微生物学报

Bacteria type	Representative species	Mineral products	Mineralization mechanism	References
Urease-producing bacteria (UPB)	Sporosarcina pasteurii, Staphylococcus sp. H6, Serratia marcescens, Enterobacter cloacae EMB19	Calcite, aragonite, Cd-Ca co-precipitates	Urease-catalyzed urea hydrolysis produces NH₃ and CO₃^2-, elevating local pH and inducing precipitation of CaCO₃ and metal ions (e.g., Cd-Ca co-precipitates)	[50-52]
Sulfate-reducing bacteria (SRB)	Desulfovibrio desulfuricans, Desulfobulbus propionicus	CdS, PbS	Sulfate-reducing bacteria reduce SO₄^2- to S^2- using lactate as an electron donor, decreasing local redox potential and inducing metal sulfide precipitation	[53]
Phosphate-solubilizing bacteria (PSB)	Burkholderia ambifaria QY14	Cd-HAP	Increased acid phosphatase activity hydrolyzes organic phosphates, releasing PO₄^3- that combines with Cd²⁺ and Ca²⁺ to precipitate as Cd-HAP	[54]
Nitrate-reducing bacteria (NRB)	Klebsiella aerogenes	Green rust, goethite	Nitrate-reducing bacteria reduce NO₃⁻ to NO₂⁻ using organic carbon as an electron donor. The produced NO₂⁻ non-enzymatically oxidizes Fe²⁺ to Fe³⁺, inducing Fe²⁺/Fe³⁺ co-precipitation as green rust and goethite	[55]
Fe-oxidizing bacteria (FOB)	Ochrobactrum sp. EEELCW01	Goethite, Fe-As minerals	Iron-oxidizing bacteria oxidize Fe²⁺ to Fe³⁺ under nitrate-reducing conditions, promoting Fe³⁺ mineral formation (e.g., goethite) and co-precipitation with As to form Fe-As minerals	[56]
Fe-reducing bacteria (FRB)	Shewanella oneidensis MR-4	Siderite (FeCO₃)	Iron-reducing bacteria reduce Fe³⁺ to Fe²⁺, which combines with CO₃^2- to form siderite	[57]

Bacteria type	Representative species	Mineral products	Mineralization mechanism	References
Urease-producing bacteria (UPB)	Sporosarcina pasteurii, Staphylococcus sp. H6, Serratia marcescens, Enterobacter cloacae EMB19	Calcite, aragonite, Cd-Ca co-precipitates	Urease-catalyzed urea hydrolysis produces NH₃ and CO₃^2-, elevating local pH and inducing precipitation of CaCO₃ and metal ions (e.g., Cd-Ca co-precipitates)	[50-52]
Sulfate-reducing bacteria (SRB)	Desulfovibrio desulfuricans, Desulfobulbus propionicus	CdS, PbS	Sulfate-reducing bacteria reduce SO₄^2- to S^2- using lactate as an electron donor, decreasing local redox potential and inducing metal sulfide precipitation	[53]
Phosphate-solubilizing bacteria (PSB)	Burkholderia ambifaria QY14	Cd-HAP	Increased acid phosphatase activity hydrolyzes organic phosphates, releasing PO₄^3- that combines with Cd²⁺ and Ca²⁺ to precipitate as Cd-HAP	[54]
Nitrate-reducing bacteria (NRB)	Klebsiella aerogenes	Green rust, goethite	Nitrate-reducing bacteria reduce NO₃⁻ to NO₂⁻ using organic carbon as an electron donor. The produced NO₂⁻ non-enzymatically oxidizes Fe²⁺ to Fe³⁺, inducing Fe²⁺/Fe³⁺ co-precipitation as green rust and goethite	[55]
Fe-oxidizing bacteria (FOB)	Ochrobactrum sp. EEELCW01	Goethite, Fe-As minerals	Iron-oxidizing bacteria oxidize Fe²⁺ to Fe³⁺ under nitrate-reducing conditions, promoting Fe³⁺ mineral formation (e.g., goethite) and co-precipitation with As to form Fe-As minerals	[56]
Fe-reducing bacteria (FRB)	Shewanella oneidensis MR-4	Siderite (FeCO₃)	Iron-reducing bacteria reduce Fe³⁺ to Fe²⁺, which combines with CO₃^2- to form siderite	[57]

细菌介导的矿化机制及其在生物医学应用的研究进展

PDF下载

聂文瑶 ¹ , 任彪 ² , 李继遥 ¹

微生物学报 | 综述 2026,66(2): 578-594

收起

微生物学报 | 综述 2026, 66(2): 578-594

细菌介导的矿化机制及其在生物医学应用的研究进展

全屏

聂文瑶¹, 任彪², 李继遥¹

作者信息

^1.四川大学华西口腔医院，牙体牙髓病科，口腔疾病防治全国重点实验室，国家口腔医学中心，口腔疾病国家临床医学研究中心，四川成都

^2.四川大学华西口腔医院，口腔疾病防治全国重点实验室，口腔疾病国家临床医学研究中心，四川成都

Mechanisms of bacteria-mediated mineralization and recent advances in their biomedical applications

Wenyao NIE¹, Biao REN², Jiyao LI¹

Affiliations

^1.State Key Laboratory of Oral Diseases & National Center for Stomatology, National Clinical Research Center for Oral Diseases, Department of Cariology and Endodontics, West China Hospital of Stomatology, Sichuan University, Chengdu, Sichuan, China

^2.State Key Laboratory of Oral Diseases, National Clinical Research Center for Oral Diseases, West China Hospital of Stomatology, Sichuan University, Chengdu, Sichuan, China

出版时间: 2026-02-04 doi: 10.13343/j.cnki.wsxb.20250652

文章导航

摘要

收起

近年来，微生物介导的矿化作为自然界中广泛存在的生物矿化过程已成为研究热点。该过程不仅深刻影响着自然界矿物的形成与地球元素循环，还参与生物体内矿物的形成，具有重要的生态学和生物学意义。细菌因代谢活性强、环境适应范围广，在微生物介导的矿化中极具代表性。本文综述了细菌介导的矿化机制及其在生物医学领域的应用，重点阐释了细菌控制矿化、细菌诱导矿化、细菌影响矿化这3种主要机制，探讨了其在医学成像、靶向治疗、组织工程等方向的应用潜力，旨在为相关研究与实践应用提供有益参考和科学启示。

关键词

细菌介导矿化 / 生物矿化 / 矿化机制 / 生物医学应用

Abstract

收起

In recent years, microbially mediated mineralization, a widespread form of biomineralization in nature, has emerged as a research hotspot. This process not only exerts profound influences on mineral formation and global biogeochemical cycling but also contributes to mineral deposition within living organisms, thereby holding significant ecological and biological importance. Among microorganisms, bacteria—characterized by high metabolic activity and remarkable environmental adaptability—represent the most prominent agents in microbial mineralization. This review summarizes the mechanisms of bacteria-mediated mineralization and their applications in the biomedical field, with a particular emphasis on three principal mechanisms: bacteria-controlled mineralization, bacteria-induced mineralization, and bacteria-influenced mineralization. Furthermore, the potential applications of these processes in medical imaging, targeted therapy, and tissue engineering are discussed. The overarching aim is to provide valuable references and scientific insights to inform future research and facilitate their translation into practical applications.

Key words

bacteria-mediated mineralization / biomineralization / mineralization mechanisms / biomedical applications

引用本文

聂文瑶, 任彪, 李继遥. 细菌介导的矿化机制及其在生物医学应用的研究进展. 微生物学报, 2026 , 66 (2) : 578 -594 . DOI: 10.13343/j.cnki.wsxb.20250652

Wenyao NIE, Biao REN, Jiyao LI. Mechanisms of bacteria-mediated mineralization and recent advances in their biomedical applications[J]. Acta Microbiologica Sinica, 2026 , 66 (2) : 578 -594 . DOI: 10.13343/j.cnki.wsxb.20250652

正文

收起

生物矿化(biomineralization)指有机生物体调控无机矿物形成的过程^[1]。这一现象广泛存在于自然界，包括真核生物，如植物硅质体的沉积^[2]、动物骨骼的形成^[3]以及微生物介导的矿物沉淀行为等。作为生物矿化的重要组成部分，微生物介导的矿化是指微生物在代谢过程中，于细胞内外形成无机矿物沉淀的过程^[4]。矿物-微生物的相互作用在环境变化、元素的生物地球循环以及矿床的形成中发挥着重要作用^[5]。其中，细菌因代谢活性强、环境适应性广，在微生物介导的生物矿化中最具代表性^[6]。

近年来，细菌介导矿化的相关研究受到广泛关注。在自然环境中，细菌活动驱动矿化，形成了微生物岩，它是稳定生态系统与记录古环境信息的关键物质载体^[7]；细菌还通过诱导碳酸盐类矿物沉淀，将无机碳固定为矿物碳，这对维持生态系统稳定运行、驱动地质沉积过程和塑造地球表层环境具有重要意义^[5]。此外，细菌介导的矿化行为也存在于生物体内。例如，海绵内的共生钙化细菌可在海绵表面生成钙化小球^[8]，模拟钙质或硅质骨骼的外围结构，从而保护宿主免受外界损伤；在人体组织中，如感染性上尿路结石^[9]和牙结石^[10-11]，也可观察到细菌矿化的痕迹。随着研究的推进，细菌矿化机制得到更深入的认识。

同时，细菌介导的矿化在生物医学领域的应用前景也逐渐显现。细菌矿化产物具有良好的生物相容性与精确的结构可控性，为精准医学带来新的技术契机^[12]。本文就细菌介导的矿化机制及其在生物医学领域的应用进展进行介绍，重点阐释细菌控制矿化(bacteria-controlled mineralization)、细菌诱导矿化(bacteria-induced mineralization)及细菌影响矿化(bacteria-influenced mineralization) 3种主要机制，并进一步探讨该机制在医学成像、靶向治疗、组织工程等方向的应用潜力，以期为相关研究与实践应用提供有益参考和科学启示。

