TS_FR+ was characterized by enrichment of metabolic enzymes and ECM proteoglycan (ALDOC, KMO, TSKU) and featured pathways of lipid, amino acid, and nucleotide metabolism (Fig. 2e, f), consistent with the upregulation of various metabolic pathways in FR+ tumor epithelial, indicating a uniform metabolic preference for stroma and the tumor they reside in. Here, TSKU is linked to neoplasm.