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Article(id=1274057655237763605, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20260168, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1772380800000, receivedDateStr=2026-03-02, revisedDate=null, revisedDateStr=null, acceptedDate=1774886400000, acceptedDateStr=2026-03-31, onlineDate=1781688615854, onlineDateStr=2026-06-17, pubDate=1780502400000, pubDateStr=2026-06-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1781688615854, onlineIssueDateStr=2026-06-17, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1781688615854, creator=13701087609, updateTime=1781688615854, updator=13701087609, issue=Issue{id=1274057338156769818, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='6', pageStart='2561', pageEnd='3114', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1781688540257, creator=13701087609, updateTime=1781688602467, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1274057599193486082, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1274057599193486083, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=3041, endPage=3051, ext={EN=ArticleExt(id=1274057655594279447, articleId=1274057655237763605, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Product characteristics of copper bioreduction and mineralization mediated by Shewanella sp. FeAMO from a deep-sea hydrothermal field, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

Objectives To screen copper-resistant and copper-reducing bacteria from deep-sea hydrothermal sediments and systematically analyze Cu(II) bioreduction process mediated by the bacteria, and mineralization product characteristics, thus filling the knowledge gap regarding microbial participation in copper cycling in deep-sea hydrothermal environments. Methods A strain, Shewanella sp. FeAMO, was obtained through anaerobic enrichment culture. With sodium lactate as the electron donor and Cu(II) as the terminal electron acceptor, inductively coupled plasma mass spectrometry, scanning electron microscopy-energy dispersive spectroscopy, X-ray diffraction, and X-ray photoelectron spectroscopy were employed to analyze bacterial growth, copper reduction kinetics, and physicochemical properties of the products. Results Strain FeAMO exhibited high tolerance to 400 μmol/L Cu(II), achieving a Cu(II) removal rate of 96.8% within 72 h. The mineralization products were spherical microspheres with diameters ranging from 5 to 10 µm. The bulk phase was predominant crystalline cuprous sulfide (Cu2S), while the surface was stably enriched with metallic copper (Cu0) nanoparticles, forming a core-shell heterostructure. Conclusion Shewanella sp. FeAMO can tolerate Cu(II) with copper transporters and generate Cu2S-Cu0 composite minerals by coupling copper reduction and sulfur reduction pathways. This study not only elucidates the mechanism of microbial-driven copper and sulfur cycling in hydrothermal environments but also provides a potential strategy for heavy metal immobilization and resource recovery.

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E-mail:
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These authors contributed equally to this work.

