MSX2 and neoplasm: In the metastasis-associated neoplastic cell states, we detected robust expression of neural crest (ZIC1, OLIG3), mesenchymal (MSX2, GDF6), and metastasis-related (DKK2) genes in addition to adrenergic genes (e.g., PHOX2B) that were shared with primary tumor cell states (Fig. 3C).