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While genome-enabled studies have identified core biosynthetic genes (e.g., CHS-like polyketide synthases) and strong co-expression modules, direct demonstration of specific transcription factors binding and activating anthraquinone biosynthetic promoters is still scarce ( Rheum species has nominated candidate R2R3-MYB and bZIP transcription factors correlated with anthraquinone accumulation, but these studies primarily provide association and expression-based evidence rather than direct TFpromoter validation (Zhao et al., 2024