好氧颗粒污泥同步硝化反硝化过程研究
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摘要
利用厌氧颗粒污泥通风培养驯化获得具有同步硝化反硝化功能的好氧颗粒污泥，使异
养菌、硝化菌和反硝化茵等共存于一个颗粒中，微生物间形成相互协作、互为共生的关系，
构成一个具有同步硝化反硝化功能的反应体系。
本文研究了同步硝化反硝化好氧颗粒污泥特性、反应过程以及微生物相分布，硝化反
硝化反应模式及通氧优化，好氧颗粒污泥的动力学常数及传质模式，温室气体N2O的产生
过程及控制方法。主要研究内容如下：
i．好氧颗粒污泥的培养
利用厌氧颗粒污泥好氧培养，93d后转化获得的好氧颗粒污泥具有良好的生物活性和同
步硝化反硝化特性，在反应过程中COD去除率达90％，氨氮去除率近98％，几乎检测不到
NO2--N和NO3--N，好氧颗粒污泥建立了良好的硝化反硝化反应体系。在过程研究中，2h
反应后73． 6％有机物被代谢去除，3h反应后COD浓度降至87m／L，NO2--N和NO3--N在反
应液中的浓度继续下降，6h的反应显示好氧颗粒污泥具有良好的同步硝化反硝化能力。
稳定状态下颗粒的平均直径为2． 26mm，好氧颗粒污泥粒径在2． 1～2． 8mm范围的占全部
颗粒污泥的60％，污泥停留时间约22d；1． 5L反应器中颗粒污泥数量约1． 1×104个，比表面
积为389m2／m3：好氧颗粒污泥的最大沉降速度约9cm／min，所能承受的最大力为23． 236N；
反应器内MLSS 8g／L，SV 15～28％，SVI 20～38ml／g。观察发现好氧颗粒污泥具有适合好氧
和厌氧反应的微环境，颗粒污泥表面结构紧密，主要是球状细菌，内部有较大的空隙，主
要为杆状细菌。
ii．好氧颗粒污泥生长特性
颗粒污泥最适宜硝化反硝化反应的温度范围为25～38℃，最适的pH范围在7～8之间，
最佳的溶解氧浓度在1-2mg／L。好氧颗粒污泥对有机物和氨氮的亲和常数分别为100mg／L
和2mg／L。当COD浓度为1500mg／L时，好氧颗粒污泥对氨氮的利用率下降；当氨氮浓度
超过400mg／L时，有机物降解速率下降，高的氨氮浓度引起硝化中间产物硝酸盐和亚硝酸
J.~_…“_........一一一止生垫鱼丝宜全丝‘一一一
盐的积累，抑制好氧微生物活性。
Luongs模型，仔=叮~
二〕卫二旦己丝{2二，较好地拟合了底物抑制实验过程，模型对coD
C:+Ks
拟合性护为0.9932，对N场+一为尸为0.9916。氨氮对颗粒污泥降解CoD的抑制常数为
凡=lll.lmg几，a=0.836;对氨氮降解的抑制常数为凡=103.lm叭，a=1.08。流加培养过程
中，采用低氨氮浓度(30m叭)流加，过程很少积累N仇一和NOZ一;氨氮流加浓度为200m叭
时，No3一和NoZ一的积累量分别为12m泌和3om叭;氨氮浓度400m叭时，2h后N仇一
的量增加到60m叭，硝化反硝化过程受抑制。
COD吸附实验发现，反应Zh后颗粒污泥吸附COD量达每g干污泥56.7mg。同时，颗
粒污泥中微生物在胞内积累PHAs，积累的PHAs以电子供体的形式参与了颗粒污泥的反硝
化反应。
111.好氧颗粒污泥生物相分析
根据分离获得的细菌形态特征和初步的生理生化特性，初步鉴定出颗粒污泥中含假单
胞菌属(Ps eudomonas)5株，亚硝化单胞菌属(Nitros口monas)7株，硝化杆菌属(从如胡ira)4株、
气单胞菌属(A eromonas)3株、黄单胞菌属〔枷nth口monas)2株、抱杆菌属(加‘illus)4株、黄色
杆菌属(知nthobacter)2株、产碱菌属(A lealigenes)2株。分离获得的菌属中包含了硝化反应
的亚硝化单胞菌属和硝化杆菌属微生物，以及具有好氧反硝化性能的Pseudomonas和
通Icaligenes菌。
摇瓶培养混合微生物进行脱氮实验，30h内有机物去除率在75%，氮去除率近50%。
无菌条件下在多孔载体上培养混合微生物，30d后，微生物在多孔载体上附着生长，氨氮去
除率达到84%，总氮(含氨氮、硝基氮和亚硝基氮)去除率63%。经过培养，含载体的反应器
利用分离得到的微生物逐步形成具有同步硝化和反硝化功能。
iv.好氧颗粒污泥同步硝化反硝化反应机制及溶解氧条件影响
用Monod方程拟合分析不同溶解氧浓度情况下的比硝化反应速率，得硝化反应半饱和
常数(尤浏)和最大比硝化反应速率常数(彻玩工)分别为1.osm叭和40.2助吮MLSS/I nin。拟合
比反硝化速率实验数据，得反硝化反应氧抑制常数(凡动和最大比反硝化反应速率常数
仇加巴)分别为o.806mg几和36.38知岁gMLSS/min。利用动力学参数，可以计算完全同步硝化
反硝化反应所需的溶解氧浓度，为o.927m妙。
不同DO情况下反应CoD去除率均在90%以上，出水cOD低于60 mg几，好氧颗粒
污泥具有良好的有机物代谢能力。高溶解氧浓度份3，omg几)可提高硝化反应速率，但易造成
反应过程积累NoZ一N和No3一N;低溶解氧浓度(夕.omg/L)，反应积累的硝化产物少，颗粒
污泥具有更好的反硝化反应能力。
中文摘要
在好氧颗粒污泥脱氮过程中，短程硝化反硝化与全程硝化反硝化途径并存。高DO情
况下，大部分NOZ一被氧化为NO3一，NO3一N再在颗粒污泥内部还原为气态氮。而在低
DO情况下，NOZ一通过短程反硝化反应直接还原为气态的NZO和NZ。同样情况通过”N
同位素实验可证实，低COD加比由于缺乏有机物而影响颗粒污泥的反硝化反应。研究反应
过程中各中间产物’SN量变化，发现部分氨氮不通过NO3一’加而直接通过 NOZ一’SN?
Anaerobic granular sludge was trained under the condition of aeration to be aerobic granular sludge with the function of simultaneous nitrification and denitrification. Microorganisms like heterotrophic bacteria; nitrification and denitrification bacteria grew in the aerobic granular sludge, and cooperated and supported each other during reaction. The aerobic granular sludge became an ideal place for simultaneous nitrification and denitrification.
