Waste excretion characteristics of Manila clams (Tapes philippinarum) under different temperature conditions

Water recirculating systems have been used in the shellfish industry for depuration and wet-storage. Knowledge of shellfish excretion characteristics is critical to recirculating system design. In this study, the excretion rate of total ammonia nitrogen (TAN), total Kjeldahl nitrogen (TKN), and 5-day biochemical oxygen demand (BOD₅) from Manila clams (Tapes philippinarum) were investigated under both laboratory and commercial conditions. The laboratory tests were conducted under temperatures ranging from 3 to 30°C. The experimental results showed that temperature was a key factor in determining the excretion rate of all the above parameters. The relationship between TAN excretion rate (R_TAN) and temperature (T) can be represented by an exponential function (R_TAN=0.57×1.25^T). For the temperature range between 3 and 20°C, the daily mean excretion rates of TAN, TKN and BOD₅ ranged between 1.5–46.1, 4.8–131.0 and 57.4–219.4 mg per kilogram of the clams (wet weight with shell on), respectively. There were linear correlations between TAN, TKN and BOD₅ production rates. The data presented in this paper can be used to estimate waste generation from a given shellfish processing operation and to size the waste treatment components for a recirculating depuration (or wet-storage) system.     

Root nitrification capacity of lettuce plants with application to aquaponics

Aquaculture and hydroponics have experienced significant growth and market presence in recent years. While aquaponics, the combination of fish and plant culture systems, is beginning to experience the same exponential growth and interest that hydroponics did many years ago, very little information is available on sizing and design of these systems. Incorporation of hydroponic plants with recirculating aquaculture systems (RAS) aids in removal of ammonia/ammonium based wastes, thus reducing the need for water discharge to control water quality. Surface only nitrification rates were quantified to be 0.83g/m²/day for inert surfaces and 0.20/m²/day for root surfaces. Direct assimilation of ammonia by the lettuce plants was less than 2% of the total ammonia and ammonium nitrogen (TAN) removed from the culture water, with the remainder being removal by oxidation of TAN into nitrate.     

Performance of sequencing microbead biofilters in a recirculating aquaculture system

Biological filtration, or biofiltration, is the key technology in recirculating aquaculture systems. Sequencing microbead biofilters, in which the media maintains a continuous up-and-down movement, are based on traditional microbead filters but offer superior filtration properties. The performance characteristics of a sequencing microbead biofilter installed in a recirculating aquaculture system for rearing Barcoo perch at 29 ± 1 °C were examined. The total ammonia-nitrogen (TAN) concentrations and the nitrite-nitrogen concentrations during a 52-day culture period were maintained blow 1.6 mg/L and 0.9 mg/L. In order to ensure efficient biofiltration, the optimal actual application of hydraulic retention time was determined to be approximately 3–5 min. The water flow produced by the reciprocating motion of the media served to wash away suspended solids, ensuring the occurrence of optimal nitrification processes. Additionally, the reciprocating motion of the media enhanced ammonia treatment efficiency significantly by improving the transport of nutrients and nitrification activity. Compared to a static situation the ammonia removal rate increased by 27% based on the application of up-and-down reciprocating movement. The biofilm on the microbead forms as a compact, complex, and homogeneous structure, consisting of numerous microscopic thin sheets. Additionally, a multitude of pores, interstitial voids, and vertical channels were widely observed to convey obviously advantageous properties in support of fluid passage, thus enhancing mass transfer and ultimately contributing to biofiltration effectiveness. The optimum biofilm thickness for providing efficient biofiltration was determined to be approximately 70 μm for this filter.     

Potential applications of indirect electrochemical ammonia oxidation within the operation of freshwater and saline-water recirculating aquaculture systems

The paper addresses two potential applications for electrochemical ammonia oxidation within the operation of recirculating aquaculture systems, in which nearly complete removal of N species is required. In one described application, a physical–chemical ammonia oxidation method is suggested to entirely replace conventional biological treatment methods (i.e. nitrification/denitrification). The second described method is suggested as a final polishing step for removing ammonia from effluents of denitrification reactors supplied with intrinsic organic matter, prior to the discharge of the water. Empirical results and cost assessment are reported for the second alternative, while the first, which was recently published, is discussed with respect to improvements, operational conditions and field tests required to induce its commercial application. The polishing alternative was shown capable of efficiently removing TAN in the effluents of RAS denitrification reactors fed with intrinsic organic solids. The cost for treating denitrification reactor effluents with TAN concentration of 10 mgN/L was estimated at 6.67 cent/m³ of discharged water. Since the chloride ion concentration in seawater and in most brackish waters is high, combining the intrinsic organic carbon denitrification process with subsequent ammonia polishing by electrochemically produced active chlorine may be a competitive approach for the removal of nitrogen species from seawater and brackish water RAS.     

