嗜热螯台球菌TAD1的脱氮性能及其好氧反硝化分子机制的初步研究
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摘要
目前，活性氮污染在世界范围内正呈现愈演愈烈的趋势，其范围不仅包括水体中的硝氮、亚硝氮及铵态氮等，还包括大气中的NO_x（主要为NO和NO_2）气体，因而如何去除活性氮的污染成为近年来的研究热点。生物法脱氮作为一种低成本且无污染的绿色技术，已引起越来越广泛的关注。然而，根据传统的反硝化理论，氧的存在抑制反硝化的进行，降低了生物脱氮的效率，因此，最初的生物反应器均是在厌氧下进行，这很明显增加了运行成本，进而限制了其大规模的工业化应用。此外，部分工业含氮废水及经过洗涤后的燃煤烟气的温度均达到了50℃左右，而当前的相关生物技术的最佳温度均为常温，因而，开发高温生物脱氮技术具有切实的工程应用价值。本文选用嗜热螯台球菌TAD1（ChelatococcusdaeguensisTAD1）为研究对象，并应用于不同的反应器内，深入分析了TAD1在多种条件下的脱氮性能，初步探讨了TAD1发生好氧反硝化的分子机制，以期为真正地工业应用提供实践与理论指导。
首先，采用PB（Plackett-Burman）设计法和基于中心组合实验设计（CCD）的响应面法，对生物滴滤塔的循环液组分进行了优化。结果表明，对脱氮率影响最大的因子是柠檬酸铵和硫酸亚铁，对实验数据进行二次多项式回归拟合后，获得响应值脱氮率（Y）对自变量柠檬酸铵（X1）和硫酸亚铁（X2）的二元二次回归方程：Y=-2.15+3.13X_1~2+204X_2-0.657X_1~2-1844X_2+5.68X_1X_2，求导后得到模型的极值点，即柠檬酸铵为2.62g/L，硫酸亚铁为0.059g/L，相应的模型预测的最大脱氮率为8.0318mg/(L·h)。
其次，在实验室内建一小型生物滴滤系统，利用优化后的循环液组分，开放条件下以TAD1挂膜后来处理模拟烟气。该系统在不同负荷和氧浓度下均可实现对NO_x的有效去除。在进口NO浓度为600ppm、112.5s的空床停留时间（EBRT）下，NO_x的去除率（RE）可达到80.2–92.3%，氧对RE没有产生负面影响。同时，在系统运行的整个过程中，TAD1始终占有优势地位，说明其具有处理实际烟气的可行性。
第三，分析TAD1在高温下的同步硝化反硝化性能，并将其应用于高温曝气生物滴滤池中。结果表明，无论是在纯种培养还是在曝气生物滴滤池中，TAD1均有良好的异养硝化－好氧反硝化性能。在纯种培养下，铵的去除能力达到6.97mg-NH_4~+-N/(L·h)，约有32.3%的总氮被TAD1转化成氮气；在曝气生物滤池中，12小时后在三种条件下的氮去除效率均达到100%，氮去除能力（平均脱氮率）分别为12.67mg/(L·h)、3.62mg/(L·h)及16.53mg/(L·h)，揭示TAD1具有应用于高温废水脱氮的潜力。
第四，在实验室生物滴滤塔上进一步探索影响脱氮效果的最佳工艺参数及条件，随后在广州电厂一生物滴滤系统内，用TAD1挂膜，以火电厂经过脱硫除尘的烟道气为研究对象。结果显示，在实验室生物滴滤塔内，初始NO-3浓度和空气流速对脱氮没有产生负面影响，综合考虑，乙酸钠为最佳碳源；在电厂生物滴滤系统中，NO_x去除效率可达到84.3-86.7%，最大污染负荷和最大处理能力分别为159.4g/(m~3·h)和137.3g/(m~3·h)，相应的NO_x进口浓度为558mg/m~3。虽然TAD1在生物膜中没有占绝对优势，但与其它多种微生物共存，共同完成处理NO_x的过程。
最后，通过实时荧光定量PCR系统分析四种反硝化基因在不同溶氧下的表达规律，从基因水平初步探讨TAD1在高温下的好氧反硝化发生机制。结果表明，TAD1的硝酸还原酶、亚硝酸还原酶和一氧化氮还原酶的基因类型分别为napA、nirK和cnorB，且含有氧化亚氮还原酶基因nosZ。随着溶氧的提高，napA、nirK、cnorB和nosZ的表达量呈下降趋势，说明高氧浓度对四种反硝化基因有抑制作用，而氧限制或微好氧环境可能更有利于基因的表达。
Recently, the pollution of reactive nitrogen is growing in intensify, which includes notonly NO-3-N, NO-2-N and NH+4-N in the water but also NO_x in the gas. Accordingly, how toremove reactive nitrogen has became a research focus. As a green technology with low costand no pollution, denitrificaion by microbial methods has attracted more and more attention.However, the oxygen inhibits denitrification and decreases removal efficiency according tothe traditionary denitrifying theory. As a result, the initial bioreactor all performed under theanaerobic condition, which obviously increased the operation cost and thus limited itsindustrial application on a large scale. In addition, the temperature of some industrial wastewater containing nitrogen and flue gas after scrub achieves to about50℃, but the presentrelative biotechnology functions well at common temperature. Therefore, it has the value ofengineering application to explore the biological nitrogen removal technology at hightemperature. In this study, thermophilic Chelatococcus daeguensis TAD1was used andinoculated to the different bioreactors. Then, the denitrification performances under differentconditions were deeply analyzed and the molecular mechanism that TAD1denitrified underaerobic condition was preliminarily discussed with the aim to provide practice and theoryguidance for the real industrial application.