With the aerobic granular sludge, the characteristics and reaction process, feature and distribution of microorganisms, the kinetic and mechanism of reaction, the generation and control of N2O were studied in a 2L reactor. Main research and corresponding results are described as follow from five aspects: i. Cultivation of aerobic granular sludge
Anaerobic granular sludge was changed to aerobic granular sludge under the condition of aeration after 93d cultivation. The aerobic granular sludge obtained had the function of simultaneous nitrification and denitrification. Over 90% COD and almost 98% NH4+-N could be removed during the reaction period with aerobic granular sludge, NO2~-N and NO3--N in effluent were not detected. Aerobic granular sludge had established the good reaction system of nitrification and denitrification. In a 6h reaction process, 73.6% COD was removed at 2h, COD degraded to 87mg/L after 3h, while the concentration of NO2~-N and NO3--N was decreased continually. The 6h reaction process showed that the aerobic granular sludge had strong ability of simultaneous nitrification and denitrification.
The aerobic granular sludge obtained had the average diameter 2.26mm, the range of 2.1 ~2.8mm granular sludge was 60% of all sludge. STR was 22d, there were about 1.1 x 104 granular in the 1.5L reaction. The special surface of granular was 389m2/m3, the maximum sedimentation speed was 9cm/min, and the maximum affording force was 23.236N. In the reaction, MLSS was 8g/L, SV 15-28%, SVI 20~38ml/g. Observed under the microscope, it
was found that the granular sludge was a suitable place for the reaction of aerobic and anaerobic reaction, the surface structure of granular sludge was mainly tighten by coccus, and the internal had a lot of gaps grown by bacilli. ii. Growth characteristics of aerobic granular sludge
The best suitable growth temperature of aerobic granular sludge for simultaneous nitrification and denitrification was 25-3 8 , the pH 7-8 and the DO 1 ~2mg/L. The affinity constants of aerobic granular sludge for ammonium and COD were less than 2mg/L and 100mg/L respectively. When the COD concentration and ammonium were over 1500mg/L and 400mg/L respectively, the consumption rates of COD and ammonium decreased correspondingly. With high ammonium concentration in broth, the midst products of nitrification, nitrite and nitrate, accumulated during the reaction that restricted the activity of aerobic granular sludge.
Lungs model, fitted the experimental data best (R2 for COD
0.9932, for NH4+-N 0.9916). The coefficients were Kt=111.1mg/L, a=0.836 for inhibition of COD consumption by ammonium, and Ki=103.1mg/L, a=1.08 for inhibition of ammonium oxidation by ammonium. During cultivation by fed-batch experiment with low concentration of ammonium (30mg/L), there were low nitrite and nitrate existed after reaction; with concentration of 200mg/L, nitrite and nitrate were 12 and 30mg/L in broth; and with concentration of 400mg/L, nitrite was 60mg/L in broth after 2h that restricted the reaction.
