Three transcription factors were identified as drivers of the SPF UP cluster (Extended Data Fig. 2g and Supplementary Table 5), namely E2f4 (control of cell cycle, increased expression in cancers), Foxm1 (oncogenic transcription factor, control of cell cycle, regulation of CRC metastasis and progression) and Nfyb (promotes invasion, metastasis and differentiation, associated with multiple cancers, activator or repressor in a context-dependent manner). Here, E2F4 is linked to cancer.