First of all, Plackett-Burman design and response surface method based on centralcomposite design were utilized to optimize the medium for recycling liquid of a bio-tricklingfilter. Results showed that the most important factors affecting denitrification rate wereammonium citrate and ferrous sulfate. With the data subject to regression fitting, thefollowing second-order polynomial equation was obtained, depicting relationship between thedenitrification rate (Y) and variables (concentrations of ammonium citrate (X1) and ferroussulfate (X2)): Y=-2.15+3.13X_1~2+204X_2-0.657X_1-1844X_2~2+5.68X_1X_2. On the basis of thisequation, extreme points were: ammonium citrate2.62g/L and ferrous sulfate0.059g/L, andthe corresponding predicted maximum denitrification rate by the model was8.0318mg/(L·h).
Then, a lab-scale bio-trickling filter was set up to treat simulated flule gas after theformation of biofilm with TAD1under the open condition using the optimized recyclingliquid. The filter could remove NO_x efficiently under conditions of different loadings and oxygen concentrations. When the inlet NO concentration was600ppm and EBRT was112.5s,the removal efficiency (RE) of NO_x achieved to80.2–92.3%and oxygen had no negativeeffect on RE. Meanwhile, TAD1predominated in the biofilm all the time during the wholeoperation, indicating that it is feasible to treat the real flue gas by TAD1.
Thirdly, the denitrification performance of simultaneous nitrification and denitrificationby TAD1at high temperature was analyzed, and then it was applied to an aerobic biofilter.Results showed that TAD1exhibited a high performance of heterotrophic nitrification-aerobicdenitrification in both pure culture and aerobic biofilter. In the pure culture, the eliminatecapacity of NH_4~+-N got up to6.97mg/(L·h), and about32.3%of total nitrogen was convertedto nitrogen gas. In the aerobic biofilter, all the nitrogen removal efficiencies of threeconditions after12h reached100%, and the eliminate capacity (average denitrification rate) ofnitrogen was12.67mg/(L·h)、3.62mg/(L·h) and16.53mg/(L·h), respectively, suggesting thatTAD1has a potential for nitrogen removal from wastewater at high temperature.
Fourthly, the best process parameters and conditions affecting denitrification efficiency ina lab-scale bio-trickling filter were further investigated. Thereafter, TAD1was inoculated to apilot-scale bio-trickling filter in a power plant (Guangzhou) using the flue gas afterdesulfuration and dedusting as an object of study. Results revealed that the initial NO_3~-concentration and air flow had no influence on the denitrification, and taken together, sodiumacetate was the optimum carbon source in a lab-scale bio-trickling filter. In a pilot-scalebio-trickling filter, the removal efficiency of NO_x attained to84.3-86.7%. The maximumpollutional loading and processing capacity were159.4g/(m~3·h) and137.3g/(m~3·h),respectively, and the corresponding inlet concentration of NO_x was558mg/m~3. AlthoughTAD1did not predominate in the biofilm, it coexisted with other microorganisms to removeNO_x together.
