In fact, spatial visualization and quantification of Wnt target genes in tumoral and non-tumoral regions, including both hepatocyte and nonparenchymal cell populatons, revealed that β-catenin-mutated tumor cells are defined by expression of bonafide Wnt targets Glul, Tbx3, Axin2, Lgr5, Lect2, and Ccnd1 (Supplementary Fig. 11a–c) with their identity intimately linked to zone 3 metabolic genes (and processes), including Cyp2e1, Cyp1a2, and Oat, with exclusion of zone 1 metabolic genes (and processes), including Cyp2f2, Ass1, and Arg1 (Supplementary Fig. 12a, b). Here, GLUL is linked to neoplasm.