, authorsList=Fei XU, Yadong GONG, Shiping WEI, Xiang ZENG), CN=ArticleExt(id=1274057658307994147, articleId=1274057655237763605, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=海底热液区希瓦氏菌(Shewanella sp.) FeAMO介导的铜生物还原与矿化产物特性, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

目的 从海底热液区沉积物中筛选高效铜耐受和铜还原菌株，并系统解析其介导的Cu(II)生物还原过程及矿化产物特性，以填补海底热液环境微生物参与铜循环研究的空白。 方法 通过厌氧富集培养获得希瓦氏菌(Shewanella sp.) FeAMO，以乳酸钠为电子供体、Cu(II)为末端电子受体，结合电感耦合等离子体质谱(inductively coupled plasma mass spectrometry, ICP-MS)、扫描电镜-能谱(scanning electron microscopy-energy dispersive spectroscopy, SEM-EDS)、X射线衍射(X-ray diffraction, XRD)及X射线光电子能谱(X-ray photoelectron spectroscopy, XPS)等技术分析菌株生长、铜还原的动力学特征及产物理化性质。 结果 菌株FeAMO对400 μmol/L Cu(II)具有高耐受性，在厌氧条件下72 h内对Cu(Ⅱ)的去除率达96.8%。矿化产物为粒径5-10 µm的球形微团，体相以结晶硫化亚铜(Cu2S)为主，表面嵌合金属铜(Cu0)的颗粒，形成核壳式异质结构。 结论 Shewanella sp. FeAMO可耐受高浓度铜离子，通过铜离子转运蛋白外排铜离子，并耦合铜还原与硫酸盐还原生成Cu2S-Cu0复合矿物，以维持自身生长。该研究不仅阐明了热液环境微生物驱动铜硫元素循环的机制，也为重金属污染生物固定与资源化回收提供了潜在策略。

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作者贡献声明

徐菲：数据分析、结果可视化、论文撰写与修订；宫亚东：方法设计、数据分析、结果可视化、论文修改；魏世平：提供资源、课题指导；曾湘：研究概念生成、方法设计、论文撰写与修订、课题监管与指导。

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A, B: SEM images of the products collected after 2 days of incubation; C: Corresponding EDS elemental mapping overlay; D, E: SEM images of the products collected after 7 days of incubation; F: Corresponding EDS elemental mapping overlay., figureFileSmall=VkdTEIhB/P8tls/QvGU/CQ==, figureFileBig=E+mV3XjFRIhCynBNf+EIOQ==, tableContent=null), ArticleFig(id=1274088130496943093, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1274057655237763605, language=CN, label=图2, caption=菌株FeAMO生物合成产物的形貌与元素组成随培养时间的演化, figureFileSmall=VkdTEIhB/P8tls/QvGU/CQ==, figureFileBig=E+mV3XjFRIhCynBNf+EIOQ==, tableContent=null), ArticleFig(id=1274088130559857654, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1274057655237763605, language=EN, label=Figure 3, caption=Evolution of crystalline phases of the biogenic products during Cu(II) reduction by strain FeAMO. A: XRD pattern of the initial precursor CuSO4·5H2O; B: XRD pattern of the products collected after 2 days of incubation; C: XRD pattern of the products collected after 7 days of incubation., figureFileSmall=T5nm4vHUxryoR+RfjoxBLw==, figureFileBig=Ll8zPjpOEjp9PPlAKmSmEw==, tableContent=null), ArticleFig(id=1274088130647938039, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1274057655237763605, language=CN, label=图3, caption=菌株FeAMO生物还原过程中产物的晶相演化, figureFileSmall=T5nm4vHUxryoR+RfjoxBLw==, figureFileBig=Ll8zPjpOEjp9PPlAKmSmEw==, tableContent=null), ArticleFig(id=1274088130715046904, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1274057655237763605, language=EN, label=Figure 4, caption=Chemical state analysis of surface Cu (3-10 nm) in the biogenic products produced by strain FeAMO. 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海底热液区希瓦氏菌(Shewanella sp.) FeAMO介导的铜生物还原与矿化产物特性
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徐菲 1, 2 , 宫亚东 1, 3, 4 , 魏士平 3 , 曾湘 1
微生物学报 | 研究报告 2026,66(6): 3041-3051
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微生物学报 | 研究报告 2026, 66(6): 3041-3051
海底热液区希瓦氏菌(Shewanella sp.) FeAMO介导的铜生物还原与矿化产物特性
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徐菲1, 2, 宫亚东1, 3, 4, 魏士平3, 曾湘1
作者信息
  • 1.自然资源部第三海洋研究所，海洋生物遗传资源重点实验室，福建 厦门
