Geobiological Perspective for the Formation of Manganiferous Deposit: Principle, Evidence, and Model
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摘要: 锰是地球系统中重要的过渡金属元素，其地球化学行为主要受到生物与环境的协同控制.沉积/成岩环境中，氧化还原条件与pH值对于锰的沉积-富集过程最为重要.在地质历史时期，大规模锰矿沉积的形成被认为与地球表层系统的氧化过程相关.锰矿的基础研究意义及重要经济属性则促使相关研究者尤为关注锰元素地球化学循环与锰矿成矿机制之间的联系.近年来，随着锰元素的地球生物学富集机制与循环过程相关研究的深入，研究者们逐渐认识到微生物活动在促进沉积物中锰质富集起到了关键控制作用.通过研究现代代表性沉积环境中所形成的富锰沉积物，明确了微生物作用与沉积环境效应共同控制了锰元素在沉积物中的富集过程.但是此前对于深时锰矿床中的地球生物学相关研究较为分散，对研究方法与成矿机制的综述工作尚存不足.本文通过检视世界范围内当前沉积型锰矿床中微生物成矿作用相关研究案例与研究进展，从中抽提出4个重要分析技术模块：（1）显微观察模块；（2）光谱分析模块；（3）同位素信号模块与（4）综合分析模块.以上4个技术模块的使用可有效识别出微生物成矿相关证据.当前锰矿床内微生物成矿作用相关证据包括：微生物成因显微结构与构造、广泛发育的生物成因自生矿物、具有显著生物信号的碳-硫及其他稳定同位素特征、与微生物活动有关的元素或组分富集现象及生物标志物等有机地球化学信号等.综合沉积型锰矿床在地球生物学视角下的成矿过程，可总结出微生物参与下的锰矿两阶段成矿机制，包括第1阶段的氧化富集阶段与第2阶段的还原保存阶段.锰氧化微生物有可能与铁氧化微生物及光合作用微生物共同构成了复杂的微生物席体系.Abstract: Manganese (Mn) is a crucial transition metal element within the Earth system, whose geochemical behavior is predominantly influenced by the synergistic interaction of biological and environmental factors. In sedimentary and diagenetic environments, redox conditions and pH levels play a pivotal role in controlling the processes of manganese precipitation and enrichment. Over geological history, the formation of large-scale manganese deposits has been closely linked to oxidation in Earth's surface systems. The fundamental research significance and considerable economic value of manganese deposits have driven a sustained focus on the relationship between the geochemical cycling of manganese and the mechanisms underlying ore formation. Recent advances in studies on the geobiological enrichment mechanisms and cycling processes of manganese have highlighted the critical role of microbial activity in manganese enrichment within sediments. Research on manganiferous sediments in representative modern sedimentary environments indicates that the enrichment of manganese is jointly governed by microbial processes and sedimentary environmental factors. Nevertheless, studies on the geobiological aspects of ancient manganese deposits have been fragmented, and comprehensive reviews of research methods and ore-forming mechanisms remain inadequate. This study systematically reviews current research cases and progress on microbial mineralization in sedimentary manganese deposits worldwide, identifying four key analytical technology modules: (1) microscopic observation, (2) spectroscopic analysis, (3) isotopic signal analysis, and (4) integrative analysis. These modules collectively enable the effective extraction of evidence related to microbial mineralization processes. Key evidence includes microstructures and textures of microbial origin, extensively developed biogenic authigenic minerals, stable isotopic characteristics (e.g., carbon-sulfur isotopes) with distinct biological signals, elemental or compositional enrichment associated with microbial activity, and organic geochemical signatures such as biomarkers. A geobiological synthesis of the ore-forming processes in sedimentary manganese deposits reveals a two-stage ore-forming mechanism involving microbial participation. This mechanism comprises an initial oxidation-enrichment stage (Stage 1) and a subsequent preservation stage under reducing conditions (Stage 2). Manganese-oxidizing microbes likely function within complex microbial mat systems, interacting with iron-oxidizing microbes and photosynthetic microbes to facilitate manganese deposition.
