KEGG   ENZYME: 5.1.3.17
Entry
EC 5.1.3.17                 Enzyme                                 
Name
heparosan-N-sulfate-glucuronate 5-epimerase;
heparosan epimerase;
heparosan-N-sulfate-D-glucuronosyl 5-epimerase;
C-5 uronosyl epimerase;
polyglucuronate epimerase;
D-glucuronyl C-5 epimerase;
poly[(1,4)-beta-D-glucuronosyl-(1,4)-N-sulfo-alpha-D-glucosaminyl] glucurono-5-epimerase
Class
Isomerases;
Racemases and epimerases;
Acting on carbohydrates and derivatives
Sysname
poly[(1->4)-beta-D-glucuronosyl-(1->4)-N-sulfo-alpha-D-glucosaminyl] glucurono-5-epimerase
Reaction(IUBMB)
Epimerization of D-glucuronate in heparosan-N-sulfate to L-iduronate. [RN:R04389]
Reaction(KEGG)
R04389
Comment
The enzyme acts on D-glucosyluronate residues in N-sulfated heparosan polymers, converting them to L-iduronate, thus modifying the polymer to heparan-N-sulfate. The enzyme requires that at least the N-acetylglucosamine residue linked to C-4 of the substrate has been deacetylated and N-sulfated, and activity is highest with fully N-sulfated substrate. It does not act on glucuronate residues that are O-sulfated or are adjacent to N-acetylglucosamine residues that are O-sulfated at the 6 position. Thus the epimerization from D-glucuronate to L-iduronate occurs after N-sulfation of glucosamine residues but before O-sulfation. Not identical with EC 5.1.3.19 chondroitin-glucuronate 5-epimerase or with EC 5.1.3.36, heparosan-glucuronate 5-epimerase.
History
EC 5.1.3.17 created 1984, modified 2015
Pathway
ec00534  Glycosaminoglycan biosynthesis - heparan sulfate / heparin
ec01100  Metabolic pathways
Orthology
K01793  heparosan-N-sulfate-glucuronate 5-epimerase
Genes
HSA26035(GLCE)
PTR453542(GLCE)
PPS100975106(GLCE)
GGO101148037(GLCE)
PON100457760(GLCE)
NLE100592444(GLCE)
HMH116482598(GLCE)
MCC694999(GLCE)
MCF102130661(GLCE)
MTHB126958145
MNI105487892(GLCE)
CSAB103245410(GLCE)
CATY105600958(GLCE)
PANU101025031(GLCE)
TGE112627967(GLCE)
MLEU105531073(GLCE)
RRO104662604(GLCE)
RBB108514748(GLCE)
TFN117070431(GLCE)
PTEH111548375(GLCE)
CANG105510505(GLCE)
CJC100402503(GLCE)
SBQ101034113(GLCE)
CIMI108303038(GLCE)
CSYR103273821(GLCE)
MMUR105873420(GLCE)
LCAT123623484(GLCE)
PCOQ105826834(GLCE)
OGA100966636(GLCE)
MMU93683(Glce)
MCAL110301842(Glce)
MPAH110327461(Glce)
RNO363073(Glce)
MCOC116073962(Glce)
ANU117699057(Glce)
MUN110550219(Glce)
CGE100754874(Glce)
MAUA101836212(Glce)
PROB127231855(Glce)
PLEU114691670(Glce)
MORG121466157(Glce)
MFOT126507456
AAMP119810193(Glce)
NGI103736825(Glce)
HGL101703064(Glce)
CPOC100733856(Glce)
CCAN109698168(Glce)
DORD105982523(Glce)
DSP122122999(Glce)
PLOP125339053(Glce)
NCAR124977563
MMMA107137979(Glce)
OCU100341828
OPI101529827(GLCE)
TUP102502937(GLCE)
GVR103592951(GLCE)
CFA487622(GLCE)
CLUD112675683(GLCE)
VVP112931528(GLCE)
VLG121484132(GLCE)
NPO129506383(GLCE)
AML100478227(GLCE)
UMR103679039(GLCE)
UAH113240917(GLCE)
UAR123800773(GLCE)
ELK111153319
LLV125104837
MPUF101670150(GLCE)
MNP131999565(GLCE)
MLK131835990(GLCE)
NVS122894189(GLCE)
ORO101386831(GLCE)
EJU114206527(GLCE)
ZCA113909686(GLCE)
MLX118003606(GLCE)
NSU110583927(GLCE)
LWW102734809(GLCE)
FCA101095983(GLCE)
PYU121041910(GLCE)
PCOO112856243(GLCE)
PBG122467608(GLCE)
PVIV125167537(GLCE)
LRUF124508447
PTG102952073(GLCE)
PPAD109262774(GLCE)
PUC125910099
AJU106980561
HHV120221934(GLCE)
BTA281195(GLCE)
BOM102274550(GLCE)
BIU109564721(GLCE)
BBUB102405908(GLCE)
BBIS104984651(GLCE)
CHX102172900(GLCE)
OAS101106592(GLCE)
BTAX128054475(GLCE)
ODA120852503 120860868
CCAD122443089(GLCE)
MBEZ129562362(GLCE)
SSC100154444(GLCE)