Finally, expression of four denitrificaiotn genes in TAD1under different levels ofdissolved oxygen (DO) was systematically analyzed by means of real-time fluorescentquantitative PCR, aiming to preliminarily discuss molecular mechanism of aerobicdenitrification by TAD1from gene level at high temperature. Results manifested that genetype of nitrate reductase, nitrite reductase and nitric oxide reductase was napA, nirK andcnorB, respectively, and there was nosZ in TAD1. With DO increasing, expressions of napA, nirK and cnorB all decreased, suggesting that high level of DO exerted an inhibitory effect onexpression of four denitrification genes, whereas oxygen-limited or microaerobic environmentprobably more favored gene expressions.
首先，采用PB（Plackett-Burman）设计法和基于中心组合实验设计（CCD）的响应面法，对生物滴滤塔的循环液组分进行了优化。结果表明，对脱氮率影响最大的因子是柠檬酸铵和硫酸亚铁，对实验数据进行二次多项式回归拟合后，获得响应值脱氮率（Y）对自变量柠檬酸铵（X1）和硫酸亚铁（X2）的二元二次回归方程：Y=-2.15+3.13X_1~2+204X_2-0.657X_1~2-1844X_2+5.68X_1X_2，求导后得到模型的极值点，即柠檬酸铵为2.62g/L，硫酸亚铁为0.059g/L，相应的模型预测的最大脱氮率为8.0318mg/(L·h)。
其次，在实验室内建一小型生物滴滤系统，利用优化后的循环液组分，开放条件下以TAD1挂膜后来处理模拟烟气。该系统在不同负荷和氧浓度下均可实现对NO_x的有效去除。在进口NO浓度为600ppm、112.5s的空床停留时间（EBRT）下，NO_x的去除率（RE）可达到80.2–92.3%，氧对RE没有产生负面影响。同时，在系统运行的整个过程中，TAD1始终占有优势地位，说明其具有处理实际烟气的可行性。
第三，分析TAD1在高温下的同步硝化反硝化性能，并将其应用于高温曝气生物滴滤池中。结果表明，无论是在纯种培养还是在曝气生物滴滤池中，TAD1均有良好的异养硝化－好氧反硝化性能。在纯种培养下，铵的去除能力达到6.97mg-NH_4~+-N/(L·h)，约有32.3%的总氮被TAD1转化成氮气；在曝气生物滤池中，12小时后在三种条件下的氮去除效率均达到100%，氮去除能力（平均脱氮率）分别为12.67mg/(L·h)、3.62mg/(L·h)及16.53mg/(L·h)，揭示TAD1具有应用于高温废水脱氮的潜力。
第四，在实验室生物滴滤塔上进一步探索影响脱氮效果的最佳工艺参数及条件，随后在广州电厂一生物滴滤系统内，用TAD1挂膜，以火电厂经过脱硫除尘的烟道气为研究对象。结果显示，在实验室生物滴滤塔内，初始NO-3浓度和空气流速对脱氮没有产生负面影响，综合考虑，乙酸钠为最佳碳源；在电厂生物滴滤系统中，NO_x去除效率可达到84.3-86.7%，最大污染负荷和最大处理能力分别为159.4g/(m~3·h)和137.3g/(m~3·h)，相应的NO_x进口浓度为558mg/m~3。虽然TAD1在生物膜中没有占绝对优势，但与其它多种微生物共存，共同完成处理NO_x的过程。
最后，通过实时荧光定量PCR系统分析四种反硝化基因在不同溶氧下的表达规律，从基因水平初步探讨TAD1在高温下的好氧反硝化发生机制。结果表明，TAD1的硝酸还原酶、亚硝酸还原酶和一氧化氮还原酶的基因类型分别为napA、nirK和cnorB，且含有氧化亚氮还原酶基因nosZ。随着溶氧的提高，napA、nirK、cnorB和nosZ的表达量呈下降趋势，说明高氧浓度对四种反硝化基因有抑制作用，而氧限制或微好氧环境可能更有利于基因的表达。
Recently, the pollution of reactive nitrogen is growing in intensify, which includes notonly NO-3-N, NO-2-N and NH+4-N in the water but also NO_x in the gas. Accordingly, how toremove reactive nitrogen has became a research focus. As a green technology with low costand no pollution, denitrificaion by microbial methods has attracted more and more attention.However, the oxygen inhibits denitrification and decreases removal efficiency according tothe traditionary denitrifying theory. As a result, the initial bioreactor all performed under theanaerobic condition, which obviously increased the operation cost and thus limited itsindustrial application on a large scale. In addition, the temperature of some industrial wastewater containing nitrogen and flue gas after scrub achieves to about50℃, but the presentrelative biotechnology functions well at common temperature. Therefore, it has the value ofengineering application to explore the biological nitrogen removal technology at hightemperature. In this study, thermophilic Chelatococcus daeguensis TAD1was used andinoculated to the different bioreactors. Then, the denitrification performances under differentconditions were deeply analyzed and the molecular mechanism that TAD1denitrified underaerobic condition was preliminarily discussed with the aim to provide practice and theoryguidance for the real industrial application.
