外加微量联氨影响全自养脱氮的机制：功能微生物群落与N_2O产生
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摘要
与传统的硝化-反硝化工艺相比，完全自养脱氮工艺（completely autotrophicnitrogen removal over nitrite, CANON）在理论上可节省62.5%的硝化所需氧量和100%的反硝化需氧有机COD量，是符合当前废水处理节能减排发展要求的可持续废水脱氮处理新技术。课题组前期对CANON过程的影响因素、脱氮贡献和外加联氨（N2H4）强化全自养脱氮能力等进行了研究。本论文从外加微量N2H4对好氧氨氧化及亚硝酸盐氧化动力学特性、CANON系统的N2O的产生特性和功能微生物群落的影响三方面进行研究。目的是通过优化CANON系统的外加N2H4水平，在强化厌氧氨氧化和抑制亚硝酸盐氧化菌（nitrite oxidation bacteria, NOB）活性的同时，降低对氨氧化菌（ammonia oxidation bacteria, AOB）活性的负面影响，综合提高系统脱氮性能、减少N2O产生，为CANON工艺的工程应用提供重要科学依据与基础数据。本论文取得的主要研究结果如下：
①建立与羟胺（NH2OH）关联的好氧氨氧化两步动力学表达，将硝化过程分为三步，应用氧利用速率（oxygen uptake rate, OUR）的呼吸测量法估计了好氧氨氧化两步过程及亚硝酸盐氧化过程的动力学参数，深化了对生物硝化过程的理解和认识。
②N2H4对好氧氨氧化及对亚硝酸盐氧化动力学抑制类型分别为竞争性和非竞争性抑制，N2H4对亚硝酸盐氧化的抑制大于其对好氧氨氧化的抑制；建立了N2H4抑制硝化过程三步动力学模型，通过模拟外加微量N2H4条件下硝化反应各过程外源性呼吸剖面线，首次得到N2H4好氧氨氧化过程基质半饱和常数和最大比基质利用速率（分别为7.96±0.811mgN2H4-N/L,0.0916±0.0188mgN2H4-N/mgCOD/h）、N2H4抑制NH2OH氧化和亚硝酸盐氧化的动力学参数估计值（分别为7.88±0.783mgN2H4-N/L和1.223±0.555mgN2H4-N/L）。
③硝化污泥中的AOB氨单加氧酶α亚基（amoA）克隆序列主要聚于亚硝化球菌（Nitrosococcu）属和亚硝化单胞菌（Nitrosomonas）属，另有少部分属于亚硝化螺旋菌（Nitrosospira）属；NOB亚硝酸盐氧化还原酶β亚基（nxrB）克隆序列聚于硝化细菌（Nitrobacter）属，大部分属于硝化细菌属中的Nitrobacterwinogradskyi，仅有少部分属于Nitrobacter vulgaris和Nitrobacter hamburgensis；长期外加微量N2H4对硝化污泥中AOB与NOB的生长均具有明显抑制作用。
④CANON污泥中的AOB amoA基因克隆序列主要聚于Nitrosomonas属和Nitrosococcus属；NOB nxrB基因克隆序列聚于Nitrobacter属；厌氧氨氧化（anaerobic ammonia oxidation, ANAMMOX）菌群落组成较为丰富，部分ANAMMOX菌联氨合酶α亚基（hzsA）克隆序列聚于Candidatus Scalindua属；长期外加微量N2H4促进ANAMMOX菌生长，对AOB生长产生部分抑制作用。
⑤长期添加微量N2H4强化CANON富集得到的CANON污泥颗粒主要呈球状和椭球状，外表层有着较紧密的微生物结构及少量细小的孔隙，导致DO不易被穿透形成内部缺/厌氧区域；好氧的AOB主要位于颗粒外层，厌氧的ANAMMOX菌主要位于颗粒污泥内部。
⑥限氧-厌氧交替运行的CANON系统中，N2O产生的途径包括NH2OH生物氧化与AOB反硝化；限氧条件下，NH2OH生物氧化的贡献大于AOB反硝化；厌氧条件下，N2O主要由AOB反硝化产生。
⑦CANON污泥的间歇试验结果表明，限氧条件下外加微量N2H4减少N2O产生；厌氧条件下外加微量N2H4促进N2O产生；在交替限氧-厌氧的CANON反应器中，外加微量N2H4在限氧阶段被完全降解，消除了其在厌氧阶段可能的负面影响；因此，在交替限氧-厌氧CANON反应器中，外加微量N2H4在提高脱氮性能的同时降低了N2O的产生；在长期外加微量N2H4的交替限氧-厌氧的CANON反应器中，N2O平均产生速率为0.066±0.047mgN/L/d,仅占总氮去除的0.018±0.013%，低于相关文献报道结果。
Comparing to traditional nitrification-denitrification process, completelyautotrophic nitrogen removal over nitrite (CANON) process, which is consistent withthe current requirements for energy saving and emission reduction and sustainabledevelopment of wastewater treatment technologies, can decrease O2consuming as muchas60%and hardly consumed COD theoretically. The previous research has beenfocused on influence factors, contributions of nitrogen removal and the capability oftrace hydrazine (N2H4) addtion enhancing for CANON process. Therefore, further studyfor the effect of trace N2H4addtion on characterization of kinetics for ammonium andnitrite oxidation, characteristics of N2O production and functional microbial communityof CANON process were investigated in this thesis. By optimizing the additive N2H4concentration in a CANON system to enhance anammox and inhibite NOB activity inmaximum extent while minimizing the inhibitory influence on AOB activity, and furtherincreasing the denitrification efficiency with N2O production decreasing, whichprovided important data and scientific foundation for engineering application ofCANON. The results in this thesis are given as follows:
