Sulfur Isotope Fractionation Mediated by Microbial Anoxygenic Photosynthetic Sulfur Oxidation Processes and Its Geological Implications
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摘要: 微生物在利用含硫物质时的同位素偏好性会导致代谢产物中硫同位素的分馏，因此地质记录中的硫同位素可以用来反演其中的微生物活动以及古海洋和大气的氧化还原条件. 对微生物参与的硫循环的传统认知中，只有微生物介导的硫还原作用和硫歧化作用会导致明显的同位素分馏现象，而微生物硫氧化过程造成的分馏效应不明显. 而最近的研究发现一株硫氧化细菌可以产生巨大的硫同位素分馏，意味着我们需要重新评估地质记录中的硫氧化过程. 综述了微生物参与的不产氧光合硫氧化过程中硫同位素分馏效应及其地质学意义，包括硫氧化微生物及绿弯菌的分布和功能、微生物介导硫氧化过程的硫同位素分馏效应、以及微生物硫氧化过程硫同位素分馏研究的地质记录. 最后对微生物参与的不产氧光合硫氧化过程中硫同位素分馏效应研究现状和未来发展方向提出总结和展望.Abstract: The isotopic preferences exhibited by microorganisms during the utilization of sulfur-containing materials result in sulfur isotopic fractionation in metabolites, Consequently, sulfur isotopes in the geological record can serve as indicators of microbial activity and redox conditions in the paleo-oceans and atmospheres. Conventionally, significant isotopic fractionation has been associated with microbially mediated sulfur reduction and sulfur disproportionation, while the fractionation effect caused by microbial sulfur oxidation processes has considered less evident. However, recent research has shown that one of sulfur-oxidizing bacteria can produce substantial sulfur isotopic fractionation, necessitating a reassessment of sulfur oxidation processes in the geological record. This paper provides a comprehensive review of sulfur isotopic fractionation effects in microbial anoxygenic photosynthetic sulfur oxidation processes and its geological implications, which includes the distribution and function of sulfur-oxidizing microorganisms and Chloroflexota, the sulfur isotopic fractionation effect of microbially mediated sulfur oxidation processes, and the geological record of sulfur isotopic fractionation studies related to microbial sulfur oxidation processes. Finally, the current status and future directions of research on sulfur isotopic fractionation mediated by microbial anoxygenic photosynthetic sulfur oxidation processes are summarized and presented.
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图 1 自然界无机硫循环的主要转换路径
实线代表硫酸盐还原过程，中间态硫歧化过程和还原态硫的氧化过程；虚线表示平衡过程形成多聚硫化物；本图基于（Canfield，2001）修改
Fig. 1. Outlined here are the principal pathways of inorganic sulfur-compound transformations in nature
图 2 绿弯菌门系统发育树
该树由基于16S rRNA基因的最大似然法构建，以Thermodesulfobium narugense DSM14796T作为树根. 右上角展示了绿弯菌门不同纲的系统发育关系和形态学特征. 黄色阴影区域表示绿弯菌纲的光合类群
Fig. 2. Phylogenetic tree of the phylum Chloroflexota
图 3 腾冲火山地热区热泉菌席，环境温度为50 ℃
图中橙黄色区域为绿弯菌，绿色区域为蓝藻
Fig. 3. Hot spring mats of Tengchong volcanic-geothermal area at ambient temperature of 50 ℃
表 1 主要不产氧光合细菌菌株介导硫氧化过程产生的硫同位素分馏特征
Table 1. Characteristic of sulfur isotope fractionation during the sulfur-oxidation process mediated by major photosynthetic sulfur-oxidizing bacterial strains
底物→产物 菌株 种属 温度(℃) pH ε产物—底物 (‰) 参考文献 H2S→S0 Chlorobium tepidum. GSB 48 7.0 1.8 Zerkle et al.(2009) S0→SO42- 48 7.0 -3.3~0 H2S→S0 Chromatium vinosum PSB 35 8.1 2.4 Howard Gest and Hayes(1984) S0→SO42- 35 8.1 0.2 SO32-→SO42- Chromatium vinosum PSB 35 8.1 5.0 Fry et al.(1985) H2S→SO42- Allochromatium vinosum PSB 25 7.0 0.1 Brabec et al.(2012) H2S→S0 Ectothiorhodospira shaposhnikovii PSB 28 8.2 2.2 Ivanov et al.(1976) H2S→SO42- 28 8.2 0.7~2.0 H2S→SO42- Chlorobaculum tepidum GSB 45 7.0 0.1 Brabec et al.(2012) 注：GSB. 绿硫细菌（green sulfur bacteria）；PSB. 紫硫细菌（purple sulfur bacteria) 
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