1 细菌介导的矿化机制

收起

生物矿化过程可基于生物体对矿化的调控程度划分为生物控制矿化(biologically controlled mineralization, BCM)和生物诱导矿化(biologically induced mineralization, BIM) 2种主要作用模式^[13-14]。随着对细胞表面等非代谢性结构在矿化中作用机制的深入认识，部分学者进一步将这部分过程独立出来，定义为生物影响矿化(biologically influenced mineralization, BIFM)^[15]。因此，现有研究中既有两分法的框架^[5]，也有三分法的体系^[16]。三分法进一步细化了细胞表面结构的化学效应在生物矿化中的作用。

随着对微生物矿化研究的深入，细菌介导的矿化也呈现出从严格控制到被动影响的多样表现。上述术语并非专用于微生物，还可用于描述动物、植物等多类生物的矿化行为^[2-3]。考虑到本文的研究对象为细菌介导的矿化过程，因此本文分别使用“细菌控制矿化” “细菌诱导矿化”与“细菌影响矿化”术语以明确“生物”的指向。

1.1 细菌控制矿化

细菌控制矿化是指细菌通过遗传调控和代谢途径严格控制晶体的类型、尺寸、形貌及空间位置的过程，其中特定的细胞器(如离子泵和通道)常用于介导离子的转运^[4]，促使矿物在细胞内部的囊泡中形成(图1)^[5]。这些矿化产物通常表现出高度有序的晶体结构，并具备一定的生物功能^[17-18]。这一理论基础源于“生物控制矿化”，该术语最早由Mann于1983年提出，用于补充Lowenstam所定义的“有机基质介导矿化(organic matrix-mediated mineralization)”^[13,19]。Lowenstam主要关注真核生物(如贝类、脊椎动物)通过构建有机框架(如囊泡或基质)引导晶体的定向沉淀，控制矿物的类型、形貌和微观结构，形成骨骼与壳体的过程^[19]。Mann在此基础上提出“生物控制矿化”作为更全面的定义，其核心是细胞活动主动调控矿物的成核、生长、形态及空间定位；早期的生物控制矿化研究主要聚焦于真核生物，其矿化过程通常在生物体严格调控的局部环境中进行，且可发生在不同空间位置^[20]。在真核生物中，这种受控矿化多发生在细胞外基质上，由细胞通过囊泡转运离子或分泌有机基质控制晶体的定向沉积，例如骨、牙、贝壳的形成^[21-22]。此外，一些单细胞真核生物也能在细胞内部进行受控矿化，例如硅藻在胞内囊泡内形成特征性的硅质骨架^[23]。由于原核生物细胞结构较为简单，其胞外基质主要用于黏附或调节局部微环境，可能难以实现严格的晶体成核和排列。若要实现类似的控制矿化，通常需要依赖胞内膜性结构形成受限的微环境^[24-25]。因此，细菌控制矿化在概念上继承了“细胞主动控制矿化”的本质内涵，但在机制上可能演化出以胞内囊泡为核心的控制策略，其典型代表为趋磁细菌(magnetotactic bacteria, MTB)形成磁小体的过程。

1975年，Blakemore^[26]首次在显微镜下观察到趋磁细菌沿着地球磁场移动。大多数已知的MTB分离株为革兰氏阴性菌，广泛分布于海洋、淡水和沉积物环境中。这些细菌具有多种形态，包括球形、弧形、杆形及螺旋形，体积通常在0.5-2 μm之间^[27]。趋磁细菌从周围环境中获取铁元素，并在严格的基因调控下于胞内囊泡中形成35-120 nm的磁铁矿(Fe₃O₄)或硫铁矿(Fe₃S₄)晶体，组装成功能化的磁小体^[28]。磁小体晶体不仅化学纯度高、粒度均一并且形态多样，包括细长的棱柱形、立方体八面体和子弹形^[29]，呈单链或多链排列。磁小体不仅能够作为微化石记录古信息，还可以响应地球磁场进行地理定位和导航，帮助细菌定位最佳生存区域^[30]。

磁小体生物合成是一个高度调控的过程。当前研究主要集中于模式菌株Magnetospirillum magneticum AMB-1和格利菲斯瓦尔德镇磁螺菌(Magnetospirillum gryphiswaldense) MSR-1，其磁小体基因簇(magnetosome gene cluster, MGCs)及编码的磁小体蛋白在膜形成、晶体成核及链状排列等多个阶段发挥协同作用，且对宿主遗传背景具有依赖性^[31]。磁小体的合成过程可大致分为磁小体膜形成、铁运输与氧化还原调控、晶体成核与生长以及链状排列4个阶段^[32]。(1) 磁小体膜从细胞质膜内陷，形成由5-6 nm厚的蛋白质磷脂双层组成的囊泡结构。多种膜相关蛋白如MamB、MamM、MamQ和MamL等协同驱动磁小体膜的形成^[32]。近期冷冻电子断层与功能基因组学研究表明，磁小体膜的尺寸并非被动生成，而是受保守丝氨酸蛋白酶MamE的蛋白水解开关主动调控；MamE在上游因子MamO和MamM的调节下剪切底物MamD，以限制或放大膜的扩张^[33]，提示磁小体膜的形成依赖一个动态的膜蛋白复合体网络。(2) 在囊泡形成后，细胞需实现铁的有效获取与精确递送，为晶体成核提供原料与化学环境^[34]；然而，关于铁离子在磁小体膜内的价态与转化机制仍存在争议。仿生研究表明，铁离子以二价形式进入磁小体膜，部分Fe²⁺被氧化后生成水铁矿等前驱体，随后通过连续的氧化还原过程转化为Fe₃O₄晶体^[35]。(3) 晶体成核、生长阶段由多种蛋白精细调控。其中MamO蛋白在成核阶段发挥关键作用，一方面激活MamE介导的矿化蛋白的水解，另一方面依赖其特有的二组氨酸金属结合位点直接与过渡金属离子结合，从而诱导晶核的形成^[36]。Mms6作为典型的晶体调控蛋白，其酸性尾部可与亚铁离子结合，调控晶体生长^[37]，从而促进具有均一尺寸与形状的磁铁矿纳米晶体形成。(4) 这些磁小体晶体组装成链。其中，MamK蛋白是一种细胞骨架相关蛋白，属于细菌特有的类肌动蛋白(actin-like protein)，能在细胞内形成纤维结构，引导磁小体按线性排列方向组织^[38]。MamY蛋白作为一种膜锚定的机械性支架蛋白，负责将磁小体链准确定位于细胞几何主轴上，使细胞的运动方向与磁力方向高度一致^[39]。值得注意的是，磁小体形成不仅受限于膜结构与磁小体蛋白的精细调控，还受到细胞代谢的影响。例如，Pang等^[40]发现反硝化代谢生成的NO信号可通过转录因子NsrR激活mamAB操纵子的转录，从而正向调控mamAB操纵子的转录，促进磁小体形成。质子转移反应质谱实时分析也证实NO与N₂O产量显著上升，表明磁小体形成受到细胞氧化还原代谢的直接调控^[41]。这些发现为长期存在的争议提供了新的线索：磁小体的矿物类型与成核机制可能并非单纯由基因决定，而是由二者交织的代谢调控网络共同塑造。近期，Liu等^[42]通过整合基因组与表型数据，构建了一个适用于多种类MTB的基因网络调控模型，用以解析磁小体的形成及其链状排列机制，为理解磁小体的形成机制提供了新的系统的视角。

除了趋磁细菌，近年来也有研究发现多种细菌可能具备控制矿化的能力，拓宽了对细菌控制矿化多样性的认识。Benzerara等^[43]报道蓝细菌(Cyanobacteriota)可在胞内形成无定形碳酸钙(amorphous calcium carbonate, ACC)。与磁小体类似，ACC具有颗粒尺寸范围窄、被脂质单层膜包裹等特点，并且其形成也受到特定基因簇的调控。在蓝细菌的矿化过程中，编码膜转运蛋白和碳酸酐酶的相关基因会显著上调表达^[44-45]，提示这些基因在离子转运与碳酸根离子形成过程中发挥着重要作用。不同的是，ACC为非晶态，并含有Mg、Sr、Ba等杂质^[46]。此外，Li等^[47]近期在一株新发现的名为WYHC-5的趋磁细菌中发现其可形成细胞内非晶态的二氧化硅球粒，并推测该过程与一个潜在的沉硅基因silicatein-1有关；初步结果表明，WYHC-5可能通过形成二氧化硅颗粒来增加细胞质量、调节浮力，从而提升其在水体中上下穿梭时的稳定性，具备一定的生物学功能。

1.2 细菌诱导矿化

细菌诱导矿化是指细菌通过新陈代谢活动改变胞外微环境(如pH、氧化还原电位、离子浓度等)，间接引发矿物的非均相成核与沉淀，该矿化产物大多沉积于细胞外或表面^[48]。这一机制可追溯至Lowenstam^[13]提出的“生物诱导矿化”，最初用于描述矿物的形成由生物体的生命活动间接促成。与细菌控制矿化不同，矿物的形成并非由遗传控制主导，而是由于代谢导致局部环境变化，进而在特定条件下引发的物理化学过程^[17]。

近年来，大量研究证实能够诱导矿化的细菌种类众多，主要包括产脲酶菌(urease-producing bacteria, UPB)、硫酸盐还原菌(sulfate-reducing bacteria, SRB)、解磷菌(phosphate-solubilizing bacteria, PSB)、硝酸盐还原菌(nitrate-reducing bacteria, NRB)、铁氧化菌(Fe-oxidizing bacteria, FOB)和铁还原菌(Fe-reducing bacteria, FRB)等^[49]。产脲酶菌如巴氏芽孢八叠球菌(Sporosarcina pasteurii)通过分解尿素产生氨和碳酸根，使pH升高，进而诱导碳酸钙、金属离子与矿物沉淀^[50-52]；硫酸盐还原菌如脱硫弧菌(Desulfovibrio desulfuricans)在厌氧(或缺氧)条件可将SO₄^2-还原为S^2-，后者与金属离子结合生成金属硫化物沉淀^[53]；解磷菌如双向伯克霍尔德氏菌(Burkholderia ambifaria) QY14则通过增加酸性磷酸酶活性，与土壤环境中的钙离子结合，形成含羟基磷灰石的镉(Cd-HAP)^[54]；硝酸盐还原菌如产气克雷伯氏菌(Klebsiella aerogenes)利用有机碳源还原NO₃⁻生成NO₂⁻，NO₂⁻促进Fe²⁺氧化为Fe³⁺，进而诱导Fe²⁺和Fe³⁺共沉淀形成针铁矿^[55]；铁氧化细菌/还原菌可通过调节Fe²⁺/Fe³⁺的氧化还原状态形成铁氧化物或铁还原沉淀^[56-57]。现将细菌诱导矿化的机制与矿化产物进行总结，见表1。