With the adsorption experiment of COD, it was found that aerobic granular sludge could adsorb COD 56.7mg/gSS after 2h. Simultaneously, PHAs was accumulated in the bacteria in granular sludge, and it served as the electron donor for effective simultaneous nitrification and denitrification. iii. Microorganism distribution and mixing cultivation of aerobic granular sludge
According to the features of morphology, physiology and biochemistry of 29 bacteria separated from the aerobic granular sludge, it concluded that there were 5 Pseudomonas, 1 Nitrosomonas, 4 Nitrospira, 3
养菌、硝化菌和反硝化茵等共存于一个颗粒中，微生物间形成相互协作、互为共生的关系，
构成一个具有同步硝化反硝化功能的反应体系。
本文研究了同步硝化反硝化好氧颗粒污泥特性、反应过程以及微生物相分布，硝化反
硝化反应模式及通氧优化，好氧颗粒污泥的动力学常数及传质模式，温室气体N2O的产生
过程及控制方法。主要研究内容如下：
i．好氧颗粒污泥的培养
利用厌氧颗粒污泥好氧培养，93d后转化获得的好氧颗粒污泥具有良好的生物活性和同
步硝化反硝化特性，在反应过程中COD去除率达90％，氨氮去除率近98％，几乎检测不到
NO2--N和NO3--N，好氧颗粒污泥建立了良好的硝化反硝化反应体系。在过程研究中，2h
反应后73． 6％有机物被代谢去除，3h反应后COD浓度降至87m／L，NO2--N和NO3--N在反
应液中的浓度继续下降，6h的反应显示好氧颗粒污泥具有良好的同步硝化反硝化能力。
稳定状态下颗粒的平均直径为2． 26mm，好氧颗粒污泥粒径在2． 1～2． 8mm范围的占全部
颗粒污泥的60％，污泥停留时间约22d；1． 5L反应器中颗粒污泥数量约1． 1×104个，比表面
积为389m2／m3：好氧颗粒污泥的最大沉降速度约9cm／min，所能承受的最大力为23． 236N；
反应器内MLSS 8g／L，SV 15～28％，SVI 20～38ml／g。观察发现好氧颗粒污泥具有适合好氧
和厌氧反应的微环境，颗粒污泥表面结构紧密，主要是球状细菌，内部有较大的空隙，主
要为杆状细菌。
ii．好氧颗粒污泥生长特性
颗粒污泥最适宜硝化反硝化反应的温度范围为25～38℃，最适的pH范围在7～8之间，
最佳的溶解氧浓度在1-2mg／L。好氧颗粒污泥对有机物和氨氮的亲和常数分别为100mg／L
和2mg／L。当COD浓度为1500mg／L时，好氧颗粒污泥对氨氮的利用率下降；当氨氮浓度
超过400mg／L时，有机物降解速率下降，高的氨氮浓度引起硝化中间产物硝酸盐和亚硝酸
J.~_…“_........一一一止生垫鱼丝宜全丝‘一一一
盐的积累，抑制好氧微生物活性。
Luongs模型，仔=叮~
二〕卫二旦己丝{2二，较好地拟合了底物抑制实验过程，模型对coD
C:+Ks
拟合性护为0.9932，对N场+一为尸为0.9916。氨氮对颗粒污泥降解CoD的抑制常数为
凡=lll.lmg几，a=0.836;对氨氮降解的抑制常数为凡=103.lm叭，a=1.08。流加培养过程
中，采用低氨氮浓度(30m叭)流加，过程很少积累N仇一和NOZ一;氨氮流加浓度为200m叭
时，No3一和NoZ一的积累量分别为12m泌和3om叭;氨氮浓度400m叭时，2h后N仇一
的量增加到60m叭，硝化反硝化过程受抑制。
COD吸附实验发现，反应Zh后颗粒污泥吸附COD量达每g干污泥56.7mg。同时，颗
粒污泥中微生物在胞内积累PHAs，积累的PHAs以电子供体的形式参与了颗粒污泥的反硝
化反应。
111.好氧颗粒污泥生物相分析
根据分离获得的细菌形态特征和初步的生理生化特性，初步鉴定出颗粒污泥中含假单
胞菌属(Ps eudomonas)5株，亚硝化单胞菌属(Nitros口monas)7株，硝化杆菌属(从如胡ira)4株、
气单胞菌属(A eromonas)3株、黄单胞菌属〔枷nth口monas)2株、抱杆菌属(加‘illus)4株、黄色
杆菌属(知nthobacter)2株、产碱菌属(A lealigenes)2株。分离获得的菌属中包含了硝化反应
的亚硝化单胞菌属和硝化杆菌属微生物，以及具有好氧反硝化性能的Pseudomonas和
通Icaligenes菌。
摇瓶培养混合微生物进行脱氮实验，30h内有机物去除率在75%，氮去除率近50%。
无菌条件下在多孔载体上培养混合微生物，30d后，微生物在多孔载体上附着生长，氨氮去
除率达到84%，总氮(含氨氮、硝基氮和亚硝基氮)去除率63%。经过培养，含载体的反应器
利用分离得到的微生物逐步形成具有同步硝化和反硝化功能。
iv.好氧颗粒污泥同步硝化反硝化反应机制及溶解氧条件影响
用Monod方程拟合分析不同溶解氧浓度情况下的比硝化反应速率，得硝化反应半饱和
常数(尤浏)和最大比硝化反应速率常数(彻玩工)分别为1.osm叭和40.2助吮MLSS/I nin。拟合
比反硝化速率实验数据，得反硝化反应氧抑制常数(凡动和最大比反硝化反应速率常数
仇加巴)分别为o.806mg几和36.38知岁gMLSS/min。利用动力学参数，可以计算完全同步硝化
反硝化反应所需的溶解氧浓度，为o.927m妙。
不同DO情况下反应CoD去除率均在90%以上，出水cOD低于60 mg几，好氧颗粒
污泥具有良好的有机物代谢能力。高溶解氧浓度份3，omg几)可提高硝化反应速率，但易造成
反应过程积累NoZ一N和No3一N;低溶解氧浓度(夕.omg/L)，反应积累的硝化产物少，颗粒
污泥具有更好的反硝化反应能力。
中文摘要
在好氧颗粒污泥脱氮过程中，短程硝化反硝化与全程硝化反硝化途径并存。高DO情
况下，大部分NOZ一被氧化为NO3一，NO3一N再在颗粒污泥内部还原为气态氮。而在低
DO情况下，NOZ一通过短程反硝化反应直接还原为气态的NZO和NZ。同样情况通过”N
同位素实验可证实，低COD加比由于缺乏有机物而影响颗粒污泥的反硝化反应。研究反应
过程中各中间产物’SN量变化，发现部分氨氮不通过NO3一’加而直接通过 NOZ一’SN?
Anaerobic granular sludge was trained under the condition of aeration to be aerobic granular sludge with the function of simultaneous nitrification and denitrification. Microorganisms like heterotrophic bacteria; nitrification and denitrification bacteria grew in the aerobic granular sludge, and cooperated and supported each other during reaction. The aerobic granular sludge became an ideal place for simultaneous nitrification and denitrification.
