Interestingly, among the LOC upregulated genes that are involved in GTPase activity (Table S3A), S100A9 gene was found to be overexpressed in HCC 63 and promote cell proliferation as well as invasion64, 65, which is consistent with our observations from functional assays in LOC-overexpressing/knockdown cells (Figure 4A and B, 5C and D). Here, S100A9 is linked to hepatocellular carcinoma.