CFR102504983(GLCE)
CBAI105069233(GLCE)
CDK105087308(GLCE)
VPC102534038(GLCE)
BACU103008441(GLCE)
BMUS118890275(GLCE)
LVE103080354(GLCE)
OOR101271197(GLCE)
DLE111177767(GLCE)
PCAD102990323(GLCE)
PSIU116749662(GLCE)
NASI112401341(GLCE)
ECB100052581(GLCE)
EPZ103543875(GLCE)
EAI106835661(GLCE)
MYB102262970(GLCE)
MYD102752823(GLCE)
MMYO118654570(GLCE)
MLF102423949(GLCE)
PKL118718777(GLCE)
EFUS103283770(GLCE)
MNA107525844(GLCE)
DRO112311999(GLCE)
SHON118992362(GLCE)
AJM119056987(GLCE)
PDIC114491549(GLCE)
PHAS123808190(GLCE)
MMF118617890(GLCE)
PPAM129064140(GLCE)
HAI109385253(GLCE)
RFQ117023538(GLCE)
PALE102891481(GLCE)
PGIG120604461(GLCE)
PVP105309629(GLCE)
RAY107511425(GLCE)
MJV108397133(GLCE)
TOD119238797(GLCE)
SARA101558339(GLCE)
LAV100673589(GLCE)
TMU101347909
ETF101654749(GLCE)
DNM101444954(GLCE)
MDO100019166(GLCE)
GAS123236126(GLCE)
SHR100925822(GLCE)
AFZ127551219
PCW110205598(GLCE)
OAA100085653(GLCE)
GGA415566(GLCE)
PCOC116225491(GLCE)
MGP100546915(GLCE)
CJO107318962(GLCE)
TPAI128075277(GLCE)
LMUT125698146(GLCE)
NMEL110403606(GLCE)
APLA101789857(GLCE)
ACYG106043673(GLCE)
CATA118259695(GLCE)
AFUL116493554(GLCE)
LSR110477573(GLCE)
SCAN103816259(GLCE)
PMOA120508939(GLCE)
OTC121343512(GLCE)
PRUF121354536(GLCE)
GFR102034065(GLCE)
FAB101818992(GLCE)
OMA130257432(GLCE)
PHI102113112(GLCE)
PMAJ107209523(GLCE)
CCAE111933852(GLCE)
CBRC103619816(GLCE)
ETL114070300(GLCE)
ZAB106630389(GLCE)
ACHL103805831(GLCE)
SVG106860151(GLCE)
MMEA130582553(GLCE)
HRT120758897(GLCE)
FPG101915406(GLCE)
FCH102054462(GLCE)
CLV102093041(GLCE)
EGZ104133817(GLCE)
NNI104017037(GLCE)
PCRI104024222(GLCE)
PLET104613898(GLCE)
EHS104512065(GLCE)
PCAO104041675(GLCE)
ACUN113484044(GLCE)
TALA104362314(GLCE)
PADL103914742(GLCE)
AFOR103899842(GLCE)
ACHC115341823(GLCE)
HALD104325661(GLCE)
HLE104834629(GLCE)
AGEN126043592
GCL127020407
CCRI104161014(GLCE)
CSTI104559768(GLCE)
CMAC104473275(GLCE)
MUI104539098(GLCE)
BREG104641025(GLCE)
FGA104081776(GLCE)
GSTE104259531(GLCE)
LDI104340123
MNB103774713(GLCE)
OHA104332008(GLCE)
NNT104411237(GLCE)
SHAB115612813(GLCE)
DPUB104303260(GLCE)
PGUU104469611(GLCE)
ACAR104521144(GLCE)
CPEA104395717(GLCE)
AVIT104270030(GLCE)
CVF104292673(GLCE)
RTD128914908(GLCE)
CUCA104065176(GLCE)
TEO104384146(GLCE)
BRHI104496092(GLCE)
AAM106495389
AROW112963593(GLCE)
NPD112948346(GLCE)
TGT104570165(GLCE)
DNE112989520(GLCE)
SCAM104139215(GLCE)
ASN102386367(GLCE)
AMJ102568999(GLCE)
CPOO109309218(GLCE)
GGN109288409(GLCE)
PSS102453691(GLCE)
CMY114019430(GLCE)
CCAY125643746(GLCE)
DCC119862903(GLCE)
CPIC101943859(GLCE)
TST117883528(GLCE)
CABI116815733(GLCE)
MRV120373550(GLCE)
ACS100556549(glce)
ASAO132762428(GLCE)
PVT110086904(GLCE)
SUND121933228(GLCE)
PBI103064301(GLCE)
PMUR107283920(GLCE)
CTIG120320136(GLCE)
TSR106540160(GLCE)
PGUT117676200(GLCE)
APRI131184689(GLCE)
PTEX113442084(GLCE)
NSS113425677(GLCE)
VKO123030701(GLCE)
PMUA114584097(GLCE)
PRAF128401359(GLCE)
ZVI118092529(GLCE)
HCG128334976(GLCE)
GJA107116570(GLCE)
STOW125424828(GLCE)
EMC129345620(GLCE)
XLA108712794(glce.S) 432242(glce.L)
XTR100489303(glce)
NPR108796035(GLCE)
RTEM120923836
BBUF120986043(GLCE)
BGAR122927478(GLCE)
MUO115459956(GLCE)
GSH117348222(GLCE)
DRE100007670(glceb) 405776(glcea)
SRX107720792 107740438 107740877 107755093
SANH107663974 107670231 107673273 107700477
SGH107553083 107557703 107559985 107569710
CCAR109050896 109066693 109093126 109109568
CAUA113043709 113052345 113079266 113106000
CGIB127962113 128014596 128016907
PTET122330363(glcea) 122348299 122349241(glceb)
LROH127156392(glcea) 127168418(glceb)
OMC131534070(glcea