蜡样芽孢杆菌和金黄色葡萄球菌对高岭土的铁浸出特性研究

高岭土是一种化学性质稳定的硅酸盐矿物，具有良好的吸附性、可塑性、分散性，用作颜料或增量剂时具有良好的遮盖力，通常作为填料和涂层材料应用于陶瓷、化妆品、造纸等工业领域。白度是评价高岭土品质的重要指标，然而天然高岭土中普遍存在三价铁杂质，降低了高岭土的白度并影响其性能。传统的物理和化学除铁方法存在成本高、污染环境、破坏高岭土结构等缺点。因此基于微生物的生物浸出法已成为高岭土行业的研究热点。目前微生物浸出加工使用的多为从自然环境中分离得到的革兰氏阴性菌，而对革兰氏阳性菌铁浸出机制和潜在应用研究较少。

本研究选用革兰氏阳性菌蜡样芽胞杆菌Bacillus cereus（B. cereus）和金黄色葡萄球菌Staphylococcus aureus（S. aureus）为实验菌株，通过调控浸出环境中的氮源碳源成分，分析了B. cereus和S. aureus对高岭土中晶态和非晶态铁杂质的还原浸出特性，通过监测细菌培养环境中的生化指标阐明铁还原浸出机制，利用工程化方法提高细菌的铁还原能力。本文主要研究内容及结果如下：

（1）探究B. cereus和S. aureus对晶态、非晶态铁氧化物的还原能力。实验中选取Fe₂O₃和高岭土作为铁还原目标，通过监测细菌浸出过程中的生化指标，分析B. cereus和S. aureus对晶态和非晶态铁氧化物的铁还原行为。实验结果显示B. cereus对晶态铁氧化物具有还原活性，其还原Fe₂O₃时培养基中亚铁离子总积累值最高达到了3 mg/L，而S. aureus仅能还原高岭土中非晶态三价铁杂质，培养基中亚铁离子总积累值最高达到0.6 mg/L。研究表明铁载体的类型和结合效率是影响革兰氏阳性菌还原高岭土中铁杂质的重要因素。

（2）探究培养基中氮源碳源成分对B. cereus和S. aureus铁还原能力的影响。实验中通过监测细菌浸出过程中的生化指标，分析革兰氏阳性菌B. cereus和S. aureus在氮源和碳源辅助氮源生长环境中的铁还原行为，揭示培养基成分与革兰氏阳性菌铁还原行为之间的关系。实验结果显示B. cereus在氮源成分稀释的0.1×LB培养基中还原晶态铁氧化物时达到最高亚铁离子浓度3 mg/L，S. aureus仅能在1×LB培养基中还原高岭土中非晶态铁氧化物。氮源培养基中添加碳源淀粉进一步提高了B. cereus的铁还原能力，亚铁离子总积累值最高提升超过0.6 mg/L。研究表明培养基中氮源碳源新陈代谢产物改变了细胞环境酸碱度，影响细胞铁载体的产生和对铁的结合能力，调节细菌的铁还原行为。

（3）探究革兰氏阳性菌B. cereus和S. aureus的铁还原通路，通过工程化提高B. cereus和S. aureus的铁还原能力。实验中利用实时荧光定量PCR分析不同培养基成分下生长细菌的铁还原基因表达水平，绘制革兰氏阳性菌还原浸出三价铁的通路，探究革兰氏阳性菌的铁还原机制。实验结果显示B. cereus的铁载体合成基因asb与细胞内外稳态调节基因sigB和napA共同调控B. cereus的铁还原过程。S. aureus铁还原相关基因包括负责三价铁转运和内化的Isd系统与Fur调节因子，负责铁载体生物合成的SbnI，负责pH稳态调节的DacA。研究表明革兰氏阳性菌B. cereus和S. aureus通过培养基中氮源碳源代谢产物改变周围环境的酸碱度，引起细胞pH响应，并利用能量代谢和pH稳态调节来影响铁载体的合成和铁复合物的转运还原。铁载体复合物进入细胞后最终被还原型烟酰胺腺嘌呤二核苷酸磷酸（NADPH）还原，释放出亚铁离子。通过转入铁载体合成基因重组质粒，B. cereus和S. aureus铁还原环境中的亚铁离子浓度最高提升50％。

本研究揭示了革兰氏阳性菌B. cereus和S. aureus的铁还原浸出机制，发现其与革兰氏阴性菌的电子传递机制不同，主要通过铁载体转运还原铁氧化物。通过工程化提高了铁载体合成量，增强B. cereus和S. aureus的铁还原能力。本研究有助于优化提高革兰氏阳性菌的铁浸出效率，为革兰氏阳性菌在生物浸出工业中的应用提供重要参考。

Kaolin is a chemically stable silicate mineral with good adsorption, plasticity and dispersion, and has good hiding power when used as pigment or additive. It is usually used as filler and coating material in ceramics, cosmetics, paper making and other industrial fields. Whiteness is an important index to evaluate the quality of kaolin. However, the presence of ferric impurities in natural kaolin reduces the whiteness of kaolin and affects its properties. The traditional physical and chemical methods of iron removal have disadvantages such as high cost, pollution of environment and destruction of kaolin structure. Therefore, bioleaching based on microorganism has become the research hotspot in kaolin industry. At present, most of the microorganisms used in microbial leaching process are Gram-negative bacteria isolated from natural environment, but there are few researches on the mechanism and potential application of iron leaching of Gram-positive bacteria.