With the aerobic granular sludge, the characteristics and reaction process, feature and distribution of microorganisms, the kinetic and mechanism of reaction, the generation and control of N2O were studied in a 2L reactor. Main research and corresponding results are described as follow from five aspects: i. Cultivation of aerobic granular sludge
Anaerobic granular sludge was changed to aerobic granular sludge under the condition of aeration after 93d cultivation. The aerobic granular sludge obtained had the function of simultaneous nitrification and denitrification. Over 90% COD and almost 98% NH4+-N could be removed during the reaction period with aerobic granular sludge, NO2~-N and NO3--N in effluent were not detected. Aerobic granular sludge had established the good reaction system of nitrification and denitrification. In a 6h reaction process, 73.6% COD was removed at 2h, COD degraded to 87mg/L after 3h, while the concentration of NO2~-N and NO3--N was decreased continually. The 6h reaction process showed that the aerobic granular sludge had strong ability of simultaneous nitrification and denitrification.
The aerobic granular sludge obtained had the average diameter 2.26mm, the range of 2.1 ~2.8mm granular sludge was 60% of all sludge. STR was 22d, there were about 1.1 x 104 granular in the 1.5L reaction. The special surface of granular was 389m2/m3, the maximum sedimentation speed was 9cm/min, and the maximum affording force was 23.236N. In the reaction, MLSS was 8g/L, SV 15-28%, SVI 20~38ml/g. Observed under the microscope, it
was found that the granular sludge was a suitable place for the reaction of aerobic and anaerobic reaction, the surface structure of granular sludge was mainly tighten by coccus, and the internal had a lot of gaps grown by bacilli. ii. Growth characteristics of aerobic granular sludge
The best suitable growth temperature of aerobic granular sludge for simultaneous nitrification and denitrification was 25-3 8 , the pH 7-8 and the DO 1 ~2mg/L. The affinity constants of aerobic granular sludge for ammonium and COD were less than 2mg/L and 100mg/L respectively. When the COD concentration and ammonium were over 1500mg/L and 400mg/L respectively, the consumption rates of COD and ammonium decreased correspondingly. With high ammonium concentration in broth, the midst products of nitrification, nitrite and nitrate, accumulated during the reaction that restricted the activity of aerobic granular sludge.
Lungs model, fitted the experimental data best (R2 for COD
0.9932, for NH4+-N 0.9916). The coefficients were Kt=111.1mg/L, a=0.836 for inhibition of COD consumption by ammonium, and Ki=103.1mg/L, a=1.08 for inhibition of ammonium oxidation by ammonium. During cultivation by fed-batch experiment with low concentration of ammonium (30mg/L), there were low nitrite and nitrate existed after reaction; with concentration of 200mg/L, nitrite and nitrate were 12 and 30mg/L in broth; and with concentration of 400mg/L, nitrite was 60mg/L in broth after 2h that restricted the reaction.