①The two-step kinetics expression of aerobic ammonium oxidation associatedwith hydroxylamine (NH2OH) was set up; the nitrifying process was dividied intothree-step and kinetic parameters of aerobic ammonium oxidation and nitrite oxidationwere estimated using the respirometry with oxygen uptake rate (OUR); this deepens thecomprehending and understanding of biological nitrification
②The kinetics for ammonium oxidation and nitrite oxidation under the inhibitionof hydrazine was competitive and noncompetitive, respectively, the inhibitive effect ofN2H4on nitrite oxidation was stronger than that on ammonia oxidation; the three-stepkinetics model of nitrification inhibited by N2H4was established, and the maximumspecific rate and half-saturation coefficient of N2H4oxidation (7.96±0.811mgN2H4-N/L,0.0916±0.0188mgN2H4-N/mgCOD/h, respectively) with the nitrifying sludge, and theinhibition coefficients of N2H4for hydroxylamine oxidation and nitrite oxidation(7.88±0.783mgN2H4-N/L,1.223±0.555mgN2H4-N/L, respectively) was estimated forthe first time by simulating exogenous OUR profiles of nitrifying reaction with traceN2H4addtion, respectively.
③In nitrifying sludge, cloning sequences of ammonia monooxygenase α subunit (amoA) of AOB were mainly related to Nitrosococcu and Nitrosomonas genuses, whilethe rest were clustering in the Nitrosospira genus; cloning sequences of nitriteoxidordeuctase β subunit (nxrB) of NOB were related to Nitrobacter genus, the majoritywere Nitrobacter winogradskyi specie, only a small part was clustering to Nitrobactervulgaris and Nitrobacter hamburgensis; long-time trace N2H4addtion couldsiginificantly restrained the growth of both AOB and NOB.
④In CANON sludge, AOB amoA cloning sequences were closely related toNitrosomonas and Nitrosococcus genuses; NOB nxrB cloning sequences were related toNitrobacter genus; the community composition of anaerobic ammonia oxidation(ANAMMOX) bacteria was abundant, and some cloning sequences of hydrazinesynthase α subunit (hzsA) of ANAMMOX bacteria were clustering in CandidatusScalindua genus; long-time trace N2H4addtion partially inhibit the growth of AOB butobviously promote the growth of ANAMMOX bacteria.
⑤The enrichment CANON granular sludge in CANON reactor with long-timeN2H4addtion are mainly spherical and ellipsoidal, and there was a more compactstructure and a few tiny microbes gap in outer layer, which made DO could not easilypenetrated the granule surface and caused anoxic/anaerobic region formed; aerobicAOB mainly located at the surface layer of granular sludge; anaerobic ANAMMOXbacteria are mainly located inside the granular sludge.
⑥In CANON system with alternant oxygen limited and anaerobic operationmode, pathways of N2O production include NH2OH biological oxidation and nitrifierdenitrification by AOB; under oxygen limited conditions, N2O produced by NH2OHbiological oxidation was more than that by nitrifier denitrification via AOB; underanaerobic conditions, N2O was mainly produced by nitrifier denitrification via AOB.
