生态环境学报 ›› 2021, Vol. 30 ›› Issue (11): 2257-2266.DOI: 10.16258/j.cnki.1674-5906.2021.11.017
李侠1(
), 兰建英2,3, 蒋海明2,3,*()
收稿日期:2021-01-10
出版日期:2021-11-18
发布日期:2021-12-29
通讯作者:
* 蒋海明（1977年生）,男,教授,主要研究方向为应用环境微生物、生物能源、环境污染物的生物治理及监测。E-mail: jhmhn@163.com作者简介:李侠（1979年生）,女（满族）,副教授,研究方向为环境微生物与微生物冶金。E-mail: lixia2002hn@163.com
基金资助:
LI Xia1(), LAN Jianying2,3, JIANG Haiming2,3,*()
Received:2021-01-10
Online:2021-11-18
Published:2021-12-29
摘要：
甲烷（CH4）是一种温室气体,其温室效应是二氧化碳的26倍。减少甲烷排放,可以缓解温室效应,有助于保护大气环境。微生物介导的甲烷厌氧氧化是调节甲烷向大气排放的关键过程。金属氧化物或金属离子可以作为甲烷厌氧氧化的电子受体,且金属依赖的甲烷厌氧氧化是减小全球甲烷排放的一个重要途径。虽然研究表明金属依赖型厌氧甲烷营养古菌可以促进这一过程,但其菌种、代谢途径及胞外电子传递途径尚缺乏深入研究。文章从金属依赖型厌氧甲烷氧化菌、金属依赖型厌氧甲烷氧化机制及金属依赖型厌氧甲烷氧化菌胞外电子传递机制等方面对金属依赖型厌氧甲烷氧化研究现状进行了概述,分析了金属依赖型厌氧甲烷氧化研究存在的问题,并讨论了其今后的研究方向,为金属依赖型厌氧甲烷氧化研究提供参考。
中图分类号:
李侠, 兰建英, 蒋海明. 金属依赖型厌氧甲烷氧化研究进展[J]. 生态环境学报, 2021, 30(11): 2257-2266.
LI Xia, LAN Jianying, JIANG Haiming. Advance in Metal Iron-dependent Anaerobic Oxidation of Methane[J]. Ecology and Environment, 2021, 30(11): 2257-2266.
图1 3种不同甲烷厌氧氧化模型（Cui et al.,2015）
Fig. 1 Three different models of anaerobic oxidation of methane (AOM) depending on the different electron acceptors (Cui et al., 2015)
| 金属 | 主要功能微生物 | 文献 |
|---|---|---|
| 铁(III) | ANME-1, ANME-2及ANME-3 | Chang et al., |
| ANME-2a和ANME-2c | Scheller et al., | |
| AAA | Ettwig et al., | |
| M. acetivorans C2A | Soo et al., Yan et al., | |
| Ca. M. ferrireducens | Cai et al., | |
| 锰(Ⅳ) | Ca. M. manganicus, Ca. M. manganireducens | Leu et al., |
| 硒 (VI) | Ca. Methanoperedens, Ca. Methylomirabilis | Luo et al., |
| Methylomonas | Lai et al., | |
| 钒(Ⅴ) | Methylomonas, Methanobacterium, Stenotrophomonas | Zhang et al., |
| 锑(V) | Methanosarcina和Sb(V)-reducing microbe (unknown) | Lai et al., |
| 铬(VI) | ANME-2d和/或Cr(VI)-reducing microbe (unknown) | Lu et al., |
| Methanobacterium, Methanosarcina, Meiothermus, ANME-2d | Dong et al., | |
| Ca. Methanoperedens和/或Cr(VI)-reducing microbe (unknown) | Luo et al., | |
| Methylococcus capsulatus (Bath) | Al-Hasin et al., | |
| 砷(V) | ANME-1和ANME-2a-c | Shi et al., |
表1 文献报道的金属依赖型厌氧甲烷氧化微生物
Table 1 Microbes reported as M-DAOM
| 金属 | 主要功能微生物 | 文献 |
|---|---|---|
| 铁(III) | ANME-1, ANME-2及ANME-3 | Chang et al., |
| ANME-2a和ANME-2c | Scheller et al., | |
| AAA | Ettwig et al., | |
| M. acetivorans C2A | Soo et al., Yan et al., | |
| Ca. M. ferrireducens | Cai et al., | |
| 锰(Ⅳ) | Ca. M. manganicus, Ca. M. manganireducens | Leu et al., |
| 硒 (VI) | Ca. Methanoperedens, Ca. Methylomirabilis | Luo et al., |
| Methylomonas | Lai et al., | |
| 钒(Ⅴ) | Methylomonas, Methanobacterium, Stenotrophomonas | Zhang et al., |
| 锑(V) | Methanosarcina和Sb(V)-reducing microbe (unknown) | Lai et al., |