产脲酶菌因其诱导机制明确、矿化效率高、环境适应性强，成为细菌诱导矿化的典型代表^[58]。产脲酶菌通过尿素水解诱导矿化，在脲酶催化下将尿素分解为CO₂和NH₃，这些化合物与水反应形成HCO₃⁻、NH₄⁺和OH⁻。OH⁻的产生促进了HCO₃⁻向CO₃^2-的转化，当环境中存在足够的Ca²⁺时可形成碳酸钙沉淀(图2)^[59]。在自然条件下，产脲酶菌诱导的矿化过程不仅依赖脲酶催化尿素水解引起的pH变化和碳酸根积累，同时Ca²⁺浓度、温度以及细菌表面性质等因素还共同决定矿化产物的类型与形貌^[60]。该过程生成的碳酸钙晶体具有多种形貌与结晶形式，包括方解石、文石和球霰石，甚至可形成非晶态碳酸钙^[61]。此外，产脲酶菌诱导的矿化在人体健康领域也具有重要意义。一方面，产脲酶细菌[如奇异变形菌(Proteus mirabilis)和摩氏摩根氏菌(Morganella morganii)等]是尿路感染性结石形成的重要病原菌^[62]，其通过尿素分解生成鸟粪石(NH₄MgPO₄·6H₂O)、碳酸磷灰石和尿酸铵等矿物^[9]。奇异变形杆菌的脲酶活性与其菌毛结构共同促进尿路导管上的结晶沉积^[63]；Schaffer等^[64]的研究表明奇异变形杆菌还可能启动膀胱结石的形成。另一方面，口腔环境中也存在编码脲酶的细菌^[65][如血链球菌(Streptococcus sanguinis)和内氏放线菌(Actinomyces naeslundii)]，它们可能通过尿素分解提高局部pH值，影响牙结石的形成^[66]。然而，关于口腔脲酶菌的作用仍存在争议，例如有研究提示唾液pH值与牙结石形成之间关系不显著^[67]。

1.3 细菌影响矿化

细菌影响矿化是指细菌的表面结构作为成核位点影响矿物沉淀的过程。在该途径中矿物沉淀本身是一个非生物过程，但受到细菌结构的物理化学属性的调控^[15]。生物诱导矿化可分为被动(passive)与主动(active) 2种；其中，被动生物诱导矿化是指阳离子与细胞表面进行非专一性结合，之后阳离子吸引溶液中的阴离子成核形成矿物，这一过程不依赖微生物代谢^[48]。Dupraz等^[15]于2009年对“被动生物诱导矿化”理论进行延伸和独立分类，提出“生物影响矿化”概念，即矿物从过饱和溶液沉淀到生物结构上；与细菌诱导矿化相比，细菌影响矿化并不强调细菌的代谢活动对矿化环境的改变，而更关注细菌残留结构与矿物界面的相互作用。即使在细胞死亡之后，其表面结构仍可能持续影响矿化过程^[68]。

随着对微生物表面结构研究的深入，细菌表面的细胞外聚合物(extracellular polymeric substances, EPS)、细胞壁(cell wall)以及S层蛋白(surface-layer protein, SLP)等可作为矿物成核模板，在无细胞代谢活动时也可介导局部矿物的沉积(图3)^[16,69]。

1.3.1 细胞外聚合物

细菌的细胞外聚合物作为生物膜的主要成分，由多糖、蛋白质、脂质和细胞外DNA等高分子聚合物组成^[70]。在细菌介导的影响矿化过程中，EPS主要通过调控局部化学环境、提供成核模板等方式促进矿物的沉积。其酸性官能团(如羧基、磷酸基)可结合钙、镁等阳离子降低成核能垒，通过静电吸附、形成氢键或络合作用等与金属离子结合，形成初始成核位点从而促进晶体的生长(图3A)^[71-72]。

EPS自身的结构特征对其矿化调控能力具有显著影响。研究表明可溶性EPS富含游离羧基，易诱导球霰石等亚稳相的成核；而紧密结合型EPS则因其较强的空间约束和稳定性，倾向于维持方解石晶体的稳定生长^[73]。此外，EPS的化学组成也影响矿化效果。例如，Ivanova等^[74]发现在蜡样芽孢杆菌(Bacillus cereus)浮游生物培养中产生的胞外多糖和带负电荷的细胞外DNA能够促进局部Ca²⁺过饱和。EPS中的蛋白质可能倾向于抑制方解石的形成，而多糖可能倾向于促进方解石的形成^[75]。除了结构与组成本身，EPS的表达量也对矿化效率和产物质量产生显著影响。EPS表达水平较高的细菌通常形成更为致密、连续的矿化层，有利于提高整体矿化效率并增强沉积物的稳定性^[76]。不同多糖成分由于电荷密度和分子结构不同，矿物成核所需的能垒也有所差异^[77]。

1.3.2 细胞壁

细菌细胞壁的主要结构成分为肽聚糖(peptidoglycan, PGN)，由N-乙酰葡糖胺(N-acetylglucosamine, NAG)和N-乙酰胞壁酸(N-acetylmuramic acid, NAM)交联^[78]。革兰氏阳性和革兰氏阴性菌的细胞壁结构存在显著差异：前者具有厚的肽聚糖层，并含有丰富的磷壁酸；后者则具有较薄的肽聚糖层，其外膜含有脂多糖(lipopolysaccharide, LPS)，外膜与肽聚糖通过短肽链交替连接形成骨架^[79]。与EPS类似，细胞壁表面也广泛分布着羧基、磷酸基、胺基和羟基等带电官能团^[80]，这些基团能与钙、镁等阳离子发生螯合，为矿物形成提供成核位点(图3B)。

细胞壁的结构类型和官能团组成进一步影响矿物的类型与晶体形貌。例如，研究表明革兰氏阳性菌枯草芽孢杆菌(Bacillus subtilis)细胞壁可诱导方解石晶体的定向生长，而革兰氏阴性的大肠埃希氏菌(Escherichia coli)则更容易形成无定形碳酸钙^[81]。在不同的饱和状态下，细胞壁对矿物沉淀模式的调控也有所不同^[82]。此外，细胞壁不同的酸性基团对金属离子的选择性吸附也是决定矿物类型的重要因素^[83]。

1.3.3 S层蛋白

在细菌和古菌中S层蛋白是最常见的表面结构之一，由蛋白质亚基组成，形成高度有序的二维晶体阵列^[84]。S层蛋白通常附着于肽聚糖层、外膜或质膜上^[85-86]。序列分析表明，大多数S层蛋白富含酸性和疏水性氨基酸，其等电点多位于弱酸性pH区间^[87]，有利于在中性或偏碱环境中吸附阳离子，为矿物成核提供条件(图3C)。

虽然S层蛋白的传统功能多与细胞形态维持和分子筛过滤相关^[88]，但也有研究表明S层蛋白在细菌表面的矿化过程中发挥重要作用。例如，S层蛋白的高度有序排列和纳米尺度孔道为异相成核提供了规则、离散的模板，有助于无机矿物的空间定位和结构导向^[89]。Kish等^[90]在古菌嗜酸热硫化叶菌(Sulfolobus acidocaldarius)的S层表面观察到了羟基氧化铁与无定形磷酸铁的沉积；此外，S层的物理结构与化学组分还可能协同调控矿物的形貌、结晶取向及晶相演变，提示这可能是微生物在极端矿化环境中维持细胞完整性与功能的关键策略之一^[91]。然而，S层蛋白对于矿物形成影响的研究多停留在形貌观察层面，缺乏分子尺度的机制解析，仍需进一步探索。

2 细菌介导的矿化在生物医学领域的应用

收起

细菌介导的矿化作为重要的自然过程，揭示了细菌在无机沉积过程中所具备的调控能力。控制矿化、诱导矿化和影响矿化是细菌的3种典型矿化机制，依次展现了细菌在细胞活动、代谢与界面水平上对矿物形成的多重调控，并在细胞内外形成结构有序、成分多样的矿化产物。这些天然生成的生物矿物通常具有优异的性能、良好的界面相容性以及可调控的功能特性^[12]，为其在生物医学领域的应用奠定了重要基础。

近年来，随着纳米医学与合成生物学的发展，研究者从细菌矿化机制中汲取启发，通过解析成核调控、界面组装和晶体生长等关键过程，将自然矿化过程转化为可设计的仿生或工程化体系。这种从“自然矿化”向“工程矿化”的转变，使细菌不再仅是矿物沉积的参与者，而成为可用于构建成像、治疗及组织修复材料的生物功能平台。基于此，本文将系统综述细菌介导的矿化在生物医学领域的研究进展，重点探讨其在医学成像、靶向治疗与组织工程等方向的应用与前景(图4)^[92]。

