Background: Cancer metabolism is highly heterogenous and flexible with the Warburg effect or oxidative phosphorylation (OXPHOS) prevailing in a cancer type- and context-dependent manner. Past studies have demonstrated that targeting OXPHOS robustly inhibits glycolysis-deficient cancer cell viability and tumorigenicity. However, the therapeutic potential of OXPHOS inhibition in metabolically flexible glycolysis-competent cancers is unclear. Moreover, whether the depletion of OXPHOS-derived ATP or the abolition of OXPHOS-supported biosynthesis is the major determinant of cancer cell susceptibility remains obscure.
Methods: A panel of metabolically flexible glycolysis-competent cancer cell lines along with those deficient in glycolysis were tested using OXPHOS inhibitors, including the mitochondrial complex I inhibitor IACS-010759. Metabolic phenotypes were determined using Seahorse metabolic flux assays. Targeted metabolomics was conducted using both GC- and LC-MS. Stable isotope tracing was carried out with uniformly labeled [15N]-/[13C]-aspartate. Patient-derived xenograft (PDX) models of colorectal cancer in NSG mice were used for in vivo validation.
Results: OXPHOS inhibition potently diminished metabolically flexible glycolysis-competent cancer cell proliferation and tumorigenicity without causing devastating energy stress. This was associated with S-phase cell cycle arrest and the enrichment of the G2/M DNA-damage checkpoint regulation pathway, suggestive of replication stress. Indeed, IACS treatment significantly reduced the purine/pyrimidine nucleotide pools, which was primarily caused by aspartate deficiency resulting from a shortage in the electron acceptor NAD+. The supplementation of exogenous nucleosides, aspartate, or pyruvate that can accept electron-generating NAD+, into the culture medium rescued cells from IACS-induced cell cycle arrest. Instructively, inhibition of GOT1, which catalyzes cytosolic aspartate biosynthesis when mitochondrial aspartate production is dampened, rendered cancer cells more susceptible to OXPHOS inhibition.
Conclusion: 1) Disruption of nucleotide homeostasis is a major determinant of cancer cell susceptibility to OXPHOS inhibition; 2) OXPHOS inhibition is a promising avenue for the treatment of cancers that are metabolic flexible and glycolysis competent, and 3) GOT1 targeting is potentially a useful approach to improve the therapeutic efficacy of OXPHOS inhibition for cancer treatment.