| 铬(VI) | ANME-2d和/或Cr(VI)-reducing microbe (unknown) | Lu et al., |
| Methanobacterium, Methanosarcina, Meiothermus, ANME-2d | Dong et al., | |
| Ca. Methanoperedens和/或Cr(VI)-reducing microbe (unknown) | Luo et al., | |
| Methylococcus capsulatus (Bath) | Al-Hasin et al., | |
| 砷(V) | ANME-1和ANME-2a-c | Shi et al., |
图2 微生物介导的M-DAOM不同机制（He et al.,2018;何丹等,2020） （A）ANME单独完成M-DAOM M-DAOM by ANME alone;（B）ANME与合作伙伴MRM共同完成M-DAOM M-DAOM by cooperation between ANME and partner MRM;（C）产甲烷古菌单独完成M-DAOM M-DAOM by archaea methanogens alone;（D）产甲烷古菌与电子穿梭体完成M-DAOM M-DAOM by archaea methanogens and electron shuttle;（E）产甲烷古菌和MRM通过甲烷代谢中间产物完成M-DAOM M-DAOM by archaea methanogens and MRM through intermediate products of methane metabolism。ANME：厌氧甲烷营养古菌 anaerobic methanotrophic archaea;MRM：金属还原微生物 metal-reducing microorganisms
Fig. 2 Different mechanisms of microbe-mediated M-DAOM (He et al., 2018; He et al., 2020)
图3 M. acetivorans C2A Fe(Ⅲ)依赖的甲烷氧化及胞外电子传递假设途径（Yan et al.,2018） CO脱氢酶/乙酰辅酶A合成酶 CO dehydrogenase/acetyl-CoA synthase（反应5）;乙酸激酶和磷酸转乙酰酶 acetate kinase and phosphotransacetylase（反应6）;辅酶F420依赖的亚甲基四氢八叠蝶呤还原酶 coenzyme F420 (F420)-dependent methylenetetrahydrosarcinapterin (H4SPT) reductase、辅酶F420依赖的亚甲基四氢八叠蝶呤脱氢酶 F420-dependent methylene-H4SPT dehydrogenase、甲基四氢八叠蝶呤环化水解酶 methenyl-H4SPT cyclohydrolase、甲酰甲烷呋喃：四氢八叠蝶呤甲酰转移酶 formylmethanofuran H4SPT formyltransferase（反应7）;甲酰甲烷呋喃脱氢酶 formylmethanofuran dehydrogenase（反应8）。Fwd：甲酰甲烷呋喃脱氢酶 formyl-methanofuran dehydrogenase;Ftr：甲酰甲烷呋喃/四氢八叠蝶呤甲酰转移酶 formylmethanofuran/H4MPT formyltransferase;Mch：甲基四氢八叠蝶呤环化水解酶 methenyl-H4MPT cyclohydrolase;Mtd：辅酶F420依赖的亚甲基四氢八叠蝶呤脱氢酶 F420-dependent methylene-H4MPT dehydrogenase;Mer：F420依赖的亚甲基四氢八叠蝶呤还原酶 F420-dependent methylene-H4MPT reductase;Mtr：转运Na+的甲基四氢甲烷蝶呤：辅酶M甲基转移酶 Na+-translocating methyl-H4MPT：coenzyme M methyltransferase;Mcr：甲基辅酶M甲基还原酶 methyl-coenzyme M reductase;Fpo：还原态辅酶F420脱氢酶 F420H2 dehydrogenase;ATPase：腺苷三磷酸合成酶 ATP synthetase;HdrDE：异二硫化物还原酶亚单位D和E heterodisulfide reductase subunits D and E;HdrA2B2C2：异二硫化物还原酶亚单位A2、B2和C2 heterodisulfide reductase subunits A2, B2, and C2;Rnf：Rnf 复合体 Rnf complex;Mrp：Na+/H+逆向转运体 Na+/H+ antiportor;MP：氧化态甲烷吩嗪 oxidized methanophenazine;MPH2：还原态甲烷吩嗪 reduced methanophenazine;H4MPT：四氢甲烷蝶呤 tetrahydromethanopterin;MFR：甲烷呋喃 methanofuran;cyt c：c型细胞色素 c-type cytochrome;F420H2：还原态辅酶F420 reduced coenzyme F420;F420：氧化态辅酶F420 oxidized coenzyme F420;CoB-SH：辅酶B coenzyme B;CoM-SH：辅酶M coenzyme M;CoM-S-S-CoB：辅酶M和辅酶B的异二硫化物 Heterodisulfide of coenzyme M and coenzyme B;Fdox：氧化态铁氧化还原蛋白 oxidized ferredoxin;Fdred-：一个电子还原的铁氧化还原蛋白 one-electron reduced ferredoxin;Fdred2-：完全还原的铁氧化还原蛋白 fully reduced ferredoxin;AQDS：蒽醌2, 6-二磺酸钠 Anthraquinone-2, 6-disulfonic Acid Disodium.