生物—电氧化法去除海水养殖循环水污染物

为提高海水养殖循环水处理效率,降低处理成本,本研究采用曝气生物滤器与电化学阳极氧化组合工艺,考察了不同阳极电势、进水氨氮和亚硝酸盐浓度下系统对氨氮及亚硝酸盐等污染物的去除效果,研究了微生物与工作电极之间的相互作用,并分析了电化学反应能耗。在水力停留时间为45 min、1.4 V阳极电压、进水氨氮和亚硝酸盐浓度分别为4.5和1.3 mg/L条件下,生物—电氧化法对氨氮去除率达88.8%,高出对照组7.6%,出水氨氮和亚硝酸盐浓度分别为0.5和0.9 mg/L,COD去除率为88.2%,高出对照组19.4%,平均能耗0.040 kWh/m~3,电极表面微生物生长对阳极电氧化过程有促进作用,微生物功能预测显示实验组硝化功能占比为0.03%,对照组为0.07%。研究表明,生物—电氧化法对海水养殖循环水的污染物有良好的去除效果,具有一定的发展应用潜力。     

不同浓度臭氧对循环水养殖系统生物膜活性及其净化效能的影响

本文通过在循环水养殖系统中添加不同浓度的臭氧,研究其对循环水养殖系统生物膜活性及其净化效能的影响.结果显示,当氧化还原电位(ORP)小于450 mV时,氨氮的去除率随着臭氧浓度升高而升高,最高去除率达39.9％,亚硝酸盐氮的平均去除率为28.2％,生物膜菌群的平均存活率为88.1％,生物膜对养殖水体氨氮和亚硝酸盐氮的处理效果良好;当氧化还原电位为500 mV时,经过臭氧24 h处理,氨氮和亚硝酸盐氮的去除率分别由36.5％、28.1％降到12.2％、8.4％,而臭氧4h处理后,生物膜对氨氮和亚硝酸盐氮的去除率分别由47.5％、32.1％降到5.0％、3.3％,水处理效果明显下降,生物膜菌群存活率由88.1％降到31.5％.由此可见臭氧添加浓度对生物膜及净化效能有重大影响.综合试验结果和分析评估,建议封闭循环水养殖系统的臭氧添加量以控制生物滤池内的氧化还原电位低于400 mV为宜,可保证循环水系统的安全性和经济性.     

Performance characteristics of fluidised bed biofilters in a novel laboratory-scale recirculation system for rainbow trout: nitrification rates, oxygen consumption and sludge collection

A laboratory-scale recirculating aquaculture system for fluidised bed biofilter evaluation was engineered. The design included all components found in typical full-scale commercial production systems. The system included two identical units each with oxygenation, UV treatment, cooling, biofiltration and a particulates separation device. Water from the two systems was mixed in a degassing unit. A 1 month test period after biofilter maturation revealed stable concentrations of total ammonia nitrogen (TAN), nitrite and nitrate within the system. Mean nitrification rate was 0.27 and 0.21 g TAN m⁻² day⁻¹. Oxygen consumption in the biofilters ranged between 56 and 64% due to nitrifying activity. Mass balances on nitrogen indicated that 48%, added via the feed, was converted to nitrate within the system, with 6% of the added nitrogen being found in the sludge. The remaining 43% was either used during fish growth, left the system, as organic nitrogenous compounds (or unidentified nitrogenous compounds), via the outlet, or was lost to the atmosphere. At least 61% of the nitrate produced was generated by the biofilters. The system proved to be an exceptional set-up for evaluation of the performance of fluidised bed biofilters, allowing both pre- and post-filter measurements of various water quality criteria.     