  • 2.北京师范大学 环境学院，区域环境安全全国重点实验室，北京
  • 3.中国地质大学(北京) 海洋学院，北京
  • 4.华北有色工程勘察院有限公司，河北 石家庄
Product characteristics of copper bioreduction and mineralization mediated by Shewanella sp. FeAMO from a deep-sea hydrothermal field
Fei XU1, 2, Yadong GONG1, 3, 4, Shiping WEI3, Xiang ZENG1
Affiliations
  • 1.Key Laboratory of Marine Genetic Resources, Third Institute of Oceanography, Ministry of Natural Resources, Xiamen, Fujian, China
  • 2.National Key Laboratory of Regional Environment Safety, School of Environment, Beijing Normal University, Beijing, China
  • 3.School of Marine Sciences, China University of Geosciences (Beijing), Beijing, China
  • 4.North China Engineering Investigation Institute Co. , Ltd. , Shijiazhuang, Hebei, China
出版时间: 2026-06-04 doi: 10.13343/j.cnki.wsxb.20260168
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摘要
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目的 从海底热液区沉积物中筛选高效铜耐受和铜还原菌株，并系统解析其介导的Cu(II)生物还原过程及矿化产物特性，以填补海底热液环境微生物参与铜循环研究的空白。 方法 通过厌氧富集培养获得希瓦氏菌(Shewanella sp.) FeAMO，以乳酸钠为电子供体、Cu(II)为末端电子受体，结合电感耦合等离子体质谱(inductively coupled plasma mass spectrometry, ICP-MS)、扫描电镜-能谱(scanning electron microscopy-energy dispersive spectroscopy, SEM-EDS)、X射线衍射(X-ray diffraction, XRD)及X射线光电子能谱(X-ray photoelectron spectroscopy, XPS)等技术分析菌株生长、铜还原的动力学特征及产物理化性质。 结果 菌株FeAMO对400 μmol/L Cu(II)具有高耐受性，在厌氧条件下72 h内对Cu(Ⅱ)的去除率达96.8%。矿化产物为粒径5-10 µm的球形微团，体相以结晶硫化亚铜(Cu2S)为主，表面嵌合金属铜(Cu0)的颗粒，形成核壳式异质结构。 结论 Shewanella sp. FeAMO可耐受高浓度铜离子，通过铜离子转运蛋白外排铜离子，并耦合铜还原与硫酸盐还原生成Cu2S-Cu0复合矿物，以维持自身生长。该研究不仅阐明了热液环境微生物驱动铜硫元素循环的机制，也为重金属污染生物固定与资源化回收提供了潜在策略。

Shewanella sp. FeAMO  /  Cu(II)生物还原  /  硫化亚铜(Cu2S)  /  金属铜(Cu0)  /  生物矿化

Objectives To screen copper-resistant and copper-reducing bacteria from deep-sea hydrothermal sediments and systematically analyze Cu(II) bioreduction process mediated by the bacteria, and mineralization product characteristics, thus filling the knowledge gap regarding microbial participation in copper cycling in deep-sea hydrothermal environments. Methods A strain, Shewanella sp. FeAMO, was obtained through anaerobic enrichment culture. With sodium lactate as the electron donor and Cu(II) as the terminal electron acceptor, inductively coupled plasma mass spectrometry, scanning electron microscopy-energy dispersive spectroscopy, X-ray diffraction, and X-ray photoelectron spectroscopy were employed to analyze bacterial growth, copper reduction kinetics, and physicochemical properties of the products. Results Strain FeAMO exhibited high tolerance to 400 μmol/L Cu(II), achieving a Cu(II) removal rate of 96.8% within 72 h. The mineralization products were spherical microspheres with diameters ranging from 5 to 10 µm. The bulk phase was predominant crystalline cuprous sulfide (Cu2S), while the surface was stably enriched with metallic copper (Cu0) nanoparticles, forming a core-shell heterostructure. Conclusion Shewanella sp. FeAMO can tolerate Cu(II) with copper transporters and generate Cu2S-Cu0 composite minerals by coupling copper reduction and sulfur reduction pathways. This study not only elucidates the mechanism of microbial-driven copper and sulfur cycling in hydrothermal environments but also provides a potential strategy for heavy metal immobilization and resource recovery.

