50 ng probes at 40°C, then eluted in hybridization for 20 minutes at 42°C.
With the most proper DNA extraction and FISH methods described above, bacterial community of BAC filter operated for 300 days was tested, and dominant bacteria were isolated and screened. It was found that five bacteria (SRO 2, SRO 11, SRO 19, SRO 20 and SRO 30) biodegraded TOC with higher efficiency, grew faster, and activated with higher dehydrigebase activity.
Five bacteria were combined and fixed on activated carbon to form BEAC process. Parameters of BEAC process were tested by comparing with BAC, including water qualities of influent and effluent water, bacterial activity, structure and dynamics with DNA extraction, ATP, SEM and FISH analysis. The results showed that dominant bacterial played an important role in improving pollutants removal efficiency and stable operation of process. Bacterial activity of BEAC filter was higher than that of BAC filter. On the 90~(th) day of BEAC process, bacterial activity was above 1000 ngATP/g carbon. On the 180~(th) day of BEAC process, bacterial activity was above 1500 ngATP/g carboon, while the highest bacterial activity was 1023.5 ngATP/g carbon during BAC process. Bacterial activity of the BEAC filter increased faster than BAC because of the dominant bacteria attached on BEAC with the properties of strong metabolism, high biological activity and low growth time. The average increase rate of bacterial activity in BEAC filter was 7.70 ngATP·g~(-1)carbon·d~(-1), while it was 5.12 ngATP·g~(-1)carbon·d~(-1) in BAC filter. The higher biological activity of BEAC improved pollutants removal efficiency. During BEAC operation, average TOC removal efficiency of BEAC was 76.24%, while its of BAC was only 58.97%. Bacterial distribution along the BEAC filter was relatively more uniform than BAC, which made bacteria on each layer of the filter act and promoted the removal efficiency.
On the 180~(th) day of BEAC operation, the amount of SRO 30 was the highest, with 50% of total biomass. The ammount of SRO 19 was 25% of total biomass. The amount of dominant bacteria was increased during BEAC process, while the amount of SRO 11 was decreased and eliminated on the 180~(th) day of operation. The study of competition between dominant bacteria on BEAC showed that the competition was related to the minus required resource (R~*). The organic resouce in influent was lower than R~* of SRO 11, so it limited the growth of SRO 11.
The competition between SRO 2, SRO 11, SRO 19 and SRO 20 of Pseudomonas genus is exploitative. The competition between SRO 2, SRO 11, SRO 19, SRO 20 and SRO 30 is interfering. The environmental load of Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 are represented by K_1 and K_2 respectively. The competition coefficient of Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 are represented byαandβ. K_1, K_2,αandβwere calculated by using Lotka-Volterra fomula. The results showed thatαwas lower than K_1/K_2, andβwas lower than K_2/K_1, which indicated that Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 could coexist in the BEAC process.
After the above study, the most proper DNA extraction and FISH methods were constructed, which could provide the reliable technologies for future study of bacteria on BAC or BEAC. It has been confirmed by this research that Bacillus subtilis SRO 30 and Pseudomonas pertucinogena SRO 19 obviously dominant achieved in BEAC ecosystem. The reasons about the dominance of BEAC were pointed out. The choice of dominant bacteria is important for BEAC process stability, during which should avoid overlapping resource, choose bacteria with low minus required resource (R~*), minimize internal competition. The forecasting results of dominant bacterial competition by using methametic fomula showed that Pseudomonas genus SRO 2, SRO 19, SRO 20 and Bacillus subtilis SRO 30 could coexist in the BEAC process, forming stable bacteria community.
引文
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