2.1 医学成像

磁共振成像(magnetic resonance imaging, MRI)是一种常用的成像方式，因其优异的空间分辨率和软组织对比度，成为临床疾病诊断重要的工具^[93]。质子密度与弛豫时间是影响MRI中图像对比度的因素^[94]。氧化铁纳米颗粒可作为MRI造影剂，增强正常组织和病变组织之间的图像对比度，而化学合成氧化铁纳米颗粒的弛豫效率常受到粒径分布、表面包覆、聚集稳定性等因素制约，影响成像灵敏度与重复性^[95]。磁小体是生物来源的磁性纳米颗粒，由于其尺寸较大、分散性较高，且单畴磁性颗粒(single-domain magnetic particle)能产生稳定磁矩与强局域磁场梯度^[28]，为MRI提供了比目前磁性氧化铁纳米颗粒制剂更好的对比度^[96]。例如，Mériaux等^[97]研究发现，纯化后磁小体的单畴磁性颗粒在超高场MRI中表现出的横向弛豫率高出市售氧化铁颗粒的4倍，即与相同剂量的人工氧化铁相比，静脉注射低剂量磁小体可使小鼠脑血管T2加权信号衰减强度提升，进而实现高灵敏度的脑血管成像效果。进一步利用磁小体膜蛋白构建仿生纳米反应系统，并对磁小体进行表面功能化，可实现精准的靶向成像。Ma等^[98]通过将Mms6蛋白集成到反胶束系统中，构建了仿生纳米反应器并合成了单域磁性纳米颗粒，该颗粒具有与天然磁小体相似但尺寸更小的特性，表现出强大的磁性靶向性和增强的肿瘤渗透性。与传统人血清白蛋白(human serum albumin, HSA)包被的氧化铁纳米颗粒相比，抗表皮生长因子受体(epidermal growth factor receptor, EGFR)修饰后的磁小体在T2加权的MRI中可显著增强对肿瘤组织的对比度，表现为肿瘤区域信号减弱(图像更暗)^[96]，从而提升了靶向肿瘤成像的清晰度与特异性。

此外，基于细菌影响矿化策略是构建多功能成像平台的另一有效途径。例如，Chen等^[99]通过在活菌表面依次沉积MnO₂与Fe₂O₃纳米颗粒，并在最外层包覆CaCO₃，制得兼具可视化与治疗功能的“生物矿化外壳”；该结构在肠道递送过程中可实现T1和T2双模式MRI实时追踪，后续Mn²⁺释放进一步实现T1成像识别。

2.2 靶向治疗

磁热疗是一种利用磁性材料在交变磁场中产热，实现肿瘤组织局部高温消融的治疗方式。该技术通过将氧化铁纳米颗粒注射到肿瘤部位并施加交变磁场诱导其局部加热，从而选择性地杀伤癌细胞^[100]。与人工合成的磁性纳米颗粒相比，磁小体链状有序排列于细菌细胞内，增强了磁各向异性，从而提高了磁热转换效率^[101]。这使得磁小体在磁热疗中的能量利用率和加热性能均优于无序或分散的磁性颗粒。Chen等^[102]表明在神经母细胞瘤裸鼠模型中，使用完整AMB-1处理组的细胞相较于使用单个磁小体或磁小体链的组展现出更明显的抗肿瘤效果。除了高效的磁热响应性能，MTB还具有独特的运动与导航能力，其鞭毛赋予细菌自主推进能力，而磁小体链则使MTB在外部磁场作用下实现定向运动^[103]。例如，MSR-1菌株与负载抗生素环丙沙星的介孔二氧化硅微管整合，在磁引导和细菌自身推进力的共同作用下，不仅将抗生素载荷有效递送至深层感染部位，还可高效破坏生物膜^[104]。

近年来，基于细菌诱导与影响矿化的靶向治疗策略日趋多样化。其中，诱导矿化依托细菌的还原代谢过程，可在细胞表面原位生成金属纳米结构。Chen等^[105]报道，奥奈达湖希瓦氏菌(Shewanella oneidensis) MR-1可还原钯盐在细胞表面形成钯单质，从而构建具有优异光热性能与肿瘤靶向性的“光热细菌”体系，用于增强光热治疗效果。相较之下，影响矿化策略更注重通过表面修饰或包被实现细菌功能化。例如，碳酸钙生物矿化的肠沙门氏菌(Salmonella enterica)可作为原位癌的疫苗载体，调节肿瘤免疫微环境并增强系统性免疫应答^[106]。Geng等^[107]则利用外源聚乙烯吡咯烷酮(polyvinylpyrrolidone, PVP)修饰细胞壁羟基基团以增强成核能力，在益生菌表面形成均匀的碳酸钙涂层，从而在炎症性肠病模型中显著改善治疗效果。此外，Zhang等^[108]通过构建单克隆抗体-多唾液酸偶联物诱导耐甲氧西林金黄色葡萄球菌(methicillin-resistant Staphylococcus aureus, MRSA)表面钙化，在系统感染治疗与免疫调控中展现出潜在治疗价值。被二氧化锰包被的矿化肠沙门氏菌可同时激活环鸟苷酸-单磷酸腺苷合成酶家族(cyclic guanosine monophosphate-adenosine monophosphate synthase, cGAS)-干扰素刺激基因(stimulator of interferon gene, STING)、NOD样受体(NOD-like receptor, NLR)及Toll样受体(Toll-like receptor, TLR)通路，并作为局部给药的免疫刺激剂在肿瘤免疫治疗中展现出强效的免疫原性^[109]。

2.3 组织工程

与化学合成的磁性纳米颗粒相比，磁小体表面富含天然脂质与蛋白质基团，为后续的化学或基因修饰提供理想界面。通过基因工程手段对磁小体膜蛋白进行适当的基因修饰，可用于连接特定肽段或蛋白质配体，进一步增强MTB系统的靶向能力与治疗功能。其中，MamC蛋白因高表达量、稳定的膜定位以及对磁小体结构的关键作用，被广泛选作基因工程改造载体，赋予磁小体特异性靶向能力^[92]。例如，Plan Sangnier等^[110]通过融合精氨酸(arginine, R)-甘氨酸(glycine, G)-天冬氨酸(aspartic acid, D)三肽(RGD)与磁小体蛋白MamC进行基因工程改造，在趋磁细菌中原位合成了表面携带RGD的磁小体(magnetosomes@RGD)，实现了对前列腺癌细胞的高效靶向和摄取。Wang等^[111]将阿霉素(doxorubicin, DOX)与转铁蛋白(transferrin, Tf)共载于磁小体(Tf-BMs-DOX)，利用Tf与肝癌细胞表面转铁蛋白受体的特异性结合，实现肝细胞癌的主动靶向。此外，磁小体表面功能化还可以用于病原体的检测。将磁小体蛋白如MamC或MamF与金黄色葡萄球菌免疫球蛋白结合蛋白(staphylococcal protein A, SPA)进行融合表达，所得的重组磁小体能够特异性结合哺乳动物抗体的可结晶区段(fragment crystallizable, Fc)，可有效捕获海产品常见病原菌副溶血弧菌(Vibrio parahaemolyticus)^[112]。

细菌诱导矿化与细菌影响矿化策略也被用于构建新型生物功能体系。例如，工程化E. coli可自分泌蛋白质纤维组装成“活性杂交胶囊”，具备生物分子螯合与酶催化功能，并通过脲酶介导的生物矿化动态调节矿化层的硬度与体积，从而在外部刺激下实现自适应机械调控^[113]。Fan等^[114]则利用生物矿化反应在E. coli MG1655表面原位负载金纳米颗粒，并构建表达肿瘤坏死因子-α (tumor necrosis factor alpha, TNF-α)的质粒系统，使细菌具备光热响应与治疗双功能。经口服后，该系统可在体内循环中靶向肿瘤区域，通过光照诱导TNF-α表达并触发肿瘤细胞凋亡。

3 总结与展望

收起

细菌介导的矿化作为一种在自然界中广泛存在的过程，近年来在矿化机制解析和生物医学应用领域均取得了显著进展。从机制层面来看，研究表明细菌的矿化行为可分为细胞内受控矿化、代谢诱导沉淀以及细胞表面介导成核等，常被归为细菌控制矿化、诱导矿化与影响矿化3种机制。然而，随着研究的深入，越来越多的证据表明细菌矿化过程可能并非完全独立的3种机制，而是存在一定的“过渡状态”或“混合机制”。例如，球形赖氨酸芽孢杆菌(Lysinibacillus sphaericus) JG-7B和鞘氨醇单胞菌(Sphingomonas sp.) S15-S1在酸性条件下可通过自身产生的酸性磷酸酶释放胞内储存的磷，在胞内或表面形成铀磷酸盐沉积，尽管未观察到典型的囊泡结构或验证矿物的生物功能性，但这一过程无需外源底物，且依赖胞内酶促反应，提示其矿化过程既具备BCM的调控能力，又具有BIM的诱导属性^[5,115]。未来的研究有必要结合细菌矿化过程，从晶体结构、形成路径和调控精度等方面进一步审视细菌矿化过程中的机制。

细菌控制矿化被认为是高度调控的过程，涉及膜结构、囊泡形成、离子通道、成核调控蛋白等多个环节^[42]。目前，对该过程的研究多集中于趋磁细菌这类模式细菌，对其他细菌物种控制矿化的过程了解有限。未来可结合多组学技术(如转录组、蛋白组与代谢组)及实时活细胞成像技术，系统解析矿化过程中调控因子的时空表达动态，以深入揭示分子机制与调控网络。在细菌诱导矿化方面，细菌在诱导过程中的行为动态仍需进一步研究，尤其是在不同环境条件下细菌的迁移性、趋化性及其与成核过程的协同关系仍缺乏系统研究^[59]。未来可借助原位成像与分子标记技术，在孔隙尺度上实现细菌运动轨迹与矿物沉积过程的同步追踪，揭示其在自然环境或应用场景中的动态行为机制。对于细菌影响矿化，尽管已有研究证实细胞分泌的胞外聚合物在成核与晶体生长中发挥关键作用，但具体哪些有机成分在其中起主导作用尚不明确。此外，不同细菌物种是否具有特异性的矿化潜能，目前也缺乏系统验证。这些问题的存在限制了对BIFM机制的深入理解。为此，未来研究可重点从以下方向展开：(1) 利用高分辨率表征技术解析细菌表面官能团与矿物间的分子互作机制；(2) 建立细菌物种、EPS组成与矿物产物之间的对应关系，进一步阐明矿化潜能与物种特异性的关系。