Shewanella sp. FeAMO  /  Cu(II) bioreduction  /  copper sulfide (Cu2S)  /  elemental copper (Cu0)  /  biomineralization
徐菲, 宫亚东, 魏士平, 曾湘. 海底热液区希瓦氏菌(Shewanella sp.) FeAMO介导的铜生物还原与矿化产物特性. 微生物学报, 2026 , 66 (6) : 3041 -3051 . DOI: 10.13343/j.cnki.wsxb.20260168
Fei XU, Yadong GONG, Shiping WEI, Xiang ZENG. Product characteristics of copper bioreduction and mineralization mediated by Shewanella sp. FeAMO from a deep-sea hydrothermal field[J]. Acta Microbiologica Sinica, 2026 , 66 (6) : 3041 -3051 . DOI: 10.13343/j.cnki.wsxb.20260168
正文
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铜元素含量在地球上排第26位，占地壳总量的0.006 8%左右[1]，其主要赋存状态为硫化铜矿、氧化铜矿和含铜沉积物[2]。在采矿冶炼、工业废水排放及农药施用等工业与农业活动中存在大量的铜污染[3]。高浓度Cu(II)可改变细胞膜通透性[4]、干扰酶活性[5]，并诱导活性氧积累[6]，进而影响细胞代谢稳态和生理功能，对生态环境构成重大威胁。
微生物介导的金属氧化还原过程在生物冶金[7]、环境修复[8]、污水处理[9]及资源回收[10]等领域展现出巨大潜力。尽管微生物对铁、锰等金属的还原过程与机制已有较为深入的研究，但对于微生物介导的Cu(II)还原过程及其生物矿化产物和调控机制的认知仍相对有限。近年来，部分研究揭示了微生物还原铜的路径。Kimber等[11]发现，硫还原地杆菌(Geobacter sulfurreducens)还原Cu(II)后在细胞表面生物矿化形成粒径均一的硫化亚铜(Cu2S)纳米颗粒。Gracioso等[12]从铜矿中分离获得芽孢杆菌(Bacillus sp.) 105，该菌可将有毒的铜离子在胞内转化为稳定的单原子铜(Cu0)，并推测了潜在的铜转运分子机制。由发酵地线菌(Geothrix fermentans)、限食固氮螺形菌(Azospira restricta)和寡养纤维单胞菌(Cellulomonas oligotrophica)为主的铜矿富集培养菌群也被证实能够还原Cu(II)并形成单质铜纳米颗粒[13]。希瓦氏菌属(Shewanella sp.)具有显著的代谢多样性，可利用Fe(III)、Mn(III, IV)、U(VI)、Cr(VI)、Pu(IV)及Pd(II)等多种金属作为末端电子受体，在多种关键金属的地球化学循环中发挥重要作用[14-15]。奥奈达湖希瓦氏菌(Shewanella oneidensis)被报道具有还原Cu(II)合成单质铜纳米颗粒和Fe-S颗粒的能力，但该过程不依赖于Mtr铁还原酶系[16-17]。Luong等[18]利用Shewanella sp. HN-41在生物电化学装置中将铜离子还原为铜氢氧氯盐矿物[Cu2Cl(OH)3]等微米级球形颗粒。
海底热液系统富含金属硫化物，是海洋中重要的金属富集环境[19]。在地壳岩石中，铜含量随岩性差异较大，通常在2-105 mg/kg之间(平均63 mg/kg)，其中碳酸盐岩中铜含量较低，而页岩等沉积岩中则相对富集[20]。相比之下，海底热液沉积物中的铜含量显著升高，可达141.15-551.58 mg/kg[21]，而热液硫化物中的铜含量甚至高达9.6-228.0 g/kg[22]。Yu等[23]分离了多株来源于海底热液区的属于γ-变形菌纲的高耐铜微生物，这些微生物通过分泌胞外聚合物吸附铜，并利用铜离子转运的相关蛋白外排铜。然而，海底热液区微生物参与金属铜还原过程的研究仍较为缺乏。基于此，本研究以从西北印度洋热液沉积物中分离的铁还原菌Shewanella sp. FeAMO为研究对象，结合电感耦合等离子体质谱(inductively coupled plasma mass spectrometry, ICP-MS)、扫描电镜-能谱(scanning electron microscopy-energy dispersive spectroscopy, SEM-EDS)、X射线衍射(X-ray diffraction, XRD)及X射线光电子能谱(X-ray photoelectron spectroscopy, XPS)等多种表征手段，分析该菌株以Cu(II)为末端电子受体形成Cu-S矿物的生物还原过程，以及矿物的晶体结构、元素组成、化学价态和形貌结构等特性，并初步推测FeAMO矿化的产物形成机制。本研究有助于深化对希瓦氏菌属参与深海热液区铜生物地球化学循环的认识，以期为基于微生物的铜污染修复与资源回收提供菌种资源和科学依据。