With the adsorption experiment of COD, it was found that aerobic granular sludge could adsorb COD 56.7mg/gSS after 2h. Simultaneously, PHAs was accumulated in the bacteria in granular sludge, and it served as the electron donor for effective simultaneous nitrification and denitrification. iii. Microorganism distribution and mixing cultivation of aerobic granular sludge
According to the features of morphology, physiology and biochemistry of 29 bacteria separated from the aerobic granular sludge, it concluded that there were 5 Pseudomonas, 1 Nitrosomonas, 4 Nitrospira, 3
引文
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Bock E, Schmidt I, Stuven R, Zart D. Nitrogen loss caused by denitrifying Nitrosomonas cells using ammonium or hydrogen as electron donors and nitrite as electron accetpor. Arch. Microbiol. 1995,163:16-20
Bock E, Wilderer PA, Freitag A. Growth of Nitrobacter in the absence of dissolved oxygen. Water Res. 1988,22:245-50
Bodelier PLE, Frenzel P. Contribution of methanotrophic and nitrifying bacteria to CH4 and NH4+ oxidation in the rhizosphere of rice plants as determined by new methods of discrimination. Appl. Environ. Microbiol. 1999,65:1826-33
Both GJ, Gerards S, Laanbroek HJ. Kinetics of nitrite oxidation in two Nitrobacter species grown in nitrite-limited chemostats. Arch. Microbiol. 1992, 157:436-41
Brock TD, Madigan MT, Martinko JM, Parker J. Biology of microorganisms. Upper Saddle River NJ, USA. Prentice Hall, 1997
Burns RC, Hardy RWF. Nitrogen fixation in bacteria and higher plants. Berlin, Springer Verlag, 1975 Carpertier B & Cerf O. Biofilms and their consequences, with particular reference to hygiene in the food industry. J Appl. Bacteriol. 1993, 75:499-511
Characklis WG. Laboratory biofilm reactors. In Characklis WG and Marshall KC (Eds.): Biofilms. New York, 1990
Costerton JW, Lewandowski Z, Debeer D. Biofilms, the customized microniche. J. Bacteriol. 1994, 176:2137-42
De Beer D, Vav Der Heuvel JC, Ottenfraf SPP. Microelectrode measurements in nitrifying aggregates. Appl Env Microbiol, 1993, 59: 573-579．
De Boer W, Gunnewiek Pak, Veenhuis M, Bock E, Laanbroek HJ. Nitrification at low pH by aggregated chemolithotrophic bacteria. Appl. Environ. Microbiol. 1991, 57:3600-4
Delong EF, Wickham GS, Pace NR. Phylogenetic strains: ribosomal RNA based probes for the identification of single cells. Science, 1989,243:1360-3
Dolfing J. Acetogenesis. Ecology of Anaerobic Microorganisms, Zehnder (Ed.). Wiley Interscience, 417-468,1988
Ebersold HR, Cordier JL, Luthy P. Bacterial mesosomes: Method dependent artifacts. Arch. Microbiol. 1981, 130: 19-22
Ehrich S, Behrens D, Lebedeva E, Ludwig W, Bock E. A new obligately chemolithoautotrophic, nitrite-oxidizing bacterium, Nitrospira moscoviensis sp nov and its phylogenetic relationship. Arch. Microbiol. 1995,164:16-23
Einsle O, Messerschmidt A, Stach P, Bourenkov GP, Bartunik HD, Huber R, Kroneck PH. Structure of cytochrome c nitrite reductase. Nature. 1999,400:476-80
Etterer T, Wilderer PA. Generation and properties of aerobic granular sludge. Wat Sci Tech, 2001,43(7) :19-26．
Ferguson SJ. Is a proton pumping oxidase essential for energy conservation in Nitrobacter? FEBS Lett.
