Transcriptional characterization of signature genes in the three meta programs was performed and validated using the scATAC-seq and stRNA-seq data (Figure 4B, 4C and Supplementary Figure 4A–4D), with results disclosing that the metabolism meta program specifically expressed REG1A, a molecular marker for ccRCC [27]. The gene discussed is REG1A; the disease is nonpapillary renal cell carcinoma.