1 材料与方法
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1.1 微生物分离与鉴定
实验所用菌株FeAMO (MCCC M27737)分离自西北印度洋卧蚕热液区(6°22′S, 60°32′E) 2 995 m深处的热液沉积物样品。该样品由“向阳红09”船实施的中国大洋38航次中“蛟龙”号利用pushcore采样器于2017年3月14日采集。该菌株分离自铁还原FRPFO培养基[24]，为兼性厌氧、具铁还原能力的革兰氏阴性细菌，目前已保藏于中国海洋微生物保藏管理中心(MCCC, https://www.mccc.org.cn/)。
1.2 培养与生长测定
所有铜还原微生物培养实验均在严格定义的CuRL液体厌氧培养基中进行，其基础组分(g/L)：NaCl 25.0，MgCl2·6H2O 4.2，KCl 0.5，NH4Cl 0.25，CaCl2·2H2O 0.7，K2HPO4 0.14，MgSO4·7H2O 3.4，PIPES 6.05。以40 mmol/L乳酸钠作为碳源与电子供体，并添加刃天青(1 g/L)作为厌氧指示剂。将20 mL培养基分装于50 mL厌氧血清瓶中，用N2:CO2 (80:20，体积比)混合气体吹扫20 min以去除氧气，随后用丁基橡胶塞密封，并于121 ℃灭菌20 min。灭菌冷却后，向每瓶培养基中补加过滤除菌的下列组分：NaHCO3 2.5 g/L，CuSO4·5H2O 0.1 g/L (400 µmol/L) (作为末端电子受体)，L-半胱氨酸0.3 g/L，以及微量元素溶液10 mL/L和维生素溶液1 mL/L。
使用处于对数生长期的预培养物进行接种，使初始OD600为0.1。接种后，所有培养物置于28 ℃恒温条件下静置培养7 d。为监测动态过程，分别于培养后的第1-7天，使用无菌注射器取样1 mL，用于后续菌体生长与产物分析。
采用LIVE/DEAD® BacLightTM细菌活力试剂盒(ThermoFisher Scientific公司)进行活菌计数。取9 µL菌液样品，与1 µL按说明书配制的染色工作液充分混合，于室温避光孵育20-30 min。孵育结束后，取10 µL染色菌液于荧光显微镜(Nikon公司)下，使用血细胞计数板进行直接镜检计数。所有生长与计数实验均设3个独立的生物学重复。
1.3 铜浓度定量分析
取不同时间点培养液，10 000 r/min离心5 min去除菌体。上清液经0.22 μm滤膜过滤后，采用电感耦合等离子体质谱(ICP-MS，ThermoFisher Scientific公司)测定溶解态总铜浓度。仪器工作条件：RF功率1 400 W，雾化气流量1.08 L/min，碰撞反应池模式(kinetic energy discrimination, KED)，以Rh作为内标进行定量校正。每个样品测定3次。
1.4 菌体形态与表面元素分析
为观察菌体在Cu(II)胁迫下的形态变化及胞外矿化产物，分别对培养至第2天与第7天的产物透析3次后收集，真空冷冻干燥，取少量溶解于无水乙醇中。将上述样品滴至硅片，临界点干燥后喷镀铂金，利用SEM观察矿物形态及元素分布，观察电压为5-15 kV。
1.5 产物的晶型与价态表征
为明确生物合成产物的矿物相与化学状态，分别进行XRD和XPS分析。与SEM制样相类似，对不同培养时间的产物透析3次后，真空冷冻干燥。XRD分析时，将真空冷冻干燥后的样品采用X射线衍射仪(Rigaku公司)进行分析，采用单色Cu靶Kα辐射，所加电流80 mA，电压40 kV，采集范围3°-70°，精度1.2 (°)/min，时间为6 s。XPS分析时，干燥样品采用X射线光电子能谱仪(Kratos公司)进行分析，激发源为单色化Al Kα (1 486.6 eV)，结合能以C 1s (284.8 eV)校准。XPS作为一种表面敏感的表征技术，其信息深度约为3-10 nm。对Cu 2p、O 1s、C 1s等核心能级进行高分辨扫描，并通过分峰拟合分析铜的化学价态。
1.6 基因组测序和注释