在应用层面，虽然细菌控制矿化的产物(如磁小体)在肿瘤治疗、靶向递送、组织工程等领域展现出良好的应用前景，但其实际转化仍面临诸多挑战，例如趋磁细菌的大规模培养难度高、磁小体产量低、生物相容性与免疫反应尚未完全明确等问题^[116]。未来研究可构建更贴近人体环境的体内验证模型，从生物安全性、功能可控性等方面开展系统评估，以推动其实际应用。目前，细菌介导的矿化在生物医学领域的应用研究主要基于细菌控制矿化机制及其矿化产物(如磁小体)，与该机制受到严格的生物调控，同时能够产生纯度高、粒径均一的磁性纳米颗粒相关。诱导矿化可通过调节细胞代谢与局部微环境实现矿化的间接调控，而影响矿化则依赖细菌表面改性，为生物涂层、药物递送及免疫调节等提供可能。尽管如此，细菌介导矿化在生物医学领域的应用仍面临多重挑战。活菌体系的安全性与免疫相容性尚未得到充分验证；同时，影响与诱导矿化过程受外界条件影响显著，难以实现时空可控的矿化行为^[117]。未来研究需通过合成生物学、材料科学与工程技术的协同策略，实现从自然矿化到可编程、可控制的功能化矿化转变。例如，通过工程菌株设计、矿化通路调控及微环境精准建模，在确保安全性的前提下实现产物的一致性与功能可调性。通过机制研究与应用探索的协同推进，有望更深入地理解不同细菌矿化机制，并推动新材料和新技术的仿生研发。

基金

收起

国家自然科学基金重大项目(81991501)
四川省自然科学基金(2024NSFSC0546)

参考文献

收起

文献

收起

参考文献引证文献

排序方式：

[1]

WANG

, HU

, WANG

, TANG

. Expanding from materials to biology inspired by biomineralization[J]. Interdisciplinary Materials, 2024, 3(2): 165-188.

[2]

, VENEKLAAS

, KUO

, LAMBERS

. Physiological and ecological significance of biomineralization in plants[J]. Trends in Plant Science, 2014, 19(3): 166-174.

[3]

MURSHED

. Mechanism of bone mineralization[J]. Cold Spring Harbor Perspectives in Medicine, 2018, 8(12): a031229.

[4]

QIN

, WANG

, MA

, SHEN

, LI

, JIAO

, TAY FR, NIU

. Microbe-mediated extracellular and intracellular mineralization: environmental, industrial, and biotechnological applications[J]. Advanced Materials, 2020, 32(22): 1907833.

[5]

DONG

, HUANG

, ZHAO

, ZENG

, LIU

, SHENG

, SHI

, WU

, JIANG

, LI

, ZHANG

, GUO

, LI

, HOU

, CHEN

. A critical review of mineral-microbe interaction and co-evolution: mechanisms and applications[J]. National Science Review, 2022, 9(10): nwac128.

[6]

DOUGLAS

, BEVERIDGE

. Mineral formation by bacteria in natural microbial communities[J]. FEMS Microbiology Ecology, 1998, 26(2): 79-88.

[7]

SPADAFORA

, PERRI

, MCKENZIE

, VASCONCELOS

. Microbial biomineralization processes forming modern Ca: Mg carbonate stromatolites[J]. Sedimentology, 2010, 57(1): 27-40.

[8]

GARATE

, SUREDA

, AGELL

, URIZ

. Endosymbiotic calcifying bacteria across sponge species and oceans[J]. Scientific Reports, 2017, 7: 43674.

[9]

麦海星, 乔庐东, 吴文起, 胡卫国, 刘凯隆, 齐士勇, 陈雪花, 何朝辉, 沈诞. 上尿路感染性结石诊断与治疗中国专家共识(2024版)[J]. 微创泌尿外科杂志, 2024, 13(4): 217-223.

MAI HX, QIAO

, WU

, HU

, LIU

, QI

, CHEN

, HE

, SHEN

. Chinese expert consensus on diagnosis and treatment of upper urinary tract infection stones (2024 edition)[J]. Journal of Minimally Invasive Urology, 2024, 13(4): 217-223 (in Chinese).

[10]

TAN

, GILLAM

, MORDAN

, GALGUT

. A preliminary investigation into the ultrastructure of dental Calculus and associated bacteria[J]. Journal of Clinical Periodontology, 2004, 31(5): 364-369.

[11]

, ZHOU

, SU

, LI

. Corynebacterium matruchotii: a confirmed calcifying bacterium with a potentially important role in the supragingival plaque[J]. Frontiers in Microbiology, 2022, 13: 940643.

[12]

ANSARI

, AHMAD

, TANWEER

, AZAM

. Magnetic iron oxide nanoparticles as a tool for the advancement of biomedical and environmental application: a review[J]. Biomedical Materials & Devices, 2024, 2(1): 139-157.

[13]

LOWENSTAM

. Minerals formed by organisms[J]. Science, 1981, 211(4487): 1126-1131.

[14]

MANN

. Biomineralization: Principles and Concepts in Bioinorganic Materials Chemistry[M]. Oxford: Oxford University Press, 2001.

[15]

DUPRAZ

, REID

, BRAISSANT

, DECHO

, NORMAN

, VISSCHER

. Processes of carbonate precipitation in modern microbial mats[J]. Earth-Science Reviews, 2009, 96(3): 141-162.

[16]

GÖRGEN

, BENZERARA

, SKOURI-PANET

, GUGGER

, CHAUVAT

, CASSIER-CHAUVAT

. The diversity of molecular mechanisms of carbonate biomineralization by bacteria[J]. Discover Materials, 2020, 1(1): 2.

[17]

BENZERARA

, MIOT

, MORIN

, ONA-NGUEMA

, SKOURI-PANET

, FÉRARD

. Significance, mechanisms and environmental implications of microbial biomineralization[J]. Comptes Rendus Geoscience, 2011, 343(2/3): 160-167.

[18]

BAZYLINSKI

. Biologically controlled mineralization in prokaryotes[J]. Reviews in Mineralogy and Geochemistry, 2003, 54(1): 217-247.

[19]

LOWENSTAM

, WEINER

. On Biomineralization[M]. Oxford: Oxford University Press, 1989.

[20]

WEINER

. An overview of biomineralization processes and the problem of the vital effect[J]. Reviews in Mineralogy and Geochemistry, 2003, 54(1): 1-29.

[21]

YAN

, ZHANG

, MA

, ZHONG

, TANG

, MAI S. The mechanism of biomineralization: progress in mineralization from intracellular generation to extracellular deposition[J]. Japanese Dental Science Review, 2023, 59: 181-190.

[22]

WEINER

, ADDADI

. Crystallization pathways in biomineralization[J]. Annual Review of Materials Research, 2011, 41: 21-40.

[23]

GÖRLICH

, PAWOLSKI

, ZLOTNIKOV

, KRÖGER

. Control of biosilica morphology and mechanical performance by the conserved diatom gene Silicanin-1[J]. Communications Biology, 2019, 2: 245.

[24]

SIMKISS

, WILBUR

. Biomineralization[M]. San Diego: Academic Press, 1989.

[25]

COSMIDIS

, BENZERARA

. Why do microbes make minerals[J]. Comptes Rendus Géoscience, 2022, 354(G1): 1-39.

[26]

BLAKEMORE

. Magnetotactic bacteria[J]. Science, 1975, 190(4212): 377-379.

[27]

JACOB

, SUTHINDHIRAN

. Magnetotactic bacteria and magnetosomes: scope and challenges[J]. Materials Science and Engineering: C, 2016, 68: 919-928.

[28]

VARGAS

, CYPRIANO

, CORREA

, LEÃO

, BAZYLINSKI

, ABREU

. Applications of magnetotactic bacteria, magnetosomes and magnetosome crystals in biotechnology and nanotechnology: mini-review[J]. Molecules, 2018, 23(10): 2438.

[29]

LEFÈVRE

, BAZYLINSKI

. Ecology, diversity, and evolution of magnetotactic bacteria[J]. Microbiology and Molecular Biology Reviews, 2013, 77(3): 497-526.

[30]

BELLINGER

, WEI

, HARTMANN

, CADIOU

, WINKLHOFER

, BANKS

. Conservation of magnetite biomineralization genes in all domains of life and implications for magnetic sensing[J]. Proceedings of the National Academy of Sciences of the United States of America, 2022, 119(3): e2108655119.

[31]

WAN

, JI

, LIU

, MA

, PAN

, LIN

. Biomineralization in magnetotactic bacteria: from diversity to molecular discovery-based applications[J]. Cell Reports, 2024, 43(12): 114995.

[32]

UEBE

, SCHÜLER

. Magnetosome biogenesis in magnetotactic bacteria[J]. Nature Reviews Microbiology, 2016, 14(10): 621-637.

[33]

WAN

, BROWNE

, HERSHEY

, MONTABANA

, IAVARONE

, DOWNING

, KOMEILI

. A protease-mediated switch regulates the growth of magnetosome organelles in Magnetospirillum magneticum [J]. Proceedings of the National Academy of Sciences of the United States of America, 2022, 119(6): e2111745119.

[34]

钟林, 李晓晨, 符军, 张友明, 李爱英. 磁小体的生物合成及用于肿瘤靶向治疗的研究进展[J]. 中国科学: 生命科学, 2017, 47(5): 572-585.

ZHONG

, LI

, FU

, ZHANG

, LI

. Biosynthesis and application of magnetosomes in tumor-targeting therapy[J]. Scientia Sinica (Vitae), 2017, 47(5): 572-585 (in Chinese).

[35]

MAO

, LIU

, SUN

, ZHAO

, AN

, WU

, FENG

, CHEN

, ZHANG

, LI

, HUANG

, GU

. Elucidating the bioinspired synthesis process of magnetosomes-like Fe₃O₄ nanoparticles[J]. Small, 2024, 20(21): 2308247.

[36]

HERSHEY

, REN

, MELNYK

, BROWNE

, OZYAMAK

, JONES

, CHANG

MCY

, HURLEY

, KOMEILI

. MamO is a repurposed serine protease that promotes magnetite biomineralization through direct transition metal binding in magnetotactic bacteria[J]. PLoS Biology, 2016, 14(3): e1002402.

