Activities in Family	Glucuronyl-galactosyl-proteoglycan 4-α-N-acetylglucosaminyltransferase (EC 2.4.1.223); Glucuronosyl-N-acetylglucosaminyl-proteoglycan α-1,4-N-acetylglucosaminyltransferase (EC 2.4.1.224);
Mechanism	Retaining
3D Structure Status	GT-A
Note	The long animal heparan synthases are made of two domains. The N-terminal domain, which adds b-1,4-GlcA residues, belongs to family GT47 while the C-terminal domain, which adds a-1,4-GlcNAc residues, belongs to family GT64. The plant homologs have a single domain.
External resources	Glymap;
Statistics	GenBank accession (724); Uniprot accession (31); PDB accession (20); 3D entries (4); cryst (0)

Eukaryota
Protein Name	EC#	Organism	GenBank	Uniprot	PDB/3D Carbohydrate Ligands Resolution (Å)
UDP-GlcUA:β-glucuronyltransferase / UDP-GlcNAc: α-N-acetylglucosaminyltransferase (exostosin-2;Ext2)	2.4.1.224	Homo sapiens	AAB07008.1 AAB62718.1 AAC50764.1 AAC51219.1 AAH10058.1 AAH68545.1 NP_000392.1	B2R5Z6 Q6NUL1 Q93063	7SCH[B] 3.10 7SCJ[B] α-D-GlcpNAc-(1-4)-β-D-GlcpA-(1-4)-α-D-GlcpNAc 3.40 7SCK[B] β-D-GlcpA-(1-4)-α-D-GlcpNAc-(1-4)-β-D-GlcpA-(1-4)-α-D-GlcpNAc 2.80 7UQX[B] 3.30 7UQY[B] 3.00 7ZAY[B] 2.80
UDP-GlcUA:β-glucuronyltransferase / UDP-GlcNAc: α-N-acetylglucosaminyltransferase (exostosin-1;Ext1)	2.4.1.224	Homo sapiens	AAB62283.1 AAH01174.1 ACS08204.1 AGV77097.1 AHB20150.1 BAG35949.1 NP_000118.1	B2R7V2 Q16394	7SCH[A] 3.10 7SCJ[A] α-D-GlcpNAc-(1-4)-β-D-GlcpA-(1-4)-α-D-GlcpNAc 3.40 7SCK[A] β-D-GlcpA-(1-4)-α-D-GlcpNAc-(1-4)-β-D-GlcpA-(1-4)-α-D-GlcpNAc 2.80 7UQX[A] 3.30 7UQY[A] 3.00 7ZAY[A] 2.80
exostosin-like 3 (Extl3;Extr1)	2.4.1.223	Homo sapiens	AAC39598.1 AAB67602.1 AAD01877.1 AAD42041.1 AAH06363.1 AAP36017.1 AAW12588.1 AAW12594.1 ACW32787.1 ADO22448.1 BAA24080.1 BAA25445.2 BAA25445.3 BAG57702.1 NP_001431.1	A0A384NPY9 D3DST8 O43909 Q53XT3	7AU2[A,B] 2.43 7AUA[A,B] 2.93 8OG1[A] 1.58 8OG4[A,B] 2.10
UDP-GlcNAc: α-1,4-N-acetylglucosaminyltransferase (Extl2)	2.4.1.223	Mus musculus	AAG17542.1 AAH31438.1 AAH94444.1 BAB31683.2 BAC26062.1 BAC27649.1 BAE34959.1 BAE36615.1 BAM48568.1 BAM48569.1 NP_067363.1	K0J1C6 Q3TSR0 Q3TXF7 Q8C089 Q8C197 Q9ES89	1OMX[A,B] 2.40 1OMZ[A,B] 2.10 1ON6[A,B] 2.30 1ON8[A,B] β-D-GlcpA-(1-3)-β-D-Galp 2.70
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