菌株FeAMO全基因测序由上海美吉生物医药科技有限公司完成，采用Illumina NovaSeq 6000平台开展双端150 bp测序(测序深度≥100×)。采用Fastp (v0.20.0)和SOPAdenovo (v2.04)软件进行序列质控和测序数据组装，使用QUAST (v5.0.2)评估组装质量。基因组注释采用Prokka (v1.13)自动化流程，使用Prodigal (v2.6)进行编码序列(coding sequence, CDS)预测，通过BLASTp (v2.7)将推断出的蛋白质序列与UniProtKB/Swiss-Prot细菌参考数据库进行比对，预测的目标蛋白质序列进一步在KEGG数据库进行BLASTp比对以获取代谢通路信息。基因组序列存储于GenBank数据库(https://www.ncbi.nlm.nih.gov/genbank/) [序列号 JBSJYJ010000002 (Chromosome)，JBSJYJ010000001 (Plasmid A)][25]。
2 结果与分析
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2.1 Shewanellasp. FeAMO的Cu(Ⅱ)还原过程
以乳酸钠为电子供体和碳源，采用250 mg/L (400 µmol/L) CuSO4作为唯一末端电子受体的培养体系中，FeAMO展现出显著的Cu(Ⅱ)还原能力(图1)。培养2 d后，培养基颜色由微蓝色逐渐变为橙红色，至第7天时变为深棕黑色；而无菌对照组在相同条件下未见颜色变化(图1A)，说明颜色转变与细菌介导的铜还原及产物沉淀密切相关。利用ICP-MS对上清液中溶解态Cu离子浓度进行监测，结果显示，在培养前3 d内，溶解态铜浓度由初始的250 mg/L (400 µmol/L)迅速降至8 mg/L (32 µmol/L)，去除率达96.8% (图1B)，表明菌株具有高效还原Cu(Ⅱ)的能力。溶解态Cu(Ⅱ)浓度显著降低后，菌株FeAMO的细胞数于第4天达到峰值(1.03×107 cells/mL)，随后进入稳定期(图1B)。
2.2 Shewanellasp. FeAMO铜还原产物的形貌变化与元素组成
采用SEM-EDS对培养第2天与第7天的菌体及其周围沉淀进行了形貌与元素分析(图2)。SEM图像显示(图2A、2D)，菌体胞外区域附着有大量粒径不均、表面光滑的球形颗粒。粒径统计表明(图2B、2E)，这些颗粒的直径主要分布在5-10 µm之间。进一步的EDS面扫描分析结果(图2C、2F)显示，铜(Cu)与硫(S)元素的特征X射线信号在颗粒区域高度重叠且连续分布，初步表明其为铜-硫化合物。为进行准确定量，对多个代表性颗粒进行了点分析(定量标准：Cu标样为纯Cu，S标样为FeS2)。结果显示，培养第2天产物的Cu、S含量(以质量百分比计)分别在45.23%-74.88%与12.06%-25.12%之间；至第7天，其含量范围趋于稳定，分别为46.71%-56.48% (Cu)与17.16%-22.81% (S)。
2.3 Shewanellasp. FeAMO铜还原产物的矿物相分析
采用XRD对不同反应阶段产物的体相晶体结构进行了分析(图3)。与CuSO4·5H2O的XRD谱图(图3A)相比，还原2 d后的产物谱图(图3B)在27.4°、31.7°、45.5°和53.9°处的衍射峰与Cu2S (PDF#84-1770)的(111)、(200)、(220)及(311)晶面相符；同时，在24.71°、26.59°、29.76°、31.20°、34.06°、35.35°、36.62°、46.84°、48.90°及54.62°等处还观察到一系列衍射峰，与Cu7S4 (PDF#23-0958)的特征峰基本吻合。上述结果表明，反应2 d后的产物为Cu2S与Cu7S4的混合相。
当还原时间延长至7 d时，产物的XRD谱图(图3C)中仅保留Cu2S对应的特征衍射峰(27.4°、31.7°、45.5°、53.9°)，而Cu7S4的衍射峰完全消失。这表明随着反应时间的增加，中间相Cu7S4被完全消耗，最终产物的主要体相晶体结构为纯相Cu2S。
2.4 Shewanellasp. FeAMO铜还原产物的元素价态分析
