TOX and hepatocellular carcinoma: These C3 cells showed high expression of inhibitory immunoreceptors and exhaustion markers, such as Pdcd1, Havcr2, Ptpn11, Lag3, Tox and Tigit. These Pdcd1+ T cells also expressed Cd247, the gene encoding PD-L1, potentially providing an autologous ligand for PD-1 binding in addition to the PD-L1 expressed on APCs and HCC cells (Fig. 2f and Extended Data Fig. 2g).