Table 2.

HS biosynthetic and modifying enzymes involved in cancer development and progression.

Enzyme	Gene	Type(s) of Cancer	Reference(s)
Xylosyltransferase1/2 (XYLT1/2)	XYLT1-2	Breast cancer/bone metastasis Salivary gland tumors	[64] [65]
β-1,4-Galactosyltransferase (b4Gal-T1-7)	B4GALT1-7	Breast cancer Colon cancer Liver cancer Leukemia Lung cancer Neuroblastoma Renal carcinoma	[66] [67] [68] [69] [70] [71] [72]
β-1,3-Glucuronyltransferase3 (GlcAT-I)	B3GAT3	Liver cancer	[73]
Exostosin like glycosyltransferase (EXTL1-3)	EXTL1-3	Breast cancer Hepatocarcinoma	[55] [74]
Exostosin1/2 (EXT1/2)	EXT1-2	Breast cancer Chondrosarcoma Osteochondroma Hepatocarcinoma Glioma Leukemia Thyroid tumor	[54,55] [75,76] [53,75,76] [77] [52] [57,58] [56]
N-deacetylase/N-sulfotransferase (1-4) (NDST1-4)	NDST1-4	Colorectal cancer Melanoma	[31,46] [78]
Glucuronyl C5-epimerase (GLCE)	GLCE	Breast cancer Lung cancer Prostate cancer	[59,60] [61] [62,63]
Hexuronyl 2-O-sulfotransferase (2-OST)	HS2ST	Breast cancer Multiple myeloma Prostate cancer	[79] [30] [49]
Glucosaminyl 6-O-sulfotransferase (6-OST)	HS6ST	Colorectal cancer Gastric cancer Glioma Ovarian cancer Pancreatic cancer	[50] [51] [52] [80,81] [82]
Glucosaminyl 3-O-sulfotransferase (3-OST)	HS3ST	Breast cancer Chondrosarcoma Colorectal cancer Leukemia Lung cancer Pancreatic cancer	[83] [47,83] [84] [85] [48] [86]
Endo-6-O-sulfatase1/2 (SULF1/2)	SULF1-2	Breast cancer Cervical cancer Liver tumors Ovarian cancer Other cancers	[87] [88] [89] [87] [90,91]
Heparanase (HPSE1/2)	HPSE1-2	Bladder cancer Brain tumors Breast cancer Gastric cancer Head and neck cancers Hepatocarcinoma Mesothelioma Myeloma Ovarian cancer Pancreatic cancer Sarcoma	[92] [93] [94,95] [96] [97] [98] [99] [100,101] [102] [103] [104]