吴川(1983—),女,教授,主要研究方向为土壤污染控制与生态修复。E-mail:
综述了铁锰氧化菌诱导成矿对重金属环境行为的影响,分别从铁/锰氧化菌与生物成矿、铁/锰氧化菌诱导铁锰氧化物沉淀耦合重金属稳定化以及铁锰氧化物对土壤中重金属的作用方面进行阐述;并从铁/锰氧化菌生物成矿方式、铁/锰氧化菌诱导生物成矿过程对土壤重金属的稳定化机制等方面进一步总结了铁/锰氧化菌在不同重金属生物成矿修复中的应用,以及微生物诱导成矿过程的调控因素,分析胞外聚合物、温度与酸碱度、共存离子和其他因素对成矿过程的影响,以期为微生物诱导成矿修复重金属污染提供理论参考。未来工作可进一步关注生成矿物稳定重金属的长效性,不同微生物菌群组合对成矿效果的调控,以及铁/锰氧化菌在重金属复合污染场地土壤修复中的应用等方面。
The effects of iron-manganese oxidizing bacteria-induced mineralization on the environmental behavior of heavy metals were reviewed, including iron/manganese-oxidizing bacteria and biomineralization, iron/manganese-oxidizing bacteria-induced iron-manganese oxide precipitation coupled with heavy metal stabilization, and the effect of iron-manganese oxide on heavy metals in soils. The application of iron/manganese-oxidizing bacteria in the bioremediation of different heavy metals was further summarized from the aspects of the biological mineralization pattern of iron/manganese-oxidizing bacteria (direct or indirect catalytic mineralization of iron oxide protein/enzyme, dual electron transfer reaction of manganese oxidase dominated by polycopper oxidase and the mineralization induced by external factors), heavy metal stabilization mechanisms of iron/manganese-oxidizing bacteria biological mineralization in soils(precipitation/coprecipitation, adsorption/complexation and redox). The effects of extracellular polymers, temperature and pH, coexisting ions and other factors on the mineralization process were also analyzed, in order to provide theoretical references for microbial-induced mineralization to remediate heavy metal pollution. Future work should focus on the long-term stability of heavy metals generated by minerals, the regulation of different microbial species combinations on mineralization, and the application of iron/manganese-oxidizing bacteria in the remediation of soil multi-heavy metal contaminated sites.
