Research progress on polycyclic aromatic hydrocarbons degrading bacteria and their applications
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摘要:
多环芳烃(PAHs)在环境中分布广泛,且具有生态和环境毒理效应,因此对PAHs污染场地的治理和修复备受关注。生物降解是去除PAHs的重要技术之一,但存在降解效率低、周期长等局限性。归纳了PAHs常见降解菌及其主要降解机制,探讨了PAHs降解菌在实际污染场地应用的研究进展与不足。结果表明:PAHs降解菌株主要包括不动杆菌属(Acinetobacter)、分枝杆菌属(Mycobacterium)和假单胞菌属(Pseudomonas),白腐真菌是常见的降解菌;相比单一菌株,复合菌群对PAHs的降解能力更强。在降解菌株降解基因(如nah基因簇)编码酶的作用下,萘、菲和芘等PAHs发生开环并逐步氧化,最终通过水杨酸或邻苯二甲酸途径进入三羧酸循环实现完全降解;而苯并[a]芘降解过程中会产生包括醇、醛、酸类中间产物,其完全降解机理仍有待研究。目前大部分针对PAHs降解菌的研究局限于实验室条件,缺少实际PAHs污染场地降解性能的验证;实际应用中,降解菌活性和PAHs的去除受温度、pH、氧气浓度和土壤有机质含量等环境因子的影响。PAHs降解菌的应用实例包括采用生物刺激和(或)生物强化的方式以促进PAHs污染场地的修复。然而,生物降解在实际应用中仍需克服降解菌失活、技术耦合困难、环境风险和成本高等限制因素。未来研究主要包括复合污染和土著菌共存条件下PAHs生物降解机制研究、降解菌生理特性调控和新型强化材料的开发;此外,应加强降解菌在实际污染场地应用的推广,以实现对PAHs污染的高效、经济、可持续治理。
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关键词:
- 多环芳烃(PAHs) /
- 降解菌 /
- 降解机制 /
- 实际应用 /
- 增强降解
Abstract:Polycyclic aromatic hydrocarbons (PAHs) are a group of pollutants widely distributed in the environment and have ecological and environmental toxicity effects. Therefore, the remediation and restoration of PAHs-contaminated sites have received significant attention. Biodegradation is one of the essential technologies for removing PAHs; however, it still faces limitations such as low degradation efficiency and long degradation periods. The common PAHs-degrading bacteria and their degradation mechanisms were summarized, focusing on discussing the research progress and limitations of applying them to real contaminated sites. The results showed that PAHs-degrading bacteria mainly included genus Acinetobacter, Mycobacterium, and Pseudomonas. White-rot fungi were common fungi that degraded PAHs. Compared to individual strains, bacterial consortia exhibited superior PAH degradation capability. For PAHs such as naphthalene, phenanthrene and pyrene, the degradation process involved ring opening catalyzed by enzymes encoded by PAHs degradation genes (e.g., nah gene cluster), followed by stepwise oxidation, ultimately leading to complete degradation through the salicylic acid or phthalic acid pathway entering the tricarboxylic acid cycle. The degradation of benzo[a]pyrene produced intermediate products, including alcohols, aldehydes, and acids. However, its complete degradation pathway was yet to be identified. Studies on PAHs degradation bacteria were mainly confined to laboratory conditions, and