First of all, Plackett-Burman design and response surface method based on centralcomposite design were utilized to optimize the medium for recycling liquid of a bio-tricklingfilter. Results showed that the most important factors affecting denitrification rate wereammonium citrate and ferrous sulfate. With the data subject to regression fitting, thefollowing second-order polynomial equation was obtained, depicting relationship between thedenitrification rate (Y) and variables (concentrations of ammonium citrate (X1) and ferroussulfate (X2)): Y=-2.15+3.13X_1~2+204X_2-0.657X_1-1844X_2~2+5.68X_1X_2. On the basis of thisequation, extreme points were: ammonium citrate2.62g/L and ferrous sulfate0.059g/L, andthe corresponding predicted maximum denitrification rate by the model was8.0318mg/(L·h).
Then, a lab-scale bio-trickling filter was set up to treat simulated flule gas after theformation of biofilm with TAD1under the open condition using the optimized recyclingliquid. The filter could remove NO_x efficiently under conditions of different loadings and oxygen concentrations. When the inlet NO concentration was600ppm and EBRT was112.5s,the removal efficiency (RE) of NO_x achieved to80.2–92.3%and oxygen had no negativeeffect on RE. Meanwhile, TAD1predominated in the biofilm all the time during the wholeoperation, indicating that it is feasible to treat the real flue gas by TAD1.
Thirdly, the denitrification performance of simultaneous nitrification and denitrificationby TAD1at high temperature was analyzed, and then it was applied to an aerobic biofilter.Results showed that TAD1exhibited a high performance of heterotrophic nitrification-aerobicdenitrification in both pure culture and aerobic biofilter. In the pure culture, the eliminatecapacity of NH_4~+-N got up to6.97mg/(L·h), and about32.3%of total nitrogen was convertedto nitrogen gas. In the aerobic biofilter, all the nitrogen removal efficiencies of threeconditions after12h reached100%, and the eliminate capacity (average denitrification rate) ofnitrogen was12.67mg/(L·h)、3.62mg/(L·h) and16.53mg/(L·h), respectively, suggesting thatTAD1has a potential for nitrogen removal from wastewater at high temperature.
Fourthly, the best process parameters and conditions affecting denitrification efficiency ina lab-scale bio-trickling filter were further investigated. Thereafter, TAD1was inoculated to apilot-scale bio-trickling filter in a power plant (Guangzhou) using the flue gas afterdesulfuration and dedusting as an object of study. Results revealed that the initial NO_3~-concentration and air flow had no influence on the denitrification, and taken together, sodiumacetate was the optimum carbon source in a lab-scale bio-trickling filter. In a pilot-scalebio-trickling filter, the removal efficiency of NO_x attained to84.3-86.7%. The maximumpollutional loading and processing capacity were159.4g/(m~3·h) and137.3g/(m~3·h),respectively, and the corresponding inlet concentration of NO_x was558mg/m~3. AlthoughTAD1did not predominate in the biofilm, it coexisted with other microorganisms to removeNO_x together.
Finally, expression of four denitrificaiotn genes in TAD1under different levels ofdissolved oxygen (DO) was systematically analyzed by means of real-time fluorescentquantitative PCR, aiming to preliminarily discuss molecular mechanism of aerobicdenitrification by TAD1from gene level at high temperature. Results manifested that genetype of nitrate reductase, nitrite reductase and nitric oxide reductase was napA, nirK andcnorB, respectively, and there was nosZ in TAD1. With DO increasing, expressions of napA, nirK and cnorB all decreased, suggesting that high level of DO exerted an inhibitory effect onexpression of four denitrification genes, whereas oxygen-limited or microaerobic environmentprobably more favored gene expressions.
引文
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