Copper also plays a pivotal role in the nucleus.32 Specifically, CCS and ATOX1 are both involved in transporting copper to the cell nucleus, where it is essential for activating different transcription factors.95–97 In human hepatocellular carcinoma (HCC) cells, the expression of genes induced under low oxygen conditions relies on the presence of CCS and copper. This evidence concerns the gene ATOX1 and hepatocellular carcinoma.