厌氧甲烷氧化古菌共生机制的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-1100

摘要/Abstract

摘要：

厌氧甲烷氧化（AOM）是全球碳循环中的关键过程，对遏制甲烷排放和缓解气候变化具有重要作用。该过程一类主要执行者为厌氧甲烷氧化古菌（ANME），它直接或与细菌协同，将AOM与不同还原电位的最终电子受体，如硫酸盐、腐殖质、金属氧化物以及硝酸盐等耦合反应。其中，AOM与硫酸盐还原耦合反应的吉布斯自由能变化极低（ΔG°' = -17 kJ/mol），且所获得的能量需要与其共生硫酸盐还原细菌（SRB）进行有效分配，因此ANME与SRB之间的高效能量转移机制自1999年ANME发现以来一直是研究核心。本文综述了AOM及其关键微生物的研究历程，重点探讨了ANME与SRB之间的3种共生模式假说（可扩散化学中间体、零价硫、直接种间电子传输），并总结了通过环境观测或富集培养发现的ANME及其共生或伴生伙伴的系统发育多样性与潜在互作方式。这些研究不仅加深了对AOM这一关键生物地球化学过程的认识，也为全面揭示其种间互作机制并拓展其应用潜力提供了参考。

关键词: 厌氧甲烷氧化, 甲烷氧化古菌, 共生机制, 直接种间电子传输, 硫酸盐还原菌

Abstract:

Anaerobic oxidation of methane (AOM) is a key process in the global carbon cycle, playing a vital role in curbing methane emissions and mitigating climate change. The primary mediators of this process are anaerobic methanotrophic archaea (ANME), which oxidize methane either independently or in cooperation with partner bacteria by coupling AOM to terminal electron acceptors with varying redox potentials, such as sulfate, humic substances, metal oxides, and nitrate. AOM coupled to sulfate reduction yields extremely low Gibbs free energy change (ΔG°' = -17 kJ/mol), and the energy yield must be efficiently allocated with their symbiotic sulfate-reducing bacterial (SRB). Consequently, the symbiotic mechanism in AOM has been a central research focus since the discovery of ANME in 1999. This review outlines the research progress on the microorganisms mediating AOM, with an emphasis on three hypothesized symbiotic models between ANME and SRB: diffusible chemical intermediates, zero-valent sulfur, and direct interspecies electron transport. Additionally, we summarize the phylogenetic diversity and potential interaction mechanisms of ANME and their symbiotic or associated partners revealed by environmental observations and enrichment cultures. Together, these advances have not only enhanced our understanding of the AOM process but also paved the way to uncover the full range of interspecies interaction mechanisms and broaden potential biotechnological applications.

Key words: anaerobic oxidation of methane, anaerobic methanotrophic archaea, symbiotic mechanism, direct interspecies electron transport, sulfate-reducing bacteria

赵雨婷, 喻航. 厌氧甲烷氧化古菌共生机制的研究进展[J]. 生物技术通报, 2026, 42(2): 3-16.

ZHAO Yu-ting, YU Hang. Advances in the Symbiotic Mechanisms of Anaerobic Methanotrophic Archaea[J]. Biotechnology Bulletin, 2026, 42(2): 3-16.

图/表 2

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厌氧甲烷氧化古菌 Anaerobic methanotrophic archaea （ANME）	共生/伴生微生物 Symbiotic/Associated microorganism	电子受体/代谢特征 Electron acceptor/Metabolic characteristics	参考文献 Reference
ANME-1 （Candidatus Methanophagaceae）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［65-66，89］
	SEEP-SRB1 （ETH-SRB1）	Fe（Ⅲ）、Mn（Ⅳ）/潜在金属还原	［22］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［70］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［39，70，109，112］
	Candidatus Desulfofervidus	SO₄^2-	［93］
	Candidatus Desulfaltia	SO₄^2-	［113］
ANME-1 （Guaymas cluster）	*HotSeep-1 （Candidatus* Desulfofervidus auxilia）**	SO₄^2-	［38-39，56，96，99，112］
ANME-1a （Candidatus Methanophagaceae）	*HotSeep-1 （Candidatus* Desulfofervidus auxilia）**	SO₄^2-	［60，100］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［60］
	Verrucomicrobia	可能利用细胞分泌的有机渗出物或胞外聚合物	［120］
ANME-1b （Candidatus Methanophagaceae）	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［50］
ANME-2 （Unknown subgroup）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［14，33，61，65，67，89-90，108-109，115-116，128］
	Desulfobulbaceae （DSB）	SO₄^2-	［116］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［70］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［39］
	Candidate division JS-1 （Atribacterota）	可能受益于AOM过程或相关代谢物	［64］
ANME-2a （Candidatus Methanocomedens）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［18，68-69］
	SEEP-SRB1	SO₄^2-	［93］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［40，92，110，112，117］
	Methanococcoides，Anaerolineales	可能依赖体系内有机物降解获取能量	［40］
ANME-2b （Candidatus Methanomarinus）	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［117］
ANME-2b （Candidatus Methanomarinus）	*SEEP-SRB1g （Candidatus* Desulfomellonium）**	SO₄^2-	［112，117］
ANME-2c （Candidatus Methanogasteraceae）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［18，68-69，114，118］
	SEEP-SRB1 （ETH-SRB1）	Fe（Ⅲ）、Mn（Ⅳ）/潜在金属还原	［22］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［92，95，110，112，117］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［60，109，112］
	Desulfobulbaceae （seepDBB/unknown subgroup）	SO₄^2-	［114，118］
	Betaproteobacteria	可能提供了新的氮代谢途径	［114］
ANME-2c or ANME-2b	non-SEEP-SRB1a Deltaproteobacteria	SO₄^2-	［95］
ANME-2d （Candidatus Methanoperedens）	Geobacteraceae	生物阳极、生物炭、腐殖质/介导电子传递	［106-107］
ANME-2d （Candidatus Methanoperedens nitroreducens）	Candidatus Methylomirabilis oxyfera Candidatus Kuenenia stuttgartiensis	NO₃^-/能够将亚硝酸盐还原为氮气	［15］［17］
ANME-3 （Candidatus Methanovorans）	Desulfobulbus （DBB）	SO₄^2-	［61，71］
ANME-3 （Candidatus Methanovorans）	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［110］

