Consistent with the signalling pathways upregulated in aggressive T1 tumours (Fig. 1), VilCreER;Apcfl/fl;KrasG12D/+;Alk5CA mice exhibit a significant enrichment of TGFβ signalling (Fig. 6e; Fessler et al. TGFβ-responsive gene signature and Hallmark TGFβ-pathway classifier22,28), alongside upregulation of KRAS and mTORC1 signalling (Fig. 6f; Hallmark KRAS and mTORC1 pathway gene sets28). Here, TGFB1 is linked to neoplasm.