采用XPS分析了还原产物表面的铜的化学价态(图4)。通过对Cu 2p谱图进行分峰拟合，定量分析了不同价态铜的相对含量。对于培养2 d的产物表面(图4B)，其Cu 2p谱图在结合能为931.9 eV和951.8 eV处出现特征峰，分别归属为金属铜(Cu0)的2p3/2和2p1/2能级；同时在932.9 eV和952.7 eV处观察到归属于一价铜[Cu(Ⅰ)]的对应信号。拟合计算表明，Cu0与Cu(Ⅰ)的表观原子含量比例为76.27%:23.73%。为确证这一价态分布，进一步分析了样品的Cu LMM俄歇谱(图4E)。该图谱在动能约918.3 eV和914.0 eV处分别出现Cu0和Cu(Ⅰ)的特征信号，进一步支持产物表面中铜以零价与一价共存。培养7 d后的产物表面表现出相似的价态特征(图4C、4F)。其Cu 2p谱图中，Cu0 (结合能：932.3、952.1 eV)与Cu(Ⅰ) (结合能：933.1、953.2 eV)的信号清晰可见，两者比例约为76.77%:23.23%。其对应的Cu LMM俄歇谱(图4F)中Cu0的特征峰位(918.6 eV)与2 d样品表面(918.3 eV)基本一致，而Cu(Ⅰ)的峰位则从914.0 eV略微移至917.5 eV，这可能反映了局部化学环境的细微变化。更重要的是，对LMM俄歇谱进行分峰面积积分发现，与第2天样品相比，第7天样品其矿物表面中Cu0组分的相对比例略有下降，而Cu(Ⅰ)的比例相应增加。
综合Cu 2p与Cu LMM谱的分析结果可知，在培养过程中，由菌株FeAMO还原CuSO4形成的固体沉淀表明始终以Cu0与Cu(Ⅰ)共存为特征且以金属态为主导，与一价态铜稳定共存。
2.5 Shewanellasp. FeAMO基因组功能注释
基因组注释结果显示[已上传ScienceDB数据库(https://www.scidb.cn/c/j00231)，CSTR编号为31253.11.sciencedb.j00231.00057]，FeAMO基因组中携带多个与重金属抗性、铁还原及硫酸盐代谢相关的功能基因。其中，耐铜相关基因包括copA、copR及cus操纵子(cusA、cusB、cusR、cusS)；耐钴/锌/铬基因包括cusA、cusB、cusC、cusD；铁还原相关基因主要涉及mtr操纵子(mtrA、mtrB、mtrC、mtrD、mtrE、mtrF、mtrH)、omcA、cymA及doxD；此外，还注释到同化硫酸盐还原相关基因cysC、cysD、cysN、cysQ、cysH、cysJ、cysI及sirA。
3 讨论
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Cu(II)作为高毒性重金属离子，在较低浓度时即可对多数微生物产生抑制与氧化应激效应[26]。本研究表明，菌株FeAMO对高浓度Cu(II)表现出显著的耐受性与还原能力。该菌株能够在初始Cu(II)浓度高达400 µmol/L的条件下生长，其耐受阈值高于已报道的G. sulfurreducens (<100 µmol/L)[11]和多种希瓦氏菌株[27] (S. oneidensis MR-1以及Shewanella sp. MB4、FB18、FS8等)。其中菌株FB18和FS8在Cu(II)浓度为150 µmol/L时即停止生长(OD600≈0)，菌株MR-1和MB4在Cu(II)浓度为400 µmol/L条件下OD600降至0[27]。这表明菌株FeAMO可能通过高效的胞外排铜机制(如Cus系统)[28]、强大的细胞膜修复能力[29]或抗氧化防御系统[30]，使其能够在高毒性环境中生存。
从还原过程来看，菌株FeAMO的生长与Cu(II)还原趋势高度同步(图1B)，表明该还原过程是与能量代谢相耦联的活跃生理活动，而非被动吸附。菌株FeAMO在400 µmol/L Cu(II)条件下72 h内去除率达到96.8%，其还原效率显著高于多数已报道的微生物金属生物还原过程。例如，G. sulfurreducens在5 μmol/L与50 μmol/L Cu(II)条件下最大去除率分别为80%和63%[11]；即使是还原能力较强的模式菌株S. oneidensis MR-1，在50 μmol/L Cu(II)条件下也需要24 h和96 h才分别达到91%和100%的去除率[16]。XRD与SEM-EDS结果表明，菌株FeAMO的铜还原矿物由CuSO4·5H2O经Cu2S与Cu7S4的混合相转化为结晶度良好的单一Cu2S相(辉铜矿) (图2-3)。然而，产物表层(~3-10 nm)的XPS结果显示，在主要体相矿物Cu2S表层存在明显的金属铜(Cu0)信号和一价铜[Cu(Ⅰ)]信号(图4)。推测产物是一种“核壳”或“镶嵌”式复合结构，其体相核心为结晶Cu2S，而表面镶嵌着纳米尺度金属铜(Cu0)颗粒。该结构与已有结果类似，部分微生物可以生成富铜中间相Cu7S4[31]，而希瓦氏菌(Shewanella sp.)可生成铜纳米颗粒以及Cu-S固相沉积物[16-17]。此外，磁鞭毛菌Magnetospirillum magneticum在特定培养条件下可合成Fe3O4或Fe3S4核被Sm2O3壳包覆的核壳纳米颗粒[32]，说明微生物矿化过程中能够形成不同矿物相共存的复合结构。
