Bioleaching of Copper Sulfide Ores: From Microbial Mechanisms to Industrial Applications

The global energy transition and rapid electrification are driving increased demand for copper. However, conventional pyrometallurgical and hydrometallurgical extraction routes are increasingly challenged by declining ore grades and stricter environmental regulations. Bioleaching involves the microbial catalysis of sulfide mineral dissolution and provides a sustainable method for copper recovery from low-grade ores, tailings and secondary resources. This review provides a critical and integrated analysis of copper sulfide bioleaching, covering microbial diversity, molecular mechanisms, mineralogical controls, operational parameters, and industrial applications. This review also examines the functional roles of prominent acidophiles, including the functional roles of prominent acidophiles, including Acidithiobacillus spp., Leptospirillum spp. and thermophilic archaea, in the oxidation of iron and sulfur, mitigation of passivation, and metal solubilization. The molecular underpinnings of these processes are explored by investigating iron and sulfur oxidation gene networks (the rus operon and sox cluster), copper resistance systems (CopA, CusCBA) and biofilm formation pathways. The mineralogical controls on the behavior of chalcopyrite (refractory/passivating), chalcocite (highly reactive) and bornite (intermediate) are critically assessed. The synergistic effects of key operational parameters (temperature, pH, redox potential, aeration and particle size) on leaching kinetics and microbial community dynamics are investigated. The scalability, efficiency and environmental footprint of industrial applications such as heap, dump, stirred-tank and in situ bioleaching are discussed. Despite more than four decades of commercial development, several challenges remain, such as slow chalcopyrite dissolution, passivation, metal toxicity, and scale-up limitations. Emerging solutions such as synthetic microbial consortia, multi-omics technologies, artificial intelligence-assisted optimization, and digital twins are identified as transformative approaches for next-generation biomining. In this review, microbiology, mineralogy, electrochemistry, and process engineering are integrated to demonstrate that biotechnological leaching is among the most promising technologies for the sustainable production of copper and to identify future directions for its industrial application.