[37]

RAWLINGS

, BRAMBLE

, HOUNSLOW

, WILLIAMSON

, MONNINGTON

, COOKE

, STANILAND

. Ferrous iron binding key to Mms6 magnetite biomineralisation: a mechanistic study to understand magnetite formation using pH titration and NMR spectroscopy[J]. Chemistry-A European Journal, 2016, 22(23): 7885-7894.

[38]

KOMEILI

, LI

, NEWMAN

, JENSEN

. Magnetosomes are cell membrane invaginations organized by the actin-like protein MamK[J]. Science, 2006, 311(5758): 242-245.

[39]

TORO-NAHUELPAN

, GIACOMELLI

, RASCHDORF

, BORG

, PLITZKO

, BRAMKAMP

, SCHÜLER

, MÜLLER

. MamY is a membrane-bound protein that aligns magnetosomes and the motility axis of helical magnetotactic bacteria[J]. Nature Microbiology, 2019, 4(11): 1978-1989.

[40]

PANG

, ZHENG

, MA

, TIAN

, WEN

. Nitric oxide sensor NsrR is the key direct regulator of magnetosome formation and nitrogen metabolism in Magnetospirillum [J]. Nucleic Acids Research, 2024, 52(6): 2924-2941.

[41]

ZHANG

, WAN

, SHEN

, ZHANG

, WANG

, WAN

, ZHANG

, MA

, LIN

, WANG

, PAN

. Influence of a 7 T magnetic field on growth, biomineralization, and denitrification metabolism in Magnetospirillum gryphiswaldense MSR-1[J]. Applied and Environmental Microbiology, 2025, 91(10): e0106925.

[42]

LIU

, ZHENG

, ZHANG

, BAI

, ZHU

, BENZERARA

, MENGUY

, ZHAO

, ROBERTS

, PAN

, LI

. Key gene networks that control magnetosome biomineralization in magnetotactic bacteria[J]. National Science Review, 2023, 10(1): nwac238.

[43]

BENZERARA

, SKOURI-PANET

, LI

, FÉRARD

, GUGGER

, LAURENT

, COURADEAU

, RAGON

, COSMIDIS

, MENGUY

, MARGARET-OLIVER

, TAVERA

, LÓPEZ-GARCÍA

, MOREIRA

. Intracellular Ca-carbonate biomineralization is widespread in cyanobacteria[J]. Proceedings of the National Academy of Sciences of the United States of America, 2014, 111(30): 10933-10938.

[44]

de WEVER

, BENZERARA

, COUTAUD

, CAUMES

, POINSOT

, SKOURI-PANET

, LAURENT

, DUPRAT

, GUGGER

. Evidence of high Ca uptake by cyanobacteria forming intracellular CaCO₃ and impact on their growth[J]. Geobiology, 2019, 17(6): 676-690.

[45]

BENZERARA

, DUPRAT

, BITARD-FEILDEL

, CAUMES

, CASSIER-CHAUVAT

, CHAUVAT

, DEZI

, DIOP

, GASCHIGNARD

, GÖRGEN

, GUGGER

, LÓPEZ-GARCÍA

, MILLET

, SKOURI-PANET

, MOREIRA

, CALLEBAUT

. A new gene family diagnostic for intracellular biomineralization of amorphous Ca carbonates by cyanobacteria[J]. Genome Biology and Evolution, 2022, 14(3): evac026.

[46]

COURADEAU

, BENZERARA

, GÉRARD

, MOREIRA

, BERNARD

, BROWN GE Jr, LÓPEZ-GARCÍA

. An early-branching microbialite Cyanobacterium forms intracellular carbonates[J]. Science, 2012, 336(6080): 459-462.

[47]

, LIU

, MENGUY

, ZHANG

, WANG

, BENZERARA

, FENG

, SUN

, ZHENG

, MENG

, GU

, LEROY

, HAO

, CHU

, PAN

. Intracellular silicification by early-branching magnetotactic bacteria[J]. Science Advances, 2022, 8(19): eabn6045.

[48]

FRANKEL

, BAZYLINSKI

. Biologically induced mineralization by bacteria[J]. Reviews in Mineralogy and Geochemistry, 2003, 54(1): 95-114.

[49]

HAN

, LI

, XUE

, CHEN

, ZHOU

, POON

. Bacterial-induced mineralization (BIM) for soil solidification and heavy metal stabilization: a critical review[J]. Science of the total Environment, 2020, 746: 140967.

[50]

VASKEVICIUS

, MALUNAVICIUS

, JANKUNEC

, LASTAUSKIENE

, TALAIKIS

, MIKOLIUNAITE

, MANEIKIS

, GUDIUKAITE

. Insights in MICP dynamics in urease-positive Staphylococcus sp. H6 and Sporosarcina pasteurii bacterium[J]. Environmental Research, 2023, 234: 116588.

[51]

BHATTACHARYA

, NAIK

, KHARE

. Harnessing the bio-mineralization ability of urease producing Serratia marcescens and Enterobacter cloacae EMB19 for remediation of heavy metal cadmium (II)[J]. Journal of Environmental Management, 2018, 215: 143-152.

[52]

MARÍN

, CABESTRERO

, DEMERGASSO

, OLIVARES

, ZETOLA

, VERA

. An indigenous bacterium with enhanced performance of microbially-induced Ca-carbonate biomineralization under extreme alkaline conditions for concrete and soil-improvement industries[J]. Acta Biomaterialia, 2021, 120: 304-317.

[53]

CHEN

, MIN

, WU

, ZHANG

, SI

. Biomineralization of Cd²⁺ and Pb²⁺ by sulfate-reducing bacteria Desulfovibrio desulfuricans and Desulfobulbus propionicus [J]. Frontiers in Environmental Science, 2025, 13: 1591564.

[54]

ZENG

, QIAO

, WANG

, SHENG

, WEI

, CHEN

, XU

. Immobilization of cadmium by Burkholderia sp. QY14 through modified microbially induced phosphate precipitation[J]. Journal of Hazardous Materials, 2021, 412: 125156.

[55]

ETIQUE

, JORAND

FPA

, ZEGEYE

, GRÉGOIRE

, DESPAS

, RUBY

. Abiotic process for Fe(II) oxidation and green rust mineralization driven by a heterotrophic nitrate reducing bacteria (Klebsiella mobilis)[J]. Environmental Science & Technology, 2014, 48(7): 3742-3751.

[56]

LUO

, JIANG

, XUE

, TANG

, ZHOU

, WU

, QIAN

, WU

. Arsenic biomineralization by iron oxidizing strain (Ochrobactrum sp.) isolated from a paddy soil in Hunan, China[J]. Land Degradation & Development, 2021, 32(6): 2082-2093.

[57]

HAN

, WANG

, ZHENG

, QIU

, LIU

, WANG

, MENGUY

, LEROY

, BOURGON

, KAPPLER

, LIU

, PAN

, LI

. Morphological, microstructural, and in situ chemical characteristics of siderite produced by iron-reducing bacteria[J]. Environmental Science & Technology, 2024, 58(25): 11016-11026.

[58]

, LI

. Biomineralization induced by cells of Sporosarcina pasteurii: mechanisms, applications and challenges[J]. Microorganisms, 2021, 9(11): 2396.

[59]

刘汉龙, 赵常, 肖杨. 微生物矿化反应原理、沉积与破坏机制及理论: 研究进展与挑战[J]. 岩土工程学报, 2024, 46(7): 1347-1358.

LIU

, ZHAO

, XIAO

. Reaction principles, deposition and failure mechanisms and theories of biomineralization: progress and challenges[J]. Chinese Journal of Geotechnical Engineering, 2024, 46(7): 1347-1358 (in Chinese).

[60]

LIU

, ALI A, SU

, LI

, HU

, WANG

. Microbial-induced calcium carbonate precipitation: influencing factors, nucleation pathways, and application in waste water remediation[J]. Science of the Total Environment, 2023, 860: 160439.

[61]

ZOU

, HABRAKEN

WJEM

, MATVEEVA

, JENSEN

ACS

, BERTINETTI

, HOOD

, SUN

, GILBERT

PUPA

, POLISHCHUK

, POKROY

, MAHAMID

, POLITI

, WEINER

, WERNER

, BETTE

, DINNEBIER

, KOLB

, ZOLOTOYABKO

, FRATZL

. A hydrated crystalline calcium carbonate phase: calcium carbonate hemihydrate[J]. Science, 2019, 363(6425): 396-400.

[62]

NORSWORTHY

, PEARSON

. From catheter to kidney stone: the uropathogenic lifestyle of Proteus mirabilis [J]. Trends in Microbiology, 2017, 25(4): 304-315.

[63]

YUAN

, HUANG

, YANG

, WANG

, LI

, YANG

, LI

. Pathogenesis of Proteus mirabilis in catheter-associated urinary tract infections[J]. Urologia Internationalis, 2021, 105(5/6): 354-361.

[64]

SCHAFFER

, NORSWORTHY

, SUN

, PEARSON

. Proteus mirabilis fimbriae- and urease-dependent clusters assemble in an extracellular niche to initiate bladder stone formation[J]. Proceedings of the National Academy of Sciences of the United States of America, 2016, 113(16): 4494-4499.

[65]

DAHLÉN

, HASSAN

, BLOMQVIST

, CARLÉN

. Rapid urease test (RUT) for evaluation of urease activity in oral bacteria in vitro and in supragingival dental plaque ex vivo [J]. BMC Oral Health, 2018, 18(1): 89.

[66]

D’SOUZA

, LAWANDE

, SAMUEL

, WISEMAN PINTO

. Effect of salivary urea, pH and ureolytic microflora on dental Calculus formation and its correlation with periodontal status[J]. Journal of Oral Biology and Craniofacial Research, 2023, 13(1): 8-12.

[67]

JHA D. Influence of salivary pH and urea level on Calculus formation: a clinico-biochemical study[J]. RGUHS Journal of Dental Sciences, 2021, 13: 2-6.

[68]

BONTOGNALI

TRR

, McKENZIE

, WARTHMANN

, VASCONCELOS

. Microbially influenced formation of Mg-calcite and Ca-dolomite in the presence of exopolymeric substances produced by sulphate-reducing bacteria[J]. Terra Nova, 2014, 26(1): 72-77.