⑦The batch tests with CANON sludge indicated that under oxygen limitedconditions, trace N2H4addition decrease N2O production; under anaerobic conditions,trace N2H4addition accelerate N2O production; in CANON reactor with alternantoxygen limited-anaerobic operation mode, trace N2H4added in oxygen limited stagewas completely degraded to eliminate the possible negative influence of N2H4onanaerobic stage; therefore, in CANON reactor with alternant oxygen limited andanaerobic operation mode, trace N2H4addtion could enhance the denitrificationefficiency while effectively reducing N2O production; in the enhanced CANON processby long-time trace N2H4addition under alternant oxygen limited and anaerobicoperation mode, N2O average production rate lower at0.066±0.047mgN/L/d was only 0.018±0.013%of the total nitrogen removal，which was below the results reported byreferences.
①建立与羟胺（NH2OH）关联的好氧氨氧化两步动力学表达，将硝化过程分为三步，应用氧利用速率（oxygen uptake rate, OUR）的呼吸测量法估计了好氧氨氧化两步过程及亚硝酸盐氧化过程的动力学参数，深化了对生物硝化过程的理解和认识。
②N2H4对好氧氨氧化及对亚硝酸盐氧化动力学抑制类型分别为竞争性和非竞争性抑制，N2H4对亚硝酸盐氧化的抑制大于其对好氧氨氧化的抑制；建立了N2H4抑制硝化过程三步动力学模型，通过模拟外加微量N2H4条件下硝化反应各过程外源性呼吸剖面线，首次得到N2H4好氧氨氧化过程基质半饱和常数和最大比基质利用速率（分别为7.96±0.811mgN2H4-N/L,0.0916±0.0188mgN2H4-N/mgCOD/h）、N2H4抑制NH2OH氧化和亚硝酸盐氧化的动力学参数估计值（分别为7.88±0.783mgN2H4-N/L和1.223±0.555mgN2H4-N/L）。
③硝化污泥中的AOB氨单加氧酶α亚基（amoA）克隆序列主要聚于亚硝化球菌（Nitrosococcu）属和亚硝化单胞菌（Nitrosomonas）属，另有少部分属于亚硝化螺旋菌（Nitrosospira）属；NOB亚硝酸盐氧化还原酶β亚基（nxrB）克隆序列聚于硝化细菌（Nitrobacter）属，大部分属于硝化细菌属中的Nitrobacterwinogradskyi，仅有少部分属于Nitrobacter vulgaris和Nitrobacter hamburgensis；长期外加微量N2H4对硝化污泥中AOB与NOB的生长均具有明显抑制作用。
④CANON污泥中的AOB amoA基因克隆序列主要聚于Nitrosomonas属和Nitrosococcus属；NOB nxrB基因克隆序列聚于Nitrobacter属；厌氧氨氧化（anaerobic ammonia oxidation, ANAMMOX）菌群落组成较为丰富，部分ANAMMOX菌联氨合酶α亚基（hzsA）克隆序列聚于Candidatus Scalindua属；长期外加微量N2H4促进ANAMMOX菌生长，对AOB生长产生部分抑制作用。
⑤长期添加微量N2H4强化CANON富集得到的CANON污泥颗粒主要呈球状和椭球状，外表层有着较紧密的微生物结构及少量细小的孔隙，导致DO不易被穿透形成内部缺/厌氧区域；好氧的AOB主要位于颗粒外层，厌氧的ANAMMOX菌主要位于颗粒污泥内部。
⑥限氧-厌氧交替运行的CANON系统中，N2O产生的途径包括NH2OH生物氧化与AOB反硝化；限氧条件下，NH2OH生物氧化的贡献大于AOB反硝化；厌氧条件下，N2O主要由AOB反硝化产生。
⑦CANON污泥的间歇试验结果表明，限氧条件下外加微量N2H4减少N2O产生；厌氧条件下外加微量N2H4促进N2O产生；在交替限氧-厌氧的CANON反应器中，外加微量N2H4在限氧阶段被完全降解，消除了其在厌氧阶段可能的负面影响；因此，在交替限氧-厌氧CANON反应器中，外加微量N2H4在提高脱氮性能的同时降低了N2O的产生；在长期外加微量N2H4的交替限氧-厌氧的CANON反应器中，N2O平均产生速率为0.066±0.047mgN/L/d,仅占总氮去除的0.018±0.013%，低于相关文献报道结果。
Comparing to traditional nitrification-denitrification process, completelyautotrophic nitrogen removal over nitrite (CANON) process, which is consistent withthe current requirements for energy saving and emission reduction and sustainabledevelopment of wastewater treatment technologies, can decrease O2consuming as muchas60%and hardly consumed COD theoretically. The previous research has beenfocused on influence factors, contributions of nitrogen removal and the capability oftrace hydrazine (N2H4) addtion enhancing for CANON process. Therefore, further studyfor the effect of trace N2H4addtion on characterization of kinetics for ammonium andnitrite oxidation, characteristics of N2O production and functional microbial communityof CANON process were investigated in this thesis. By optimizing the additive N2H4concentration in a CANON system to enhance anammox and inhibite NOB activity inmaximum extent while minimizing the inhibitory influence on AOB activity, and furtherincreasing the denitrification efficiency with N2O production decreasing, whichprovided important data and scientific foundation for engineering application ofCANON. The results in this thesis are given as follows:
①The two-step kinetics expression of aerobic ammonium oxidation associatedwith hydroxylamine (NH2OH) was set up; the nitrifying process was dividied intothree-step and kinetic parameters of aerobic ammonium oxidation and nitrite oxidationwere estimated using the respirometry with oxygen uptake rate (OUR); this deepens thecomprehending and understanding of biological nitrification