重金属暴露是全世界普遍存在的环境问题。据报道,自然和人为原因导致水、土壤和沉积物中的重金属含量持续增加。重金属天然存在于富含金属的铁(氢)氧化物和/或硫化物矿物中,是污染的主要地质成因与来源[
微生物能形成矿物、分解以及转化矿物,而矿物的存在能够促进微生物的生长,使微生物与矿物之间协同作用得到强化,微生物和矿物的交互作用影响着土壤中重金属的环境行为[
生物成矿是在生物的参与下,无机元素选择性地沉积在环境中特殊有机物表面的过程。生物成矿与非生物成矿的显著区别在于是否存在生物基质的调控,生物成矿产物的形态、特征及晶型等会受到生物代谢过程或环境因素的影响调控[
环境中的重金属污染具有隐蔽性、持久性、不可逆转性等特点,其无法被生物降解修复,主要通过凝聚、吸附沉淀、氧化还原、甲基化等作用进行转化以降低其在环境中的生物利用性。而传统的物理化学修复技术成本较高,易破坏土壤生态环境,造成二次污染等。微生物诱导的生物成矿是一种很有前景的环境重金属污染原位修复技术。生物矿物的组成各不相同,根据成矿产物的类型主要可分为碳酸盐矿物、铁锰氧化物、硫化物和磷酸盐矿物[
铁锰氧化物(矿物)形成过程包括非生物过程和生物过程,铁锰氧化物的形成机制可能涉及非生物反应、酶催化或两者的结合。如在中性pH附近氧化条件下Fe(Ⅱ)可通过非生物氧化使之转化为溶解性较小的Fe(Ⅲ)矿,活性铁渗透Fe(Ⅱ)-针铁矿系统,或过氧化氢反应可形成的Fe(Ⅵ)等[
铁/锰氧化菌是一个系统发育多样的类群,能够催化Fe(Ⅱ)、Mn(Ⅱ)形成铁锰氧化物,其广泛分布于土壤环境中[
铁/锰氧化菌诱导生物成矿[
The biomineralization induced by iron/manganese-oxidizing bacteria[
土壤矿物是土壤固相的主要组成部分,约占土壤质量的95%以上。其中铁(氢)氧化物是土壤中重要的次生黏粒矿物,它们不仅具有巨大的比表面积,而且表面还拥有大量的活性官能团,对污染物有很强的界面反应能力,对重金属离子的固定能力强。已有研究[
锰氧化物是环境中重要的强氧化剂,已证明可氧化As(Ⅲ)、Fe(Ⅱ)、Co(Ⅱ)、U(Ⅳ)和Cr(Ⅲ)等。Fe和Mn的氢氧化物特别是Mn的氢氧化物对Pb2+、Cd2+有很强的专性吸附能力。由于生物锰氧化物的表面积和单位面积的结合能均较高,生物锰氧化物较非生物锰氧化物吸附重金属的能力更强[
不同铁/锰氧化菌通过不同的氧化和转运机制,导致铁锰矿物与细胞之间的各种相互作用。在适宜环境条件下,不同的Fe(Ⅱ)氧化菌使用O2(需氧和微需氧Fe(Ⅱ)氧化菌)或硝酸盐/氯酸盐(厌氧Fe(Ⅱ)氧化菌)作为电子受体进行酶催化,将Fe(Ⅱ)氧化为Fe(Ⅲ),而锰氧化菌也可利用O2作为电子受体将Mn(Ⅱ)氧化为Mn(Ⅲ)。此外,重金属(As和Cd等)可能与细胞膜上的膜羟基、磷酰基、氨基等相互作用[
铁/锰氧化菌可通过直接或间接途径氧化Fe(Ⅱ)和/或Mn(Ⅱ),该氧化过程有两种方式:(1)通过蛋白/酶,例如,铁氧化酶、多铜氧化酶(MCO)催化氧化Fe(Ⅱ)和/或Mn(Ⅱ)生成矿物;(2)通过改变环境因子产生相同的结果[
Fe(Ⅱ)氧化蛋白/酶可直接氧化Fe(Ⅱ)。可溶性Fe(Ⅱ)游离至细胞表面,可通过扩散吸收或运输机制进入周质或结合至细胞表面,被分别位于周质或细胞表面的铁氧化酶蛋白氧化[
蛋白/酶可间接氧化Fe(Ⅱ)。在缺氧且存在硝酸盐条件下,硝酸盐依赖型铁氧化菌氧化Fe(Ⅱ)存在不同途径[
Mn(Ⅱ)氧化酶可催化Mn(Ⅱ)进行双电子转移反应。锰氧化是双电子转移过程,锰氧化菌使用细胞表面或胞内独特的Mn(Ⅱ)氧化酶通过单电子转移直接介导Mn(Ⅱ)生成Mn(Ⅲ),随后可能歧化为Mn(Ⅱ)和Mn(Ⅳ)[
在复杂环境中,细胞产生的蛋白质与外界干扰因素有关,重金属胁迫可使细胞产生蛋白质。(1)生物成矿过程中存在的不同矿物表面可改变成矿产物[
铁/锰氧化菌可通过诱导铁锰氧化物沉淀固化/稳定化修复土壤重金属污染。以专性吸附形式吸附在土壤铁锰氧化物上的铁锰结合态重金属迁移性很低,相对较稳定,且相对而言不具有生物有效性。(1)固化修复主要是通过微生物产生的矿物联结土壤颗粒或增加土壤密度来固化土壤[