Fig. 3 Pathway proposed for Fe(III)-dependent methane oxidation and conservation of energy by M. acetivorans (Yan et al., 2018) 5CH4+32Fe(III)+13ADP+13HPO4=→CH3COOH+32Fe(II)+19H++13ATP +3CO2+5H2O ΔG°′= -2880.7 kJ
图4 胞质异二硫化物还原酶HdrA2B2C2利用CoM-S-S-CoB和F420H2还原Fdox假设机制（Yan et al.,2017） Fdred-：一个电子还原的铁氧化还原蛋白 one electron reduced ferredoxin;Fdox：氧化态铁氧化还原蛋白 oxidized ferredoxin;F420：氧化态辅酶F420 oxidized coenzyme F420;F420H2：还原态辅酶F420 reduced coenzyme F420;CoB-SH：辅酶B coenzyme B;CoM-SH：辅酶M coenzyme M;CoM-S-S-CoB：辅酶M和辅酶B的异二硫化物 Heterodisulfide of coenzyme M and coenzyme B;HdrA2B2C2：异质二硫化物还原酶亚单位A2、B2和C2 heterodisulfide reductase subunits A2, B2, and C2;FAD：黄素二核苷酸?avin adenine dinucleotide;Fe-S：铁硫簇 iron-sulfur cluster
Fig. 4 Proposed scheme for the reduction of Fdox with CoM-S-S-CoB and F420H2 by cytoplasmic electron confurcating heterodisulfide reductase HdrA2B2C2 complex (Yan et al., 2017)
图5 M. ferrireducens Fe(Ⅲ)依赖的甲烷氧化及胞外电子传递假设途径（Cai et al.,2018） H4MPT：四氢甲烷蝶呤 tetrahydromethanopterin;MFR：甲烷呋喃 methanofuran;Fwd：甲酰甲烷呋喃脱氢酶 formyl-methanofuran dehydrogenase;Ftr：甲酰甲烷呋喃/四氢甲烷蝶呤甲酰转移酶 formylmethanofuran/H4MPT formyltransferase;Mch：甲基四氢甲烷蝶呤环化水解酶 methenyl-H4MPT cyclohydrolase;Mtd：辅酶F420依赖的亚甲基四氢甲烷蝶呤脱氢酶 F420-dependent methylene-H4MPT dehydrogenase;Mer：F420依赖的亚甲基四氢甲烷蝶呤还原酶 F420-dependent methylene-H4MPT reductase;Mtr：转运Na+的甲基四氢甲烷蝶呤：辅酶M甲基转移酶 Na+-translocating methyl-H4MPT：coenzyme M methyltransferase;Mcr：甲基辅酶M甲基还原酶 methyl-coenzyme M reductase;Fpo：还原态辅酶F420脱氢酶 F420H2 dehydrogenase;ATPase：腺苷三磷酸合成酶 ATP synthetase;HdrDE：异二硫化物还原酶亚单位D和E subunits D and E of heterodisulfide reductase;HdrABC：异二硫化物还原酶亚单位A、B和C2 subunits A, B, and C of heterodisulfide reductase;FrhB：辅酶F420还原氢化酶B亚单位 subunit B of F420-reducing hydrogenase;cytb：b型细胞色素 b-type cytochrome;NrfD：多硫化物还原酶D亚单位 subunit D of polysulfide reductase;FeS：铁氧还蛋白铁硫蛋白 ferredoxin iron sulfur protein;MK：氧化态甲基萘醌 oxidized menaquinone;MKH2：还原态甲基萘醌reduced menaquinone;F420H2：还原态辅酶F420 reduced coenzyme F420;F420：氧化态辅酶F420 oxidized coenzyme F420;CoB-SH：辅酶B coenzyme B;CoM-SH：辅酶M coenzyme M;CoM-S-S-CoB：辅酶M和辅酶B的异二硫化物 Heterodisulfide of coenzyme M and coenzyme B;Fdox：氧化态铁氧化还原蛋白 oxidized ferredoxin;Fdred2-：完全还原的铁氧化还原蛋白 fully reduced ferredoxin。下同 The same below
Fig. 5 Metabolic construction of the putative pathway for AOM coupled to Fe(III) reduction in “M. ferrireducens” (Cai et al., 2018)
图6 Ca. Methanoperedens基因组中AOM与Mn(IV)还原偶联的代谢途径构建（Leu et al.,2020）
Fig. 6 Metabolic construction of the putative pathway for AOM coupled to Mn(IV) reduction in the “Ca. Methanoperedens” genomes (Leu et al., 2020)
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