1982,146:239-43
Ferguson SJ. Denitrification and its control. Antonie van Leeuwenhoek Int. J.Gen. Mol.MicrobioI. 1994, 66:89-110
Ferguson SJ. Nitrogen cycle enzymology. Curr. Opin. Chem. Biol. 1998:182-93
Focht DD, Chang AC. Nitrification and denitrification processes related to wastewater. Adv. Appl. Microbiol, 1975, 19:153-86
Freitag A, Bock E. Energy conservation in Nitrobacter, FEMS Microbiol. Lett. 1990,66:157-62
Gee CS, Pfeffer JT, Suidan MT. Nitrosomonas and Nitrobacter interactions in biological nitrification. J. Environ. Eng. 1990, 116(1) :4-17
Goreau TJ, Kaplan WA, Wofsy SC, McElroy MB, Valois FW, Watson SW. Production of NO2-and N2O by nitrifying bacteria at reduced concentrations of oxygen. Appl. Environ. Microbiol. 1980, 40:526-32
Guiot SR, Pauss A, Costerton WJ. A structured model of the anaerobic granule consortium. Wat. Sci. Tech. 1992,25:1-10
Halling-Sorensen B, Hjuler H. Simultaneous nitrification and denitrification with and upflow fixed bed reactor applying clinoptilolite as media. Wat. Treat, 1992, 7: 77-88
Hanaki K, Hong Z, Matsuo. Production of nitrous oxide gas during denitrification of wastewater. Water Sci. Technol. 1992, 26:1027-36
Harada H, Uemura S, Momonoi K. Interaction between sulfate-reducing bacteria and methane producing bacteria in UASB reactors fed with low strength wastes containing different levels of sulfate. Wat. Res. 1994, 28:355-67
Hellinga C, Schellen AAJC, Mulder JW, Van Loosdrecht MCM, Heijnen JJ. The SHARON process: an innovative method for nitrogen removal from ammonium-rich waste water. Water Sci. Technol. 1998, 37:135-42
Helmer C, Kunst S. Simultaneous nitrification/denitrification in an aerobic biofilm system. Wat. Sci. Tech. 1998, 37(4-5) : 183-877
Hirorns WD, Hastings RC, Head IM, McCarthy AJ, Saunders JR, Pickup RW, Hall GH. Amplification of 16s ribosomal RNA genes of autotrophic ammonia-oxidizing bacteria demonstrates the ubiquity of nitrosospiras in the environment. Microbiology, 1995,141:2793-800
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Juretschko S, Timmermann G, Schmid M, Schleifer KH, Pommereningroser A, Koops HP, Wagner M. Combined molecular and conventional analyses of nitrifying bacterium diversity in activated sludge-nitrosococcus mobilis and nitrospira-like bacteria as dominant populations. Appl. Environ. Microbiol. 1998,64:3042-51
Kester RA, De Boer W, Laanbroek HJ. Production of NO and N2O by pure cultures of nitrifying and denitrifying bacteria during changes in aeration. Appl. Environ. Microbiol. 1997,63:3872-7
Kuenen JG, Robertson LA. Combined nitrification-denitrification processes. FEMS Microbiol. Rev. 1994,15: 109-17
Lettinga G, Hulshoff PLW, Zeeman G. Biological wastewater treatment: Anaerobic Treatment). Department of Environmental Technology, Wageningen Agricultural University, The Netherlands, 1993．
Lewis DL, Gattie DK. Effects of cellular aggregation on the ecology of microorganisms. American Soc. Microbiol. News. 1990,56:263-268
Lloyd H, Ketchum Jr. Design an physical features of sequencing batch reactors. Water Sci. Tech. 1997, 35(1) :11-18
Lipschultz F, Zafiriou OC, Wofsy SC, McElroy MB, Valois FW, Watson SW. Production of NO and N2O by soil nitrifying bacteria. Nature,1981, 294:641-3
Madigan MT, Martinko JM, Parker J. Brock Biology of Microorganisms, Prentice Hall, 473-531, 1997
Masuda S, Watanabe Y, Ishiguro M. Biofilm properties and simultaneous nitrification and denitrification in aerobic rotating biological contactors. Wat. Sci. Tech. 1991, 23(7-9) : 1355-63
Mateju V, Cizinska S, Krejci J. Biological water denitrification-a revies. Enzyme Microb. Technol. 1992, 14:170-183
Mishima K, Makamura M. Self-immovilization of aerobic activated sludge-a pilot study of the aerobic upflow sludge blanket process in municipal sewage treatment. Wat Sci Tech, 1991, 23: 981-990．
Mitchell P. Chemiosmotic coupling and energy transduction. Theor. Exp. Biophys, 1969,2:159-261
Moriyama K, Sato K, Harada Y, Washiyama K and Okamoto K. Simultaneous biologicalremoval of nitrogen and phosphorus using oxic-anaerobic-oxic process. Wat. Sci. Tech. 1990,22(7-8) :61-66
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Munch EV, Lant P, Keller J. Simultaneous nitrification and denitrification in bench-scale sequencing batch reactor. Wat. Res. 1996,30(2) : 277-284
Okayasu Y, Abe I, Matsuo Y. Emission of nitrous oxide from high-rate nitrification and denitrification by mixed liquor circulating process and sequencing batch reactor process. Water Sci. Technol. 1997,36:39-45
