The sulfation patterns and chain lengths of proteoglycans can be modified within the tumor milieu by N-deacetylase/N-sulfotransferases (NDSTs), O-sulfotransferases (HS2ST1, HS3STs, HS6STs), endosulfotransferases (SULF1 and SULF2), heparanases (HPSE), sheddases and sulfatases (Figure 2B) [99]. Here, HPSE is linked to neoplasm.