生物强化移动床生物膜反应器处理水产养殖循环水初步研究

利用自制的硝化细菌菌剂促进移动床生物膜反应器(Moving bed biofilm reactor,MBBR)的挂膜启动,分析不同载体氨氮负荷、碳氮比条件下反应器运行状况,并进一步进行了实验室模拟循环水养殖草金鱼实验。结果显示,利用自制硝化菌剂能够完成整个移动床反应器的启动过程,在接种15 d后使循环出水氨氮稳定在1 mg/L以下。单位体积载体氨氮负荷实验表明,MBBR能够在100 mg TAN/(L填料·d)条件下,使出水满足一般水产养殖水质要求(氨氮0.5 mg/L,亚硝氮0.1 mg/L)。进水碳氮比在1以内时MBBR能够稳定高效运行。在实验室模拟循环水养殖过程中,经菌剂强化的MBBR能维持循环出水氨氮低于0.5 mg/L,亚硝氮低于0.05 mg/L。     

封闭循环水系统生物滤池气水比对水质净化效能的影响

通过在全封闭循环水系统中养殖半滑舌鳎(Cynoglossus semilaevis Gunthe), 研究了不同气水比对曝气生物滤池净化效能, 以及对DO、pH值的影响。结果表明: 本试验系统在温度为(19±1)℃, 系统循环次数为15次, 养殖池DO保持在12 mg/L以上的运行条件下, 随着气水比由0.75 : 1~1.50 : 1的增加, 生物滤池氨氮的去除率由35.0%增加至52.0%, NO₂-N的去除率由8.2%增加至44.6%, 气水比对硝化反应影响显著, 但对化学需氧量COD的去除率影响并不显著, 其平均去除率为10.14%; pH值有增加的趋势, 生物滤池进水口到出水口的pH值由7.97增加至 8.08; 气水比最佳运行参数为1.25:1。同时还发现1级生物滤池进水口DO接近饱和, 1级到末级滤池间DO仅降低了10%左右, 系统pH值在7.9~8.1。本研究所获参数, 可供生物膜法处理养殖循环水的条件优化作参考。

     

Assessment of AquaMats for removing ammonia in intensive commercial Pacific white shrimp Litopenaeus vannamei aquaculture systems

AquaMats are high surface–area polymer filters whose use produces higher yields with reduced health risks for the aquaculture product. We used AquaMats in pilot-scale systems and in intensive commercial Pacific white shrimp Litopenaeus vannamei production systems to stabilize and improve water quality by removing ammonia. In the pilot-scale systems, evaluation of the effects of temperature and hydraulic retention time (HRT) on ammonia removal rate indicated that the surface total ammonia nitrogen (TAN) conversion rate (STR, mg TAN/m²-day) increased with increasing temperature and decreasing HRT. The highest STR of 319.8 mg TAN/m²-day was observed at a temperature of 30 °C and a HRT of 5 min. In the commercial shrimp production systems, ammonia levels were significantly greater in the control systems (without AquaMats) than in the treatment systems (with AquaMats) after 6 days (P < 0.05). Results suggested that eight 150 cm × 90 cm pieces of AquaMats (0.057 m² surface area per m³ culture volume) were sufficient for promoting nitrification in this system. The growth rate of juvenile shrimp was most enhanced in treatment C (with 12 pieces of AquaMats, 0.085 m²/m³), which exhibited a significant decrease in ammonia.     

Effects of body weight, water temperature and ration size on ammonia excretion by the areolated grouper (Epinephelus areolatus) and mangrove snapper (Lutjanus argentimaculatus)

The effects of body weight, water temperature and ration size on ammonia excretion rates of the areolated grouper Epinephelus areolatus and the mangrove snapper Lutjanus argentimaculatus were investigated. Under given experimental conditions, L. argentimaculatus had a higher weight-specific ammonia excretion rate than E. areolatus. Weight-specific ammonia excretion rates of fasted individuals of both species showed an inverse relationship with body weight (W, g wet wt.), but a positive relationship with water temperature (t, °C). The relationships for total ammonia nitrogen (TAN) were: E. areolatus: TAN (mg N kg⁻¹ d⁻¹)=21.4·exp^0.11t·W^−0.43 (r²=0.919, n=60); L. argentimaculatus: TAN (mg N kg⁻¹ d⁻¹)=121.5·exp^0.12t·W^−0.55 (r²=0.931, n=60). Following feeding, the weight-specific ammonia excretion rate of E. areolatus increased, peaked at 2 to 12 h (depending on temperature), and returned to pre-feeding levels within 24 h. A similar pattern was observed for L. argentimaculatus, with a peak of TAN excretion being found 6 to 12 h after feeding. Stepwise multiple regression analysis indicated that weight-specific TAN excretion rates of both species increased with increasing temperature and ration (R, percent body wt. d⁻¹): E. areolatus: TAN (mg N kg⁻¹ d⁻¹)=22.8·t−28.8·R−378.2 (r²=0.832, n=24); L. argentimaculatus: TAN (mg N kg⁻¹ d⁻¹)=22.9·t−25.4·R−216.4 (r²=0.611, n=24). The effect of body weight on weight-specific postprandial TAN excretion was not significant in either species (p>0.05). This study provides empirical data for estimating ammonia excretion of these two species under varying conditions. This has application for culture management.     