重金属离子与细菌分泌的阴离子结合形成不溶性沉淀[
铁/锰氧化菌通过沉淀/共沉淀对重金属的修复应用
Application of heavy metal remediation by iron/manganese-oxidizing bacteria through the precipitation/coprecipitation
微生物 |
重金属 |
特征描述 |
锰氧化物/铁氧化物 |
机制 |
重金属浓度 |
缺点 |
参考文献 |
As | 在40 mg·kg-1的Mn (Ⅱ)和20%~30%水分含量下,锰氧化菌(MnOB) 可同时增强As (Ⅲ)的去除、As的固定化和邻苯二甲酸二丁酯(DBP)的降解 | 砷酸盐(如Mg3 (AsO4)2、FeAsO4)、镁铁氧化物和生物锰氧化物 | 细胞色素c (Ccm);吡咯并喹啉醌(PQQ) | 土壤实验—As (Ⅲ): mg·kg-1 | 在土壤试验中,某些微生物可在缺氧条件下将As(Ⅴ)还原为As (Ⅲ) | [ |
|
Cd | 生成多孔海绵状的水钠锰矿过程中,Cd被捕获并将其嵌人固定在晶格结构中 | 水钠锰矿 | 未提及多铜氧化酶(MCO) | 土壤试验—现场采样(稻田土壤中总Cd含量为0.78 mg·kg-1) | 揭示了稻田生物土壤中的八种锰氧化菌株,但未能明确主要作用菌株 | [ |
|
As | 菌株极大地加速As (Ⅴ) -Fe(Ⅲ) 和As (Ⅲ)-Fe (Ⅲ)氢氧化物的形成,对As去除率高达85% | 混合As (Ⅴ) -Fe (Ⅲ)和As (Ⅲ)-Fe (Ⅲ)氢氧化物和图水羟砷铁矶(Fe6(AsO3)4(SO4)(OH)4·4H2O) | 外膜细胞色素c (Cyc2); Petl操纵子 | 实验室试验—现场采样(酸性矿山废水中As: Fe摩尔比高达0.6~0.8) | 该酸性矿山废水中砷氧化速率的变化可能与细菌活动的空间和季节变化有关,这些变化的起源尚不清楚 | [ |
|
As | 锰氧化菌介导矿物生成中,As去除的最佳Mn (Ⅱ)/Fe (Ⅱ)比约为1 : 3 (mol:mol) | 羟基氧化铁(FeOOH)和二氧化锰(MnO2) | 多铜氧化酶-MCO (CumA) | 实验室水溶液—10 μmol·L-1 As、100 μmol·L-1 Mn (Ⅱ) 和不同的Fe (Ⅱ)初始浓度 | 当地下水中含有高浓度的Mn (Ⅱ)、Fe (Ⅱ)和As (Ⅲ/Ⅴ)时,可使用生物强化技术来消除或消除 | [ |
细菌表面官能团和细胞周围胞外聚合物对重金属的吸附与络合是稳定重金属的第二大途径[
铁/锰氧化菌通过吸附/络合对重金属的修复应用
Application of heavy metal remediation by iron/manganese-oxidizing bacteria through the adsorption/complexation
微生物 |
重金属 |
特征描述 |
锰氧化物/铁氧化物 |
机制 |
重金属浓度 |
缺点 |
参考文献 |
Cd | 与 |
中间产物为绿锈,最终产物为针铁矿 | 硝酸还原酶,亚硝酸盐积累非生物氧化Fe (Ⅱ) | 温室大麦宮养试验—添加5 mg·kg-1 CdCl2 和15 mg·kg-1 Cd (NO3)2 | 针铁矿初始吸附Cd最低量和pH为5.5下,温度为20℃时,累积Cd脱附达71% | [ |
|
Cd | 当Fe (Ⅱ)和Mn (Ⅱ)用作电子给体时,Cd (Ⅱ)的最大去除效率分别为80.63%和84.58% | 羟基氧化铁(FeOOH) 和二氧化锰(MnO2) | 细胞色素cdl; 亚硝酸盐还原酶;硝酸还原酶 | 实验室水溶液试验—初始Cd (Ⅱ)浓度为10 mg·L-1 | 该菌株产生的气体中检测出氮气(N2)和一氧化氮(N2O),培养瓶中最终产物为N2,但实际应用中N2O是否会还原为N2有待商榷 | [ |
|
U、Co | 铁氧化菌产生的氧化物对铀的去除率达55%,对钴的去除率达81% | 以磁铁矿为主的铁矿物,铀与铁氧化物形成双齿和三齿内球络合物 | 未提及,猜测为硝酸还原酶,亚硝酸盐积累非生物氧化Fe (Ⅱ) | 实验室水溶液实验—100 μmol·L-1 U(Ⅳ) 和100 μmol·L-1 Co (Ⅲ) | 关于该菌株的作用酶/蛋白研究较少 | [ |