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图 1 中国南方地区成冰系至二叠系代表性锰矿石内的微生物记录镜下证据
其中a、b、c图分别为成冰系大塘坡组、石炭系巴平组、二叠系茅口组锰矿石的岩石薄片照片.a1至a3为成冰系大塘坡组纹层状锰矿石中微生物相关构造（均为正交镜下观察结果，放大40倍，红色箭头指示微生物结构）.b1与b2指示了样品中的微米级结核及类似生物席的弱层状纹理，b3为b2的阴极发光照片.c1至c3为矿化的微生物产生的微观纹，明显可见结核和颗粒悬浮在微晶基质中.综合修改自Yu et al.（2019，2021a，2021b）
Fig. 1. Microscopic evidence of microbial records within representative manganese deposits from the Nanhua System to the Permian in southern China
图 2 扫描电子显微镜（SEM）观察到的中国南方地区成冰系大塘坡组（a、b），石炭系巴平组（c、d）及二叠系茅口组（e、f）锰矿石中似平行状微层理、交织状微层理及微团块与微球粒等与微生物成矿作用相关的显微沉积构造
Fig. 2. Microbial mineralization-related micro-sedimentary structures observed under scanning electron microscopy (SEM) in manganese ores from the Cryogenian Datangpo Formation (a, b), the Carboniferous Baping Formation (c, d), and the Permian Maokou Formation (e, f) in southern China, including parallel-like micro-laminations, interwoven micro- laminations, micro-nodules, and micro-spherules
图 3 广西中部忻城地区石炭系巴平组锰矿样品MS-2的岩石薄片及拉曼光谱分析所得矿物和有机质分布情况
拉曼光谱分析路径沿图中的L4-L4′.红色箭头标注了锐钛矿产出层位.修改自Yu et al.（2021b）
Fig. 3. Mineralogical and organic matter distribution in the thin section and Raman analysis of the Carboniferous Baping Formation manganese ore sample MS-2 from the Xincheng area, central Guangxi
图 4 地质历史时期碳酸锰矿床中碳同位素记录与锰含量相关关系
其中图4a为无机碳同位素（δ13Ccarb），图4b为有机碳同位素（δ13Corg）.数据来源于Dong et al.（2023）、Gao et al.（2021）、Gutzmer and Beukes（1998）、Hein et al.（1999）、Huang et al.（2022）、Kuleshov and Bych（2002）、Okita（1992）、Polgari et al.（1991）、Tan et al.（2021）、Wang et al.（2019）、Wu et al.（2016）、Yu et al.（2016，2021b，2024）、Zhang et al.（2020，2021b）
Fig. 4. Correlation between carbon isotope record
表 1 锰氧化细菌及真菌的常见种属及分类
Table 1. Common genera and classifications of manganese-oxidizing bacteria and fungi
分类 细菌 真菌 门类 变形菌门Proteobacteria 放线菌门Actinobacteria 厚壁菌门Firmicutes 子囊菌门Ascomycota 属种 纤发菌属Leptothrix 阿霉素属Agromyces 芽孢杆菌属Bacillus 黄瓜织球壳菌Plectosphaerella cucumerina 假单胞菌属Pseudomonas 微杆菌属Microbacterium 短芽胞杆菌属Brevibacillus 棘壳孢属Pyrenochaeta sp. 纤维素单胞菌Cellulomonas 铜绿假单胞菌属Cupriavidus 壳多孢属Stagonospora sp. 罗氏菌属Ralstonia 枝顶孢霉菌Acremonium strictum 贪噬菌属Variovorax 大肠杆菌属Escherichia 土微菌属Pedomicrobium 赤细菌属Erythrobacter Albidiferax属 柠檬酸杆菌属 Citrobacter 黄杆菌属 Flavobacterium 橙单胞菌属 Aurantimonas 玫瑰杆菌属Roseobacter 假交替单胞菌属Pseudoalteromonas 海洋交替单胞菌属Alteromonas 海杆菌属Marinobacter 盐单胞菌属Halomonas 甲基盒菌属Methylarcula 不动杆菌属Acinetobacter 泛菌属Pantoea 磺脲类属Sulfurimonas 注：修改自Santelli et al. (2011)、Hansel and Learman (2015)、段国文等(2020). 
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