②The kinetics for ammonium oxidation and nitrite oxidation under the inhibitionof hydrazine was competitive and noncompetitive, respectively, the inhibitive effect ofN2H4on nitrite oxidation was stronger than that on ammonia oxidation; the three-stepkinetics model of nitrification inhibited by N2H4was established, and the maximumspecific rate and half-saturation coefficient of N2H4oxidation (7.96±0.811mgN2H4-N/L,0.0916±0.0188mgN2H4-N/mgCOD/h, respectively) with the nitrifying sludge, and theinhibition coefficients of N2H4for hydroxylamine oxidation and nitrite oxidation(7.88±0.783mgN2H4-N/L,1.223±0.555mgN2H4-N/L, respectively) was estimated forthe first time by simulating exogenous OUR profiles of nitrifying reaction with traceN2H4addtion, respectively.
③In nitrifying sludge, cloning sequences of ammonia monooxygenase α subunit (amoA) of AOB were mainly related to Nitrosococcu and Nitrosomonas genuses, whilethe rest were clustering in the Nitrosospira genus; cloning sequences of nitriteoxidordeuctase β subunit (nxrB) of NOB were related to Nitrobacter genus, the majoritywere Nitrobacter winogradskyi specie, only a small part was clustering to Nitrobactervulgaris and Nitrobacter hamburgensis; long-time trace N2H4addtion couldsiginificantly restrained the growth of both AOB and NOB.
④In CANON sludge, AOB amoA cloning sequences were closely related toNitrosomonas and Nitrosococcus genuses; NOB nxrB cloning sequences were related toNitrobacter genus; the community composition of anaerobic ammonia oxidation(ANAMMOX) bacteria was abundant, and some cloning sequences of hydrazinesynthase α subunit (hzsA) of ANAMMOX bacteria were clustering in CandidatusScalindua genus; long-time trace N2H4addtion partially inhibit the growth of AOB butobviously promote the growth of ANAMMOX bacteria.
⑤The enrichment CANON granular sludge in CANON reactor with long-timeN2H4addtion are mainly spherical and ellipsoidal, and there was a more compactstructure and a few tiny microbes gap in outer layer, which made DO could not easilypenetrated the granule surface and caused anoxic/anaerobic region formed; aerobicAOB mainly located at the surface layer of granular sludge; anaerobic ANAMMOXbacteria are mainly located inside the granular sludge.
⑥In CANON system with alternant oxygen limited and anaerobic operationmode, pathways of N2O production include NH2OH biological oxidation and nitrifierdenitrification by AOB; under oxygen limited conditions, N2O produced by NH2OHbiological oxidation was more than that by nitrifier denitrification via AOB; underanaerobic conditions, N2O was mainly produced by nitrifier denitrification via AOB.
⑦The batch tests with CANON sludge indicated that under oxygen limitedconditions, trace N2H4addition decrease N2O production; under anaerobic conditions,trace N2H4addition accelerate N2O production; in CANON reactor with alternantoxygen limited-anaerobic operation mode, trace N2H4added in oxygen limited stagewas completely degraded to eliminate the possible negative influence of N2H4onanaerobic stage; therefore, in CANON reactor with alternant oxygen limited andanaerobic operation mode, trace N2H4addtion could enhance the denitrificationefficiency while effectively reducing N2O production; in the enhanced CANON processby long-time trace N2H4addition under alternant oxygen limited and anaerobicoperation mode, N2O average production rate lower at0.066±0.047mgN/L/d was only 0.018±0.013%of the total nitrogen removal，which was below the results reported byreferences.
引文
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