另外，菌株FeAMO介导的Cu(II)生物矿化过程可能存在铜还原过程和硫还原过程(图5)。基于对菌株FeAMO的基因组(GenBank登录号为JBSJYJ010000002)分析(CSTR编号为31253.11.sciencedb.j00231.00057)，结果发现该菌株编码了CopA P-type ATPase及RND外排泵家族复合物，包括铜外排泵操作子CusCBA和CusRS，以及钴-锌-镉(Co-Zn-Cd)外排泵操纵子CzcCBA及其调控因子CzcD，可将过量的铜离子主动排出至胞外或周质空间。同时，内膜醌池(quinol pool)中的电子经由内膜醌氧化还原酶(quinone oxidoreductase)传递至周质蛋白MtrA，并进一步通过Mtr途径(MtrA-MtrB-MtrC/OmcA复合物)跨越外膜传递至细胞表面。结合文献所示希瓦氏菌外膜电子传递系统(如MtrCAB体系)在金属还原中的作用[33-35]，推测在胞外，Cu(II)接受电子被还原为Cu(I)或Cu0。另一方面，在菌株FeAMO的基因组中未发现编码异化硫酸盐还原途径，但注释到编码完整同化硫酸盐还原途径(cysCDNQHJ基因簇和sir基因)，具备将SO42-还原为H2S的遗传基础，其缺乏cysK和cysE基因，减少了胞内半胱氨酸合成，大部分S2-被转运或扩散作用至胞外，极易与溶液中的Cu(I)反应，这为形成Cu-S矿物提供了直接的硫源。菌株FeAMO来源于热液区沉积物，该环境富含金属与硫化物，该过程可能模拟了深海热液系统中Cu-S矿物沉积的微生物学路径，具有潜在地质指示意义[36]。
4 结论
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本研究表明，菌株FeAMO能够耐受高浓度铜离子，并以Cu(Ⅱ)作为唯一末端电子受体实现高效生物还原。在乳酸钠作为电子供体和碳源的条件下，该菌株可在72 h内去除96.8%的Cu(Ⅱ)并生成固相沉积物，显示出较强的铜胁迫适应能力与金属转化效率。多尺度表征结果显示，还原产物呈球形微团状沉积，是一种以结晶Cu2S为体相、表面镶嵌金属铜(Cu0)颗粒的“核壳”复合结构。该过程对于解析微生物参与海底热液区的铜/硫元素循环，为揭示金属硫化矿物的微生物介导沉积机制提供了新的研究线索，同时对铜污染环境的原位固化、废铜资源生物回收及生物冶金构建具有潜在应用价值。
基金
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  • 国家重点研发项目(2021YFF0501304)
  • 国家自然科学基金(42576120)
文献
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2026年第66卷第6期
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doi: 10.13343/j.cnki.wsxb.20260168
  • 接收时间：2026-03-02
  • 首发时间：2026-06-17
  • 出版时间：2026-06-04
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  • 收稿日期：2026-03-02
  • 录用日期：2026-03-31
基金
the National Key Research and Development Program of China(2021YFF0501304)
国家重点研发项目(2021YFF0501304)
the National Natural Science Foundation of China(42576120)
国家自然科学基金(42576120)
作者信息
    1.自然资源部第三海洋研究所，海洋生物遗传资源重点实验室，福建 厦门
    2.北京师范大学 环境学院，区域环境安全全国重点实验室，北京
    3.中国地质大学(北京) 海洋学院，北京
    4.华北有色工程勘察院有限公司，河北 石家庄
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https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20260168
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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