Oleszkiewicz JA. Granulation in Anaerobic sludge bed reactors treating food industry wastes. Biological waste. 1988
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Rittmann BE, Langeland WE. Simultaneous denitrification with nitrification in single-channel oxidation ditches. Journal WPCF. 1985, 57(4) :300-308
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Bock E, Wilderer PA, Freitag A. Growth of Nitrobacter in the absence of dissolved oxygen. Water Res. 1988,22:245-50
Bodelier PLE, Frenzel P. Contribution of methanotrophic and nitrifying bacteria to CH4 and NH4+ oxidation in the rhizosphere of rice plants as determined by new methods of discrimination. Appl. Environ. Microbiol. 1999,65:1826-33
Both GJ, Gerards S, Laanbroek HJ. Kinetics of nitrite oxidation in two Nitrobacter species grown in nitrite-limited chemostats. Arch. Microbiol. 1992, 157:436-41
Brock TD, Madigan MT, Martinko JM, Parker J. Biology of microorganisms. Upper Saddle River NJ, USA. Prentice Hall, 1997
Burns RC, Hardy RWF. Nitrogen fixation in bacteria and higher plants. Berlin, Springer Verlag, 1975 Carpertier B & Cerf O. Biofilms and their consequences, with particular reference to hygiene in the food industry. J Appl. Bacteriol. 1993, 75:499-511
Characklis WG. Laboratory biofilm reactors. In Characklis WG and Marshall KC (Eds.): Biofilms. New York, 1990
Costerton JW, Lewandowski Z, Debeer D. Biofilms, the customized microniche. J. Bacteriol. 1994, 176:2137-42
De Beer D, Vav Der Heuvel JC, Ottenfraf SPP. Microelectrode measurements in nitrifying aggregates. Appl Env Microbiol, 1993, 59: 573-579．
De Boer W, Gunnewiek Pak, Veenhuis M, Bock E, Laanbroek HJ. Nitrification at low pH by aggregated chemolithotrophic bacteria. Appl. Environ. Microbiol. 1991, 57:3600-4
Delong EF, Wickham GS, Pace NR. Phylogenetic strains: ribosomal RNA based probes for the identification of single cells. Science, 1989,243:1360-3
Dolfing J. Acetogenesis. Ecology of Anaerobic Microorganisms, Zehnder (Ed.). Wiley Interscience, 417-468,1988
Ebersold HR, Cordier JL, Luthy P. Bacterial mesosomes: Method dependent artifacts. Arch. Microbiol. 1981, 130: 19-22
Ehrich S, Behrens D, Lebedeva E, Ludwig W, Bock E. A new obligately chemolithoautotrophic, nitrite-oxidizing bacterium, Nitrospira moscoviensis sp nov and its phylogenetic relationship. Arch. Microbiol. 1995,164:16-23
Einsle O, Messerschmidt A, Stach P, Bourenkov GP, Bartunik HD, Huber R, Kroneck PH. Structure of cytochrome c nitrite reductase. Nature. 1999,400:476-80
Etterer T, Wilderer PA. Generation and properties of aerobic granular sludge. Wat Sci Tech, 2001,43(7) :19-26．
Ferguson SJ. Is a proton pumping oxidase essential for energy conservation in Nitrobacter? FEBS Lett.
1982,146:239-43
Ferguson SJ. Denitrification and its control. Antonie van Leeuwenhoek Int. J.Gen. Mol.MicrobioI. 1994, 66:89-110
Ferguson SJ. Nitrogen cycle enzymology. Curr. Opin. Chem. Biol. 1998:182-93
Focht DD, Chang AC. Nitrification and denitrification processes related to wastewater. Adv. Appl. Microbiol, 1975, 19:153-86
Freitag A, Bock E. Energy conservation in Nitrobacter, FEMS Microbiol. Lett. 1990,66:157-62
Gee CS, Pfeffer JT, Suidan MT. Nitrosomonas and Nitrobacter interactions in biological nitrification. J. Environ. Eng. 1990, 116(1) :4-17
Goreau TJ, Kaplan WA, Wofsy SC, McElroy MB, Valois FW, Watson SW. Production of NO2-and N2O by nitrifying bacteria at reduced concentrations of oxygen. Appl. Environ. Microbiol. 1980, 40:526-32
Guiot SR, Pauss A, Costerton WJ. A structured model of the anaerobic granule consortium. Wat. Sci. Tech. 1992,25:1-10
Halling-Sorensen B, Hjuler H. Simultaneous nitrification and denitrification with and upflow fixed bed reactor applying clinoptilolite as media. Wat. Treat, 1992, 7: 77-88
Hanaki K, Hong Z, Matsuo. Production of nitrous oxide gas during denitrification of wastewater. Water Sci. Technol. 1992, 26:1027-36
Harada H, Uemura S, Momonoi K. Interaction between sulfate-reducing bacteria and methane producing bacteria in UASB reactors fed with low strength wastes containing different levels of sulfate. Wat. Res. 1994, 28:355-67
Hellinga C, Schellen AAJC, Mulder JW, Van Loosdrecht MCM, Heijnen JJ. The SHARON process: an innovative method for nitrogen removal from ammonium-rich waste water. Water Sci. Technol. 1998, 37:135-42
Helmer C, Kunst S. Simultaneous nitrification/denitrification in an aerobic biofilm system. Wat. Sci. Tech. 1998, 37(4-5) : 183-877