|
Sb、As | 生物铁锰氧化物氧化Sb (Ⅲ) 的效率高于As (Ⅲ),但吸附As的效率高于Sb | 羟基氧化铁(FeOOH)、氢氧化锑/生物锰氧化物(Sb(OH)3 /BMO) 络合物 | 多铜氧化酶-MCO (CumA) | 实验室水溶液试验—10 μmol·L-1 As (Ⅲ) 和Sb (Ⅲ) | Fe (Ⅱ)、As (Ⅲ)和Sb (Ⅲ) 的存在加速了Mn (Ⅱ)的氧化,但抑制了代表微生物活性的16S rRNA基因表达水平 | [ |
对于铁还原菌、铁氧化菌等微生物,与其代谢相关的氧化还原过程可改变剧毒重金属离子的价态,形成毒性较小的离子[
铁/锰氧化菌通过氧化还原对重金属的修复应用
Application of heavy metal remediation by iron/manganese-oxidizing bacteria through redox
微生物 |
重金属 |
特征描述 |
锰氧化物/ |
机制 |
重金属浓度 |
缺点 |
参考文献 |
Cr | 生物锰氧化物氧化Cr(Ⅲ)的初始速率较等量合成的二氧化锰氧化Cr(Ⅲ)的速率快约7倍 | 水钠锰矿 | 多铜氧化酶- MCO(MnxEFG) | 实验室模拟海水条件—0、10、100 µmol·L–1 Cr(Ⅲ) | 未加实验介质前,大于5 µmol·L–1的Cr(Ⅲ)浓度抑制Mn(Ⅱ)和Cr(Ⅲ)氧化 | [ |
|
Cr | 100 µmol·L–1Cr(Ⅲ)在24h内几乎100%被氧化为Cr(Ⅵ) | 水钠锰矿 | 多铜氧化酶-MCO(CumA,MnxG,McoA); |
实验室培养基实验—0、10、50、100 µmol·L–1 Cr(Ⅲ) | Cr(Ⅲ)可能进入细胞,间接抑制Mn(Ⅱ)氧化 | [ |
生物矿物的形成通常受到各种因素的共同作用,包括生物体与有机基质、反应条件(酸碱度、温度等)、共存离子、腐殖质与其他因素等。
生物矿物中的有机基质可定义为与组分表面结合的任何有机物质,如蛋白质、磷脂、胶原和碳水化合物等。特别是细胞代谢所产生的大分子有机物——胞外聚合物(EPS),在微生物成矿过程中扮演着至关重要的角色(
胞外聚合物(EPS)在生物成矿过程中的作用[
The role of extracellular polymeric substances(EPS) in the biomineralization[
温度是影响生物成矿反应过程的重要因素。通常,温度对生物成矿的影响因微生物而异,而且温度和酸碱度的影响通常会有协同作用。pH的升高会影响菌株的生长和重金属离子的沉淀。酸性条件不利于Mn(Ⅱ)的氧化,而Fe(Ⅲ)在碱性条件下易形成铁(氢)氧化物沉淀[
反应体系中存在的各种离子、腐殖质以及其他因素也是影响生物成矿多样性和复杂性的重要因素之一(
共存离子与腐殖质等其他因素在生物成矿过程中的作用[
The role of coexisting ions and other factors in the biomineralization process[
腐殖质作为天然有机质,是微生物生长的重要营养物质,主要由腐殖酸、黄腐酸和胡敏素等组成[
生物成矿研究的发展使其在土壤重金属固化/稳定化修复方面成为一个有前途的可持续解决方案。不同铁/锰氧化菌诱导生物成矿形成的矿物类型、晶体结构等性质显著不同,在一定条件下,铁锰矿物还可进行二次成矿,耦合和固定更多的重金属。不同晶型的矿物固定重金属及促使重金属价态变化的机制更为复杂,未来研究可进一步聚焦:(1)矿物稳定重金属的长效性研究:固化/稳定化之后的重金属可能重新活化、随环境条件变化等重新溶出,未来的研究可提高生成矿物的晶格稳定性和抗侵蚀性;(2)复合菌效果研究:铁/锰氧化菌的成矿方式类似,两种类别菌株复合对As、Cr、Cd等重金属是否存在协同效果;(3)研究生物成矿在场地土壤复合重金属污染的修复应用:大多数关于重金属铁锰(氢)氧化物生物成矿的研究是在实验室、水溶液条件及针对单一重金属的探讨,实际的土壤污染通常是多种金属的复合污染,且不同土壤的理化性质存在较大差异,因此,亟需研究生物成矿对实际污染场地多种重金属复合污染的修复机制及应用。
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