Removal of nitrogen by Algal Turf Scrubber Technology in recirculating aquaculture system

Ongoing research in recirculation aquaculture focuses on evaluating and improving the purification potential of different types of filters. Algal Turf Scrubber (ATS) are special as they combine sedimentation and biofiltration. An ATS was subjected to high nutrient loads of catfish effluent to examine the effect of total suspended solids (TSS), sludge accumulation and nutrient loading rate on total ammonia nitrogen (TAN), nitrite and nitrate removal. Nutrient removal rates were not affected at TSS concentration of up to 0.08 g L^?1 (P > 0.05). TAN removal rate was higher (0.656 ± 0.088 g m^?² day^?1 TAN) in young biofilm than (0.302 ± 0.098 g m^?² day^?1 TAN) in mature biofilm at loading rates of 3.81 and 3.76 g m^?² day^?1 TAN (P < 0.05), respectively, which were considered close to maximum loading. TAN removal increased with TAN loading, which increased with hydraulic loading rate. There was no significant difference in removal rate for both nitrite and nitrate between young and mature biofilms (P > 0.05). The ATS ably removed nitrogen at high rates from catfish effluent at high loading rates. ATS‐based nitrogen removal exhibits high potential for use with high feed loads in intensive aquaculture.     

生物膜法SBR（BSBR）在循环养殖水处理中影响因素分析

膜法SBR（sequence batch reactor）是将SBR法与接触氧化法相结合的一种新型生物膜法处理工艺。此研究以总氨氮（TAN）及总氮（TN）的去除反应速度作为考察指标，分析生物膜法SBR（biofilm sequence batch reactor，BSBR）处理水产循环养殖系统水体中影响TAN及TN去除效果的主要因素。其中，pH和碱度对硝化反应有很大的影响，pH控制在6．3以上时TAN处理效果较好。溶解氧（DO）对反硝化反应也有较大的影响，同时考虑到水生生物的生长需求，在此试验系统中进入反应器的水体DO最好能控制在4．5～6．5mg·L^-1。水温保持在20％左右，可以保证有一个较好的脱氮效果。     

Effects of abrupt salinity increase on nitrification processes in a freshwater moving bed biofilter

The nitrification process is a widely used biological approach responsible for ammonia and nitrite removal in recirculating aquaculture system (RAS) biofilters. Given this pivotal role, the influence of different water quality parameter on nitrification efficiency is important information for RAS operations. One influencing parameter is salinity, and salinity fluctuations in freshwater RAS biofilters are reported to affect the nitrifying bacteria. This study investigated the effects of abrupt increase in salinity in freshwater RAS on substrate-dependent (1’-order) as well as substrate independent (0’-order) nitrification rates. A 100% inhibition was found for surface specific removal (STR) of total ammonia nitrogen (TAN) and surface specific nitrite removal (SNR) when salinity was abruptly increased to 25‰ and above. A fast turnover (i.e. steep decline in [NH₄-N⁺] and [NO₂-N⁻]) were observed at lower salinities (≤10‰), while limited/no degradation of either ammonia or nitrite was seen at salinities above 25‰. At low substrate loading (1’-order process), removal rate constants (k_1a) of 0.22 and 0.23 m d^-1 were observed for ammonia and nitrite degradation, respectively, declining to 0.01 m d^-1when adding marine RAS water increasing the salinity to 15‰. Similar observations followed at high nutrient loadings (0’-order process) with STR and SNR of 0.10 and 0.12 g N m^-2 d^-1, respectively, declining to 0.01 g N m^-2 d^-1 at 15‰. When salinities of 25‰ and 35‰ were applied, neither TAN nor nitrite degradation was seen. The results thus demonstrate a pronounced effect of salinity changes when freshwater RAS biofilters are subjected to fast/abrupt changes in salinity. RAS facility operators should be aware of such potential effects and take relevant precautions.     