Hirorns WD, Hastings RC, Head IM, McCarthy AJ, Saunders JR, Pickup RW, Hall GH. Amplification of 16s ribosomal RNA genes of autotrophic ammonia-oxidizing bacteria demonstrates the ubiquity of nitrosospiras in the environment. Microbiology, 1995,141:2793-800
Ho K. Biological nutrient removal in activated sludge processes with low F/M, sludge bulking control. Ph.D. thesis. The University of Queensland, Australia, 1994
Hooper AB, Vannelli T, Bergmann DJ, Arciero DM. Enzymology of the oxidation of ammonia to nitrite by bacteria. J. Gen. Mol. Microbiol., 1997,71:59-67
Janssen LPBM, Warmoeskerken MMCG. Transport phenomena data companion. Delft, Delft University Press, 1987
Juretschko S, Timmermann G, Schmid M, Schleifer KH, Pommereningroser A, Koops HP, Wagner M. Combined molecular and conventional analyses of nitrifying bacterium diversity in activated sludge-nitrosococcus mobilis and nitrospira-like bacteria as dominant populations. Appl. Environ. Microbiol. 1998,64:3042-51
Kester RA, De Boer W, Laanbroek HJ. Production of NO and N2O by pure cultures of nitrifying and denitrifying bacteria during changes in aeration. Appl. Environ. Microbiol. 1997,63:3872-7
Kuenen JG, Robertson LA. Combined nitrification-denitrification processes. FEMS Microbiol. Rev. 1994,15: 109-17
Lettinga G, Hulshoff PLW, Zeeman G. Biological wastewater treatment: Anaerobic Treatment). Department of Environmental Technology, Wageningen Agricultural University, The Netherlands, 1993．
Lewis DL, Gattie DK. Effects of cellular aggregation on the ecology of microorganisms. American Soc. Microbiol. News. 1990,56:263-268
Lloyd H, Ketchum Jr. Design an physical features of sequencing batch reactors. Water Sci. Tech. 1997, 35(1) :11-18
Lipschultz F, Zafiriou OC, Wofsy SC, McElroy MB, Valois FW, Watson SW. Production of NO and N2O by soil nitrifying bacteria. Nature,1981, 294:641-3
Madigan MT, Martinko JM, Parker J. Brock Biology of Microorganisms, Prentice Hall, 473-531, 1997
Masuda S, Watanabe Y, Ishiguro M. Biofilm properties and simultaneous nitrification and denitrification in aerobic rotating biological contactors. Wat. Sci. Tech. 1991, 23(7-9) : 1355-63
Mateju V, Cizinska S, Krejci J. Biological water denitrification-a revies. Enzyme Microb. Technol. 1992, 14:170-183
Mishima K, Makamura M. Self-immovilization of aerobic activated sludge-a pilot study of the aerobic upflow sludge blanket process in municipal sewage treatment. Wat Sci Tech, 1991, 23: 981-990．
Mitchell P. Chemiosmotic coupling and energy transduction. Theor. Exp. Biophys, 1969,2:159-261
Moriyama K, Sato K, Harada Y, Washiyama K and Okamoto K. Simultaneous biologicalremoval of nitrogen and phosphorus using oxic-anaerobic-oxic process. Wat. Sci. Tech. 1990,22(7-8) :61-66
Murray RGE, Watson SW. Structure of nitrocyctis oceanus and comparison with Nitrosomonas and Nitrobacter.J. Bacterial, 1965,89:1594-609
Munch EV, Lant P, Keller J. Simultaneous nitrification and denitrification in bench-scale sequencing batch reactor. Wat. Res. 1996,30(2) : 277-284
Okayasu Y, Abe I, Matsuo Y. Emission of nitrous oxide from high-rate nitrification and denitrification by mixed liquor circulating process and sequencing batch reactor process. Water Sci. Technol. 1997,36:39-45
Oleszkiewicz JA. Granulation in Anaerobic sludge bed reactors treating food industry wastes. Biological waste. 1988
Pachana K, Keller J. Study of factors affecting simultaneous nitrification and denitrification (SND). Wat. Sci. Tech. 1999, 39(6) :600-605
Payne W. Denitrification. New York, Wiley, 1981
Poth M, Focht DD. 15N Kinetic analysis of N2O production by Nitrosomonas europaea: an wxamination of nitrifier denitrification. Appl. Environ. Microbiol, 1985,49:1134-41
Rittmann BE, Langeland WE. Simultaneous denitrification with nitrification in single-channel oxidation ditches. Journal WPCF. 1985, 57(4) :300-308
Robertson LA, Gijs KJ. Aerobic denitrification: a controversy revived. Arch. Microbiol. 1984, 139:351-354
Robertson LA, Kuenen JG. Nitrogen removal from water and waste. In: Microbial control of pollution. Fry JC, Gadd GM, Herbert RA et al. (Eds.). Cambridge University Press, Cambridge, 1992, 227-267
Robertson LA, Van Neil EWJ. Simultaneous denitrification with nitrification in aerobic chemostat cultures of thiophaera pantotropha. Appl. Environ. Microbiol. 1988, 54(1) : 2812-18
Schlegel HG, Bowien B. Autotrophic Bacteria. Berlin, Springer Verlag, 1987
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