Therefore, this study successfully elucidated the correlated extracellular and intracellular mechanisms of metal detoxification by yeast. However, the Ves started to deform and gradually lost their activity at 200 mg/L. Under transmission electron microscopy, the intracellular Ves showed evident sorption of Cu cations (100 mg/L). These accelerated the transport of Cu cations into intracellular, but the redox reaction in both cell membrane and intracellular region was limited. Moreover, the thickness of cell membrane decreased from 450 to 116 nm during the elevation of Cu stress. Then, the contrast of Cu concentration between the extracellular and intracellular regions was enlarged. Subsequently, the potential Haber-Weiss reaction in EPS lowered fungal ability to shield against the Cu toxicity. Moreover, redox peaks at − 0.35 V (reduction) and − 0.02 V (oxidation) in EPS were observed based on electrochemical analysis. The Cu removal was decreased from ~ 35 to ~ 0% as Cu stress raised from 0 to 200 mg/L, which confirmed the low binding of Cu cations to EPS. In particular, GSH content in EPS did not show significant changes. In contrast to other heavy metals (e.g., Pb²⁺), low Cu²⁺ stress has no evident stimulation to EPS production. The Cu²⁺ concentrations were set from 0 to 200 mg/L. This study explored the mechanisms of Rho to resist Cu toxicity from extracellular to intracellular, i.e., EPS, membrane, and Ves. The red yeast (Rhodotorula mucilaginosa: Rho) has abundant extracellular polymeric substances (EPS) and intracellular vesicles (Ves). Cluster analysis, based on Cu abundance levels, reveals that correlations exist between the chemical forms of Cu and their intracellular locations, which in turn indicates a relationship between their migration activity and toxicity. These results indicated that Cu ions were integrated with compounds of low bioavailable, such as undissolved phosphate or oxalate, which contributed to the metal tolerance and detoxification of E. In addition, the binding capacity of Cu to soluble salts or proteins decreased and its binding capacity to insoluble oxalate increased with the increasing Cu supply. The distribution characteristics of Cu, particularly in the roots, were closely related to the Cu tolerance of E. Most of the Cu in the roots, stems and leaves were extracted by 2 % hydrate acetic acid (HAc) in controls. splendens, the highest proportion of which was stored in its cell walls (68 %) and vacuoles (42 %). Various chemical forms of Cu were absorbed by E. This study develops a systematic method of detecting the intracellular localization and different chemical forms of Cu to comprehensively understand the mechanisms involved in Cu tolerance and detoxification of E. Elsholtzia splendens, a Cu-tolerant plant growing in Cu mine areas, can accumulate high levels of Cu. Copper (Cu) accumulation in soils is becoming a major environmental problem.
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