Copper (Cu) is an essential micronutrient for plant growth; however, excessive accumulation of Cu resulting from the intensive use of Cu-based agrochemicals has become a major environmental concern, adversely affecting crop productivity and ecosystem sustainability. Developing eco-friendly strategies to mitigate Cu toxicity is therefore crucial for sustainable agriculture. This study evaluated fungal VOCs from Fusarium verticillioides and Prunella vulgaris extract (Pv) as biostimulants to improve copper (Cu) tolerance in rice (Oryza sativa L. cv. TNG67).
Exposure to FvVCs promoted shoot elongation while inhibiting root growth. Transcriptomic analysis revealed that FvVCs induced tissue-specific gene expression patterns, including the upregulation of genes associated with cytokinin signaling, cell division, and ribosome biogenesis in shoots, whereas genes involved in ethylene biosynthesis, auxin synthesis, and abscisic acid signaling were upregulated in roots. Furthermore, pretreatment with FvVCs enhanced seedling survival under Cu stress and increased proline accumulation, suggesting that fungal volatiles function as signaling molecules that prime plant defense responses and improve stress adaptation.
In other way, Pv treatment markedly improved plant growth and reduced Cu-induced damage. Transcriptomic and molecular analyses demonstrated that Pv stimulated the expression of genes involved in cell wall remodeling, Casparian strip formation, proline biosynthesis, deoxymugineic acid production, metallochaperones, and vacuolar and efflux transporters. These responses contributed to enhanced metal sequestration, reactive oxygen species detoxification, while limiting Cu uptake, thereby mitigating oxidative damage and improving Cu tolerance. LC–MS analysis identified several bioactive compounds, including caffeic acid, rosmarinic acid, and melatonin, which may contribute to the protective effects of Pv. Among the tested concentrations, 0.25% (w/v) Pv was the most effective in enhancing Cu tolerance.
Overall, the results demonstrate that both fungal VOCs and P. vulgaris extract improve rice adaptation to Cu stress through coordinated physiological, anatomical, and molecular mechanisms. While FvVCs primarily enhance stress tolerance through defense priming and hormonal regulation, Pv mitigates Cu toxicity by restricting metal accumulation and promoting detoxification processes. These findings provide new insights into plant responses to biologically derived signaling compounds and biostimulants and highlight their potential as sustainable alternatives to conventional chemical inputs for improving crop performance in Cu-contaminated agricultural soils.

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