there was a lack of verification in real contaminated soils. In application, the activity of degrading bacteria and the efficiency of PAHs removal were influenced by various environmental factors, including temperature, pH, oxygen levels, and soil organic matter content. In addition, some cases utilized biological stimulation and/or bioaugmentation to significantly improve the bioremediation of PAH-contaminated sites. Nevertheless, the application must overcome multiple limiting factors, including reduced degrading bacteria activity, failed integration with multiple technologies, and high environmental risks and costs. Further researches should include the mechanisms of PAHs biodegradation under conditions with combined pollution and the presence of indigenous microorganisms, the regulation of physiological characteristics of degrading bacteria, and the development of novel materials. Furthermore, promoting the application of PAHs-degrading bacteria in real contaminated sites should be strengthened to achieve efficient, economical, and sustainable control of PAHs contamination
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表 1 PAHs降解菌株及其降解率
Table 1. PAHs degrading bacteria strains and degradation efficiencies
菌株属名 PAHs(降解时间,降解率) 培养介质 数据来源 微杆菌(Microbacterium) 萘(45 d,85%);菲(45 d,约60%);芴(45 d,78%);
苯并[b]荧蒽(45 d,约35%)液体培养基 文献[15] 金黄杆菌(Chryseobacterium) 菲(4 d,100%) 基础盐培养基 文献[16] 枝芽孢杆菌(Virgibacillus) 菲(10 d,94%) 缺氧矿物培养基 文献[17] 肠杆菌(Enterobacter) 芘(120 h,42%~77%) 选择性培养基 文献[18] 新鞘氨醇菌(Novosphingobium) 芘(12 d,<10%) 无碳矿物培养基 文献[13] 骨干杆菌(Diaphorobacter) 苯并[a]芘(52 h,96%) 基础盐培养基 文献[12] 伯克霍尔德氏菌(Paraburkholderia) 二苯并噻吩(30 d,100%) 基本培养基 文献[19] 不动杆菌(Acinetobacter) 菲、芴、苊、荧蒽、芘(7 d,>90%) 土壤 文献[20] 芽胞杆菌(Bacillus) 芘(12 d,约20%) 无碳矿物培养基 文献[13] 菲、苯并[a]荧蒽、芘(56 d,99%) 焦化厂土壤 文献[21] 分枝杆菌(Mycobacterium) 芘(7 d,11%~86%) 矿物培养基 文献[22] 芘(6 d,约100%) 无碳矿物培养基 文献[13] 菲、芘、苯并[a]荧蒽、苯并[a]芘、䓛(150 d,∑PAHs为76%) 液体培养基 文献[23] 苍白杆菌(Ochrobactrum) 芘(12 d,<10%) 无碳矿物培养基 文献[13] 萘(45 d,约70%);菲(45 d,约60%);芴(45 d,约40%);
苯并[b]荧蒽(45 d,约50%)液体培养基 文献[15] 厄氏菌(Oerskovia) 萘(30 d,69%);苊(30 d,47%) 液体矿物培养基 文献[24] 分枝菌酸杆形菌(Mycolicibacterium) 菲(3 d,100%);荧蒽(7 d,100%);芘(3 d,99%) 培养基 文献[25] 红球菌(Rhodococcus) 菲(3 d,23%);芴(7 d,100%);荧蒽(14 d,27%) 培养基 文献[25] 寡养单胞菌(Stenotrophomonas) 萘(45 d,约60%);菲(45 d,约50%);芴(45 d,48%);
苯并[b]荧蒽(45 d,约50%)液体培养基 文献[15] 假单胞菌(Pseudomonas) 萘(45 d,约70%);菲(45 d,40%~60%);芴(45 d,约40%);