[69]

WEI

, XIAO

, DONG

, CHEN

. Carbonate polymorph formation in microbially induced calcium carbonate precipitation (MICP): influencing factors, mechanisms, and knowledge gaps[J]. Earth-Science Reviews, 2025, 265: 105129.

[70]

WANG

, ZHANG

, DUAN

, SHI

, ZHANG

, GUAN

, SAND

, HOU

. Extracellular polymeric substances and biocorrosion/biofouling: recent advances and future perspectives[J]. International Journal of Molecular Sciences, 2022, 23(10): 5566.

[71]

FANG

, HUANG

, WEI

, LIANG

, RONG

, CHEN

, CAI

. Microcalorimetric and potentiometric titration studies on the adsorption of copper by extracellular polymeric substances (EPS), minerals and their composites[J]. Bioresource Technology, 2010, 101(15): 5774-5779.

[72]

KANG

, QU

, ALVAREZ

PJJ

, ZHU

. Extracellular saccharide-mediated reduction of Au³⁺ to gold nanoparticles: new insights for heavy metals biomineralization on microbial surfaces[J]. Environmental Science & Technology, 2017, 51(5): 2776-2785.

[73]

, YAO

, WANG

, HUANG

, FU

, ZHOU

. Insights into the formation mechanism of vaterite mediated by a deep-sea bacterium Shewanella piezotolerans WP3[J]. Geochimica et Cosmochimica Acta, 2019, 256: 35-48.

[74]

IVANOVA

, EGOROV

, ZABRODSKAYA

, SHALDZHYAN

, BARANCHIKOV

, TSVIGUN

, LYKHOLAY

, YAPRYNTSEV

, LEBEDEV

, KULMINSKAYA

. Matrix is everywhere: extracellular DNA is a link between biofilm and mineralization in Bacillus cereus planktonic lifestyle[J]. NPJ Biofilms and Microbiomes, 2023, 9: 9.

[75]

LIU

, HUANG

, ZHANG

, SONG

, HE

, WANG

, LIAN

. Bio-mineralisation, characterization, and stability of calcium carbonate containing organic matter[J]. RSC Advances, 2021, 11(24): 14415-14425.

[76]

LOU

, CHANG

, CUI

, QIAN

, HAO

, ZHANG

. Microbiologically influenced corrosion inhibition induced by S. putriefaciens mineralization under extracellular polymeric substance regulation via FlrA and FlhG genes[J]. Corrosion Science, 2023, 221: 111350.

[77]

GIUFFRE

, HAMM

, HAN

, de YOREO

, DOVE

. Polysaccharide chemistry regulates kinetics of calcite nucleation through competition of interfacial energies[J]. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(23): 9261-9266.

[78]

DITTRICH

, KURZ

, WEHRLI

. The role of autotrophic picocyanobacteria in calcite precipitation in an oligotrophic lake[J]. Geomicrobiology Journal, 2004, 21(1): 45-53.

[79]

SILHAVY

, KAHNE

, WALKER

. The bacterial cell envelope[J]. Cold Spring Harbor Perspectives in Biology, 2010, 2(5): a000414.

[80]

SOUTHAM

. Bacterial surface-mediated mineral formation[M]//Environmental Microbe-Metal Interactions. Washington, DC, USA: ASM Press, 2014: 257-276.

[81]

, HE

, REN

, DONG

, LIAN

. Mechanisms of carbonate precipitation induced by two model bacteria[J]. Chemical Geology, 2023, 628: 121461.

[82]

DUNHAM-CHEATHAM

, RUI

, BUNKER

, MENGUY

, HELLMANN

, FEIN

. The effects of non-metabolizing bacterial cells on the precipitation of U, Pb and Ca phosphates[J]. Geochimica et Cosmochimica Acta, 2011, 75(10): 2828-2847.

[83]

THOMAS

, RICE

. Revised model of calcium and magnesium binding to the bacterial cell wall[J]. BioMetals, 2014, 27(6): 1361-1370.

[84]

Von KÜGELGEN

, TANG

, HARDY

, KUREISAITE-CIZIENE

, BRUN

, STANSFELD

, ROBINSON

, BHARAT

TAM

. In situ structure of an intact lipopolysaccharide-bound bacterial surface layer[J]. Cell, 2020, 180(2): 348-358.e15.

[85]

SÁRA

, SLEYTR

. Crystalline bacterial cell surface layers (S-layers): from cell structure to biomimetics[J]. Progress in Biophysics and Molecular Biology, 1996, 65(1/2): 83-111.

[86]

BHARAT

TAM

, von KÜGELGEN

, ALVA

. Molecular logic of prokaryotic surface layer structures[J]. Trends in Microbiology, 2021, 29(5): 405-415.

[87]

SMIT

, OLING

, DEMEL

, MARTINEZ

, POUWELS

. The S-layer protein of Lactobacillus acidophilus ATCC 4356: identification and characterisation of domains responsible for S-protein assembly and cell wall binding[J]. Journal of Molecular Biology, 2001, 305(2): 245-257.

[88]

GERBINO

, CARASI

, MOBILI

, SERRADELL

, GÓMEZ-ZAVAGLIA

. Role of S-layer proteins in bacteria[J]. World Journal of Microbiology and Biotechnology, 2015, 31(12): 1877-1887.

[89]

SCHULTZE-LAM

, HARAUZ

, BEVERIDGE

. Participation of a cyanobacterial S layer in fine-grain mineral formation[J]. Journal of Bacteriology, 1992, 174(24): 7971-7981.

[90]

KISH

, MIOT

, LOMBARD

, GUIGNER

, BERNARD

, ZIRAH

, GUYOT

. Preservation of archaeal surface layer structure during mineralization[J]. Scientific Reports, 2016, 6: 26152.

[91]

SIMON

, POMPE

, GRUNER

, STURM

, OSTERMANN

, MATYS

, VOGEL

, RÖDEL

. Nested formation of calcium carbonate polymorphs in a bacterial surface membrane with a graded nanoconfinement: an evolutionary strategy to ensure bacterial survival[J]. ACS Biomaterials Science & Engineering, 2022, 8(2): 526-539.

[92]

REN

, ZHOU

, LONG

, XIE

, KANKALA

, WANG

, ZHANG

, LIU

. Biomedical applications of magnetosomes: state of the art and perspectives[J]. Bioactive Materials, 2023, 28: 27-49.

[93]

GROVER

VPB

, TOGNARELLI

, CROSSEY

MME

, COX IJ, TAYLOR-ROBINSON

, McPHAIL

MJW

. Magnetic resonance imaging: principles and techniques: lessons for clinicians[J]. Journal of Clinical and Experimental Hepatology, 2015, 5(3): 246-255.

[94]

KUPFER

, OGLE

. Advanced imaging approaches for regenerative medicine: emerging technologies for monitoring stem cell fate in vitro and in vivo [J]. Biotechnology Journal, 2015, 10(10): 1515-1528.

[95]

YAN

, DA HY, ZHANG

, LÓPEZ

, WANG

. Bacterial magnetosome and its potential application[J]. Microbiological Research, 2017, 203: 19-28.

[96]

ERDAL

, DEMIRBILEK

, YEH

, AKBAL

, RUFF

, BOZKURT

, CABUK

, SENEL

, GUMUSKAYA

, ALGIN

, COLAK

, ESENER

, DENKBAS

. A comparative study of receptor-targeted magnetosome and HSA-coated iron oxide nanoparticles as MRI contrast-enhancing agent in animal cancer model[J]. Applied Biochemistry and Biotechnology, 2018, 185(1): 91-113.

[97]

MÉRIAUX

, BOUCHER

, MARTY

, LALATONNE

, PRÉVÉRAL

, MOTTE

, LEFÈVRE

, GEFFROY

, LETHIMONNIER

, PÉAN

, GARCIA

, ADRYANCZYK-PERRIER

, PIGNOL

, GINET

. Magnetosomes, biogenic magnetic nanomaterials for brain molecular imaging with 17.2 T MRI scanner[J]. Advanced Healthcare Materials, 2015, 4(7): 1076-1083.

[98]

, XU

, TAO

, QIAN

, CUI

, REHMAN

, ZHU

, CHEN

, ZHAO

, WANG

, QI

, DAI

, ZHANG

, XIE

, LU

, WANG

. Magnetosome-inspired synthesis of soft ferrimagnetic nanoparticles for magnetic tumor targeting[J]. Proceedings of the National Academy of Sciences of the United States of America, 2022, 119(45): e2211228119.

[99]

CHEN

, SUN

, LIN

, ZHAO

, ZHOU

, WU

, ZHAO

, LIU

. Generating theranostic bacteria by depositing a protective visual therapeutic nanocoating via in situ sequential mineralization[J]. Science Advances, 2025, 11(41): eady9183.

[100]

LIU

, ZHANG

, WANG

, ZHU

, LI

, MA

, ZHANG

, CHEN

, TIWARI

, SHI

, ZHANG

, FAN

, ZHAO

, LIANG

. Comprehensive understanding of magnetic hyperthermia for improving antitumor therapeutic efficacy[J]. Theranostics, 2020, 10(8): 3793-3815.

[101]

VILLANUEVA

, GUBIEDA A

, GANDARIAS

, ABAD DÍAZ de CERIO A, ORUE

, ÁNGEL GARCÍA

, de

COS D

, ALONSO

, FDEZ-GUBIEDA

. Heating efficiency of different magnetotactic bacterial species: influence of magnetosome morphology and chain arrangement[J]. ACS Applied Materials & Interfaces, 2024, 16(49): 67216-67224.

[102]

CHEN

, WANG

, CHEN

, WANG

, HALGAMUGE

, CHEN

, SONG

. Smart magnetotactic bacteria enable the inhibition of neuroblastoma under an alternating magnetic field[J]. ACS Applied Materials & Interfaces, 2022, 14(12): 14049-14058.