厌氧甲烷氧化古菌 Anaerobic methanotrophic archaea （ANME）	共生/伴生微生物 Symbiotic/Associated microorganism	电子受体/代谢特征 Electron acceptor/Metabolic characteristics	参考文献 Reference
ANME-1 （Candidatus Methanophagaceae）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［65-66，89］
	SEEP-SRB1 （ETH-SRB1）	Fe（Ⅲ）、Mn（Ⅳ）/潜在金属还原	［22］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［70］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［39，70，109，112］
	Candidatus Desulfofervidus	SO₄^2-	［93］
	Candidatus Desulfaltia	SO₄^2-	［113］
ANME-1 （Guaymas cluster）	*HotSeep-1 （Candidatus* Desulfofervidus auxilia）**	SO₄^2-	［38-39，56，96，99，112］
ANME-1a （Candidatus Methanophagaceae）	*HotSeep-1 （Candidatus* Desulfofervidus auxilia）**	SO₄^2-	［60，100］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［60］
	Verrucomicrobia	可能利用细胞分泌的有机渗出物或胞外聚合物	［120］
ANME-1b （Candidatus Methanophagaceae）	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［50］
ANME-2 （Unknown subgroup）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［14，33，61，65，67，89-90，108-109，115-116，128］
	Desulfobulbaceae （DSB）	SO₄^2-	［116］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［70］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［39］
	Candidate division JS-1 （Atribacterota）	可能受益于AOM过程或相关代谢物	［64］
ANME-2a （Candidatus Methanocomedens）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［18，68-69］
	SEEP-SRB1	SO₄^2-	［93］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［40，92，110，112，117］
	Methanococcoides，Anaerolineales	可能依赖体系内有机物降解获取能量	［40］
ANME-2b （Candidatus Methanomarinus）	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［117］
ANME-2b （Candidatus Methanomarinus）	*SEEP-SRB1g （Candidatus* Desulfomellonium）**	SO₄^2-	［112，117］
ANME-2c （Candidatus Methanogasteraceae）	*Desulfobacteraceae （Desulfosarcina/Desulfococcus）（DSS）*	SO₄^2-	［18，68-69，114，118］
	SEEP-SRB1 （ETH-SRB1）	Fe（Ⅲ）、Mn（Ⅳ）/潜在金属还原	［22］
	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［92，95，110，112，117］
	*SEEP-SRB2 （Candidatus* Desulfomithrium）**	SO₄^2-	［60，109，112］
	Desulfobulbaceae （seepDBB/unknown subgroup）	SO₄^2-	［114，118］
	Betaproteobacteria	可能提供了新的氮代谢途径	［114］
ANME-2c or ANME-2b	non-SEEP-SRB1a Deltaproteobacteria	SO₄^2-	［95］
ANME-2d （Candidatus Methanoperedens）	Geobacteraceae	生物阳极、生物炭、腐殖质/介导电子传递	［106-107］
ANME-2d （Candidatus Methanoperedens nitroreducens）	Candidatus Methylomirabilis oxyfera Candidatus Kuenenia stuttgartiensis	NO₃^-/能够将亚硝酸盐还原为氮气	［15］［17］
ANME-3 （Candidatus Methanovorans）	Desulfobulbus （DBB）	SO₄^2-	［61，71］
ANME-3 （Candidatus Methanovorans）	*SEEP-SRB1a （Candidatus* Syntrophophila）**	SO₄^2-	［110］