苯并[b]荧蒽(45 d,30%~60%)液体培养基 文献[15] 萘(96 h,100%);芴(72 h,40%);二苯并呋喃(96 h,约66%);二苯并噻吩(96 h,约32%) 基础盐培养基 文献[14] 解氢芽胞杆菌(Hydrogenibacillus) 萘(20 h,100%);菲(20 h,20%);芴(20 h,约95%);二苯并呋喃(20 d,100%);二苯并噻吩(20 d,约70%);咔唑(20 d,约30%) 培养基 文献[26] 表 2 PAHs降解菌群及其降解率
Table 2. PAHs degrading consortia and degradation efficiencies
菌株属名 PAHs(降解时间,降解率) 培养介质 数据来源 分枝杆菌(Mycobacterium)、梭状芽胞菌
(Clostridium)菲(7 d,82%) 地下水 文献[34] 芽胞杆菌(Bacillus)、分枝杆菌
(Mycobacterium)、苍白杆菌(Ochrobactrum)、
新鞘氨醇菌(Novosphingobium)芘(6 d,100%) 无碳矿物
培养基文献[13] 戴氏菌(Dyella)、索氏菌(Thauera)、地杆菌
(Geobacter)、泰氏菌 (Tissierella)、骨干杆菌
(Diaphorobacter)苯并[a]芘(30 d,11%~20%) 焦化废水 文献[35] 分枝杆菌(Mycobacterium)、鞘氨醇单胞
(Sphingomonas)菲(3 d,100%);荧蒽(3 d,71.2%);芘(3 d,50%) 基础盐培养基 文献[32] 脱硫弧菌(Desulfovibrio)、佩特里单胞菌
(Petrimonas)菲、芴、苊、蒽、芘、苯并[a]荧蒽、苯并[b]荧蒽、苯并[k]荧蒽、苯并[a]芘、
二苯并[a,h]蒽(∑PAH,55 d,21%~29%)矿物盐培养基 文献[36] 鞘氨醇菌(Sphingobium)、假单胞菌
(Pseudomonas)萘(8 h,约85%);菲(60 h,100%);芴(16 h,约80%);苊(30 h,约25%);
蒽(30 h,约100%);荧蒽(8 h,100%);二苯并[a]荧蒽(30 h,约60%);
二苯并呋喃(30 h,约50%);二苯并噻吩(60 h,94%);咔唑(30 h,约20%)基础盐培养基 文献[33] 微杆菌(Microbacterium)、苍白杆菌
(Ochrobactrum)、假单胞菌(Pseudomonas)、
寡养单胞菌(Stenotrophomonas)萘(45 d,97%);菲(45 d,97%);芴(45 d,76%);
苯并[b]荧蒽(45 d,73%)液体培养基 文献[15] 表 3 PAHs降解菌、降解基因和编码酶
Table 3. PAHs degrading bacteria, associated genes and encoding enzymes
宿主菌 PAHs 降解基因 编码酶 数据来源 Pseudomonas sp. MPDS 萘、芴、二苯并呋喃、
二苯并噻吩nahAa、nahAb、nahAc、nahAd 萘双加氧酶 文献[14] Pseudomonas fluorescens AH-40、
Sphingomonas koreensis strain ASU-06萘、菲、蒽、芘 nahAc 文献
[41-42]Herbaspirillum sp. strain RV1423 萘 nag 文献[43] Polaromonas naphthalenivorans CJ2 萘 nagAc、nagAd 文献[44] Mycobacterium vanbaalenii PRY-1、Mycobacterium sp. PYR10、
Mycobacterium sp. PYR15菲、芘 nidAB、nidA3B3 环羟基化双加氧酶 文献
[45-46]Rhodococcus sp. P14 蒽、菲、芘、苯并[a]蒽 baaA、baaB 文献[47] Delftia acidovorans Cs1-4 菲 phn 文献[48] Acidovorax strain NA3 萘、菲、苯并[a]蒽、苯并[a]芘 phnAc、phnB、phnC 文献[49] Mycobacterium sp. strain CH-2 萘、菲、荧蒽 pdoA2B2、nidAB 文献[50] Mycobacterium sp. strain 6PY1 芘 pdoA1B1 文献[51] Mycobacterium vanbaalenii strain PYR-1 菲 nidA、nidB、nidC、nidD 文献[52] Roseobacter clade 菲、芘、苯并[a]芘 pahE PAH水合酶-醛缩酶 文献[53] Mycobacterium vanbaalenii PYR-1 芘 phdG 文献[54] Nocardioides sp. strain KP7 萘、菲、蒽、1-甲氧基萘 phdABCD 文献[55] Mycobacterium vanbaalenii PYR-1 芘 phdF 环裂解双加氧酶 文献[54] Sphingomonas koreensis strain ASU-06、Pseudomonas fluorescens AH-40、Pseudomonas aeruginosa、Pseudomonas sp.、Sphingomonas koreensis strain ASU-06、Ralstonia sp. 萘、菲、蒽、荧蒽、芘 C12O、C23O 儿茶酚双加氧酶 文献[41-42,56] -
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