Serine-473 is phosphorylated by Tor complex 2, (TORC2) [46], and in yeast and human cancer cells, TORC2 phosphorylation of Akt is stimulated by mature ribosomes, which can bind to both TORC2 and Akt to mediate their interaction, and TORC2 activity is blocked by defects in ribosome biogenesis [60]. This evidence concerns the gene CRTC2 and cancer.