This study selected the Gram-positive bacteria Bacillus cereus (B. cereus) and Staphylococcus aureus (S. aureus) as experimental strains, by regulating the composition of nitrogen and carbon sources in the leaching environment, the reduction leaching characteristics of B. cereus and S. aureus on crystalline and amorphous iron impurities in kaolin were analyzed. And the iron reductive leaching mechanism was elucidated by monitoring the biochemical indicators in the bacterial culture environment. The iron reduction abilities of bacteria were improved by engineering bacteria. The main research content and results of this thesis are as follows:

(1) Investigate the reduction ability of B. cereus and S. aureus on crystalline and amorphous ferric iron. In the experiment, Fe₂O₃ and kaolin were selected as iron reduction targets, and the iron reduction behaviors of B. cereus and S. aureus on crystalline and amorphous ferric oxides were analyzed by monitoring the biochemical indicators in the bacterial leaching process. The experimental results showed that B. cereus had reduction activity on crystalline iron oxides, and the total accumulation value of ferrous ions in the medium of Fe₂O₃ reduction reached 3 mg/L, while S. aureus could only reduce amorphous ferric impurities in kaolin, and the total accumulation value of ferrous ions in the medium reached 0.6 mg/L. The results showed that the type and binding efficiency of siderophores are important factors affecting the reduction of iron impurities in kaolin by Gram-positive bacteria.

(2) Investigate the effects of nitrogen and carbon compositions on the iron reduction abilities of B. cereus and S. aureus. In the experiment, by monitoring the biochemical indicators in the process of bacterial leaching, the iron reduction behaviors of Gram-positive bacteria B. cereus and S. aureus in the growth environment of nitrogen source and carbon source-assisted nitrogen source were analyzed, revealing the relationship between the medium composition and iron-reducing behaviors of Gram-positive bacteria. The experimental results showed that B. cereus achieved the highest concentration of ferrous ion 3 mg/L when reducing crystalline iron oxides in 0.1´LB medium with diluted nitrogen source composition, while S. aureus could only reduce amorphous iron oxide of kaolin in 1´LB medium. The addition of carbon source starch in nitrogen source medium further improved the iron reduction ability of B. cereus, and the maximum increase of the total accumulation value of ferrous ions was more than 0.6 mg/L. The results showed that the metabolic products of nitrogen source and carbon source in the medium changed the pH of the cell environment, affected the production of siderophore and its binding ability to iron, and regulated the iron reduction behaviors of bacteria.

(3) Investigate the iron reduction pathway of Gram-positive bacteria B. cereus and S. aureus, and improve their iron reduction ability through bacterial engineering. In the experiment, real-time fluorescent quantitative PCR was used to analyze the iron-reducing gene expression level of bacteria grown under different medium compositions, and the pathway of ferric iron reduction by Gram-positive bacteria was mapped to explore their iron reduction mechanism. The experimental results showed that the siderophore synthesis gene cluster asb, the extracellular and extracellular homeostasis regulation genes sigB and napA jointly regulated the iron reduction process of B. cereus. S. aureus iron reduction-related genes include the Isd system and Fur regulator responsible for ferric iron transport and internalization, SbnI for siderophore biosynthesis, and DacA for pH homeostasis regulation. The results showed that Gram-positive bacteria B. cereus and S. aureus change the pH of the surrounding environment through the metabolites of nitrogen and carbon sources in the medium, causing cell pH response, affecting the syntheses of siderophore and the transport and reduction of iron complexes through the regulation of energy metabolism and pH homeostasis. The siderophore complexes enter the cell and are reduced by the reduced nicotinamide adenine dinucleotide phosphate (NADPH) to release ferrous ions. The concentration of ferrous ions in B. cereus and S. aureus iron-reducing environment was increased by up to 50% by transferring the siderophore synthesized gene recombinant plasmid.

In this study, the iron leaching mechanisms of Gram-positive bacteria B. cereus and S. aureus were revealed, and found that it is different from the electron transport mechanism of Gram-negative bacteria, the ferric oxides were mainly transported through siderophores. Through bacterial engineering, the synthesis of siderophores was increased and the iron reducing abilities of B. cereus and S. aureus were enhanced. This study is helpful to optimize and improve the iron leaching efficiency of Gram-positive bacteria, and provides an important reference for the application of Gram-positive bacteria in bioleaching industry.

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