[103]

ALONSO

, GANDÍA

, JEFREMOVAS

, GANDARIAS

, VILLANUEVA

, LETE

, MARCANO

, GARCÍA-PRIETO

, ORUE

, BARQUÍN

, de COS ELICES

, GARCÍA-ARRIBAS

, MUELA

, FDEZ-GUBIEDA

. Magnetotactic bacteria: biorobots for targeted therapies and model nanomagnetic systems[C]//2023 IEEE International Magnetic Conference - Short Papers (INTERMAG Short Papers). Sendai, Japan: IEEE, 2023: 1-2.

[104]

STANTON

, PARK

, VILELA

, BENTE

, FAIVRE

, SITTI

, SÁNCHEZ

. Magnetotactic bacteria powered biohybrids target E. coli biofilms[J]. ACS Nano, 2017, 11(10): 9968-9978.

[105]

CHEN

, LIU

, FAN

, PENG

, WANG

, ZHANG

, LIU

, ZHANG

. Self-mineralized photothermal bacteria hybridizing with mitochondria-targeted metal-organic frameworks for augmenting photothermal tumor therapy[J]. Advanced Functional Materials, 2020, 30(14): 1909806.

[106]

GUO

, DING

, ZHOU

. Converting bacteria into autologous tumor vaccine via surface biomineralization of calcium carbonate for enhanced immunotherapy[J]. Acta Pharmaceutica Sinica B, 2023, 13(12): 5074-5090.

[107]

GENG

, WANG

, WU

, CAO

, LIU

. Biointerface mineralization generates ultraresistant gut microbes as oral biotherapeutics[J]. Science Advances, 2023, 9(11): eade0997.

[108]

ZHANG

, LIU

, ZHANG

, LU

, LI

, HUANG

, ZHANG

, LI

, LU

, MING

, YANG

, SHOU

, WANG

, XIA

, XU

, WANG

. Inducing bacterial calcification for systematic treatment and immunomodulation against methicillin-resistant Staphylococcus aureus [J]. Nature Biotechnology, 2025: 1-13.

[109]

WANG

, ZHONG

, XU

, ZHUANG

, GONG

, CHEN

, TAO

, HU

, HUANG

, YANG

, LI

, ZHAO

, SUN

, HUO

, CHEN

, ZHAO

, PENG

, LIU

. Oncolytic mineralized bacteria as potent locally administered immunotherapeutics[J]. Nature Biomedical Engineering, 2024, 8(5): 561-578.

[110]

PLAN SANGNIER

, PREVERAL

, CURCIO

, SILVA

AKA

, LEFÈVRE

, PIGNOL

, LALATONNE

, WILHELM

. Targeted thermal therapy with genetically engineered magnetite magnetosomes@RGD: photothermia is far more efficient than magnetic hyperthermia[J]. Journal of Controlled Release, 2018, 279: 271-281.

[111]

WANG

, GENG

, ZHANG

, WANG

, LIU

, BASIT

, MIAO

, LIU

, JIANG

. Bacterial magnetosomes loaded with doxorubicin and transferrin improve targeted therapy of hepatocellular carcinoma[J]. Nanotheranostics, 2019, 3(3): 284-298.

[112]

, HU

, LIU

, LI

, WANG

, ZHANG

, JIANG

, TIAN

, LI

. Surface expression of protein A on magnetosomes and capture of pathogenic bacteria by magnetosome/antibody complexes[J]. Frontiers in Microbiology, 2014, 5: 136.

[113]

BIRNBAUM

, MANJULA-BASAVANNA

, KAN

, TARDY

, JOSHI

. Hybrid living capsules autonomously produced by engineered bacteria[J]. Advanced Science, 2021, 8(11): 2004699.

[114]

FAN

, LI

, LIU

, ZHENG

, CHEN

, ZHANG

. Bacteria-mediated tumor therapy utilizing photothermally-controlled TNF-α expression via oral administration[J]. Nano Letters, 2018, 18(4): 2373-2380.

[115]

MERROUN

, NEDELKOVA

, OJEDA

, REITZ

, FERNÁNDEZ

, ARIAS

, ROMERO-GONZÁLEZ

, SELENSKA-POBELL

. Bio-precipitation of uranium by two bacterial isolates recovered from extreme environments as estimated by potentiometric titration, TEM and X-ray absorption spectroscopic analyses[J]. Journal of Hazardous Materials, 2011, 197: 1-10.

[116]

MATHURIYA

, YADAV

, KAUSHIK

. Magnetotactic bacteria: performances and challenges[J]. Geomicrobiology Journal, 2015, 32(9): 780-788.

[117]

LIU

, YUAN

, BREMMER

, HU

. Convergence of nanotechnology and bacteriotherapy for biomedical applications[J]. Advanced Science, 2024, 11(16): 2309295.

2026年第66卷第2期

PDF下载

236

引用本文

BibTeX

文章信息

doi: 10.13343/j.cnki.wsxb.20250652

接收时间：2025-08-24
首发时间：2026-02-05
出版时间：2026-02-04

补充材料

相关文章

文章信息

作者

出版历史

收稿日期：2025-08-24
录用日期：2025-11-10

基金

the Major Program of National Natural Science Foundation of China(81991501)

国家自然科学基金重大项目(81991501)

the Sichuan Provincial Natural Science Foundation(2024NSFSC0546)

四川省自然科学基金(2024NSFSC0546)

作者信息

^1.四川大学华西口腔医院，牙体牙髓病科，口腔疾病防治全国重点实验室，国家口腔医学中心，口腔疾病国家临床医学研究中心，四川成都

^2.四川大学华西口腔医院，口腔疾病防治全国重点实验室，口腔疾病国家临床医学研究中心，四川成都

参考文献

分享链接

https://castjournals.cast.org.cn/joweb/wswxb/CN/10.13343/j.cnki.wsxb.20250652

分享至

全文二维码

扫描看全文

引用本文

BibTeX

本文的引用情况

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

关闭全屏

BibTeX
EndNote
RefWorks
TxT

Bacteria type	Representative species	Mineral products	Mineralization mechanism	References
Urease-producing bacteria (UPB)	Sporosarcina pasteurii, Staphylococcus sp. H6, Serratia marcescens, Enterobacter cloacae EMB19	Calcite, aragonite, Cd-Ca co-precipitates	Urease-catalyzed urea hydrolysis produces NH₃ and CO₃^2-, elevating local pH and inducing precipitation of CaCO₃ and metal ions (e.g., Cd-Ca co-precipitates)	[50-52]
Sulfate-reducing bacteria (SRB)	Desulfovibrio desulfuricans, Desulfobulbus propionicus	CdS, PbS	Sulfate-reducing bacteria reduce SO₄^2- to S^2- using lactate as an electron donor, decreasing local redox potential and inducing metal sulfide precipitation	[53]
Phosphate-solubilizing bacteria (PSB)	Burkholderia ambifaria QY14	Cd-HAP	Increased acid phosphatase activity hydrolyzes organic phosphates, releasing PO₄^3- that combines with Cd²⁺ and Ca²⁺ to precipitate as Cd-HAP	[54]
Nitrate-reducing bacteria (NRB)	Klebsiella aerogenes	Green rust, goethite	Nitrate-reducing bacteria reduce NO₃⁻ to NO₂⁻ using organic carbon as an electron donor. The produced NO₂⁻ non-enzymatically oxidizes Fe²⁺ to Fe³⁺, inducing Fe²⁺/Fe³⁺ co-precipitation as green rust and goethite	[55]
Fe-oxidizing bacteria (FOB)	Ochrobactrum sp. EEELCW01	Goethite, Fe-As minerals	Iron-oxidizing bacteria oxidize Fe²⁺ to Fe³⁺ under nitrate-reducing conditions, promoting Fe³⁺ mineral formation (e.g., goethite) and co-precipitation with As to form Fe-As minerals	[56]
Fe-reducing bacteria (FRB)	Shewanella oneidensis MR-4	Siderite (FeCO₃)	Iron-reducing bacteria reduce Fe³⁺ to Fe²⁺, which combines with CO₃^2- to form siderite	[57]

Bacteria type

Representative species

Mineral products

Mineralization mechanism

References

Urease-producing bacteria (UPB)

Sporosarcina pasteurii, Staphylococcus sp. H6, Serratia marcescens, Enterobacter cloacae EMB19

Calcite, aragonite, Cd-Ca co-precipitates

Urease-catalyzed urea hydrolysis produces NH₃ and CO₃^2-, elevating local pH and inducing precipitation of CaCO₃ and metal ions (e.g., Cd-Ca co-precipitates)

[50-52]

Sulfate-reducing bacteria (SRB)

Desulfovibrio desulfuricans, Desulfobulbus propionicus

CdS, PbS

Sulfate-reducing bacteria reduce SO₄^2- to S^2- using lactate as an electron donor, decreasing local redox potential and inducing metal sulfide precipitation

[53]

Phosphate-solubilizing bacteria (PSB)

Burkholderia ambifaria QY14

Cd-HAP

Increased acid phosphatase activity hydrolyzes organic phosphates, releasing PO₄^3- that combines with Cd²⁺ and Ca²⁺ to precipitate as Cd-HAP

[54]

Nitrate-reducing bacteria (NRB)

Klebsiella aerogenes

Green rust, goethite

Nitrate-reducing bacteria reduce NO₃⁻ to NO₂⁻ using organic carbon as an electron donor. The produced NO₂⁻ non-enzymatically oxidizes Fe²⁺ to Fe³⁺, inducing Fe²⁺/Fe³⁺ co-precipitation as green rust and goethite

[55]

Fe-oxidizing bacteria (FOB)

Ochrobactrum sp. EEELCW01

Goethite, Fe-As minerals

Iron-oxidizing bacteria oxidize Fe²⁺ to Fe³⁺ under nitrate-reducing conditions, promoting Fe³⁺ mineral formation (e.g., goethite) and co-precipitation with As to form Fe-As minerals

[56]

Fe-reducing bacteria (FRB)

Shewanella oneidensis MR-4

Siderite (FeCO₃)

Iron-reducing bacteria reduce Fe³⁺ to Fe²⁺, which combines with CO₃^2- to form siderite

[57]