Variability of the nitrifying bacteria in the biofilm and water column of a recirculating aquaculture system for tilapia (Oreochromis niloticus) production

The aim of this study was to evaluate variability of nitrifying bacterial community in the biofilm and in the water of a recirculating aquaculture systems (RAS) in a tilapia farming in order to determine if nitrification process is dependent, or not, of nitrifying bacteria abundance. Biofilm and water samples were collected periodically for 30 days and analysed with the fluorescent in situ hybridization (FISH) technique, used to quantify ammonia‐oxidizing bacteria (AOB) and nitrite‐oxidizing bacteria (NOB). Ammonia presented the peak in the first week, while the nitrite's maximum was recorded in the second week. Nitrate increased steadily, indicating nitrification activity. Total bacterial abundance in biofilm increased continuously, while in water, it did not change significantly. In the biofilm, number of AOB was high at beginning, decreased after few days and increased again following augment of ammonia. Number of NOB also showed an increase in abundance in biofilm following the increment of nitrite and nitrate. In water, AOB and NOB did not show major variability. Relative abundance of nitrifying bacteria represented more than 30% of total bacteria in biofilm at beginning of the experiment. Their contribution decreased to >3% in last days. It indicates that nitrifying bacteria are biofilm colonizers, and that their activity seems to be directly related to the concentration of nitrogen compounds. However, contribution of nitrifying bacteria did not vary much along the time. We may conclude that the biofilm‐nitrifying bacteria plays major role in nitrification process in RAS and that the activity of these organisms is dependent of their abundance in response to the concentration of nitrogen compounds.     

Effect of particulate organic carbon on heterotrophic bacterial populations and nitrification efficiency in biological filters

Competition between heterotrophic and nitrifying bacteria is of major practical importance in aquaculture biofilter design and operation. This competition must be understood to minimize the negative impact of heterotrophic bacteria on an aquaculture system. On the other hand, the heterotrophic population is suspected of having a positive effect against pathogenic bacteria. Little information is available on the bacterial communities present within aquaculture systems, except for nitrifying bacteria, but a combination of traditional aquacultural engineering research methods and novel microbiological techniques offers new opportunities for the study of these communities.

The heterotrophic bacterial population activity and the nitrification efficiency of a submerged biological filter were studied for an influent TAN concentration of 2 mg/l and varying C/N ratios. The TAN removal rate was found to be 30% lower at a C/N ratio of 0.5 than at a C/N ratio of 0. For higher C/N ratios the reduction in nitrification efficiency was 50% while the attached bacterial abundance was doubled. Moreover, results confirm that abundance of sheared and attached bacteria are correlated. It is not known to what extent biofilter configuration might influence the relationship between heterotrophic and nitrifying bacteria, and further work will be carried out with moving bed and fluidized filters. A better understanding of the role of the heterotrophic bacteria in RAS will help to optimize any positive “biocontrol” effect and to minimize the microbial degradation of rearing water and the reduction of nitrification rates.     

Effects of formaldehyde on nitrification in biofilters of small‐scale recirculating systems

Florfenicol (Aquaflor®) is the only U.S. Food and Drug Administration (FDA) approved drug for treating diseased fish reared in recirculating aquaculture systems (RAS). Treating diseased fish in RAS is challenging because of the potential to damage nitrifying bacteria in the biofilters. Impaired nitrification can lead to concentrations of ammonia and nitrite that compromise fish welfare. The objective of this study was to determine the effects of a FDA‐approved parasiticide and fungicide, Parasite‐S^® (formalin), on biofilter nitrification. Stable biofilters were exposed once to 0, 9.25, 18.5, 37, or 55.5 mg/L formaldehyde. Total ammonia nitrogen (TAN) and nitrite nitrogen were monitored daily before and throughout the study to quantify biofilter function. Formaldehyde concentrations ≥37 mg/L increased TAN and nitrite nitrogen concentrations, and nitrification did not recover to pre‐exposure concentrations up to 8 day postexposure. On the basis of those results, a second trial was conducted. Stable biofilters were exposed once or on four consecutive days to 9.25 or 18.5 mg/L formaldehyde. Biofilters repeatedly exposed to formaldehyde showed signs of impairment and had variable recovery relative to single exposures. Results of this study may help identify formaldehyde concentrations that can be safely applied to RAS when treating diseased fish.     

An Analytical Model for Predicting the Carrying Capacity of Submerged Biofilters Used in Aquaculture

Laboratory data and information from the literature were used to develop an equation that models ammonia removal by submerged biofilters. The equation is based on the half-order/zero-order kinetics model and fixed biofilm nitrification, which indicates that the nitrification rate is a linear function of the ammonia concentration at ammonia concentrations <2.0 mg/l. Input data for the equation include easily gathered information of flow rate, biofilter size, maximum permissible ammonia level, biofilter influent and biofilter effluent ammonia concentrations, temperature, and ammonia production rate. The equation can be used to determine the carrying capacity of the submerged biofilters, to estimate biofilter sizes needed to support various fish loads, and to investigate the effects of changes in biofilter system parameters on carrying capacity. Comparisons of the predicted and actual carrying capacities of several experimental biofilters indicated that the equation accurately predicted carrying capacity under most conditions. Studies of the sensitivity of the equation to changes in system parameters demonstrated that flow rate limits biofilter performance and suggested an approach for determining cost-effective flow rates for biofilter operation.     

Performance and operation of a rotating biological contactor in a tilapia recirculating aquaculture system

This paper describes the performance characteristics of an industrial-scale air-driven rotating biological contactor (RBC) installed in a recirculating aquaculture system (RAS) rearing tilapia at 28 °C. This three-staged RBC system was configured with stages 1 and 2 possessing approximately the same total surface area and stage 3 having approximately 25% smaller. The total surface area provided by the RBC equaled 13,380 m². Ammonia removal efficiency averaged 31.5% per pass for all systems examined, which equated to an average (± standard deviation) total ammonia nitrogen (TAN) areal removal rate of 0.43 ± 0.16 g/m²/day. First-order ammonia removal rate (K₁) constants for stages 1–3 were 2.4, 1.5, and 3.0 h⁻¹, respectively. The nitrite first-order rate constants (K₂) were higher, averaging 16.2 h⁻¹ for stage 1, 7.7 h⁻¹ for stage 2, and 9.0 h⁻¹ stage 3. Dissolved organic carbon (DOC) levels decreased an averaged 6.6% per pass across the RBC. Concurrently, increasing influent DOC concentrations decreased ammonia removal efficiency. With respect to dissolved gas conditioning, the RBC system reduced carbon dioxide concentrations approximately 39% as the water flowed through the vessel. The cumulative feed burden – describes the mass of food delivered to the system per unit volume of freshwater added to the system daily – ranged between 5.5 and 7.3 kg feed/m³ of freshwater; however, there was no detectable relationship between the feed loading rate and ammonia oxidation performance.     

循环海水养殖系统硝化滤器中氨氧化微生物分析

研究循环水养殖硝化滤器载体上附着生物膜的微生物群落结构可以为提高其处理速率和效率,并为特异性工程菌构建提供依据。采用改良的AFLP方法分析了循环水养殖硝化滤器载体上附着的氨氧化细菌16S rRNA基因和氨单加氧酶amoA基因片段及其系统发育情况。结果表明:分析16S rRNA基因得到的序列片段比分析amoA基因片段得到了更多信息,准确度较高,可作为分析循环水养殖硝化滤器氨氧化菌群组成的有效方法。克隆测序所得序列与网上公布数据比对,可见存在于循环水养殖硝化滤器载体上的氨氧化细菌与Nitrosomonas cryotolerans、Nitrosomonas oligotropha、Nitrosospira tenuis、Nitrosomonas marina相似度达100%,与Nitrosomonas aestuarii相似度为87%。大部分属于亚硝化单胞菌属(Nitrosomonas),仅少数序列属于亚硝化螺菌属(Nitrosospira)。采用16S rRNA基因和amoA片段分析方法得到的附着于封闭循环海水养殖硝化滤器载体上的氨氧化细菌主要为变形菌(Proteobacteria)的β-亚类的亚硝化单胞菌属(Nitrosomonas)和少量的亚硝化螺菌属(Nitrosospira)氨氧化细菌,以及一定数量的γ-亚类氨氧化细菌。