The increased intracellular ratio L-lactic acid/Glucose monophosphate and D-Fructose monophosphate in all 3D tumor spheroids compared to their relative 2D cultures well agreed with the up-regulation of the glycolytic enzymes ALDOC and ENO2, at both gene and protein levels (Fig. 4C; Fig. S2 in additional file 8). The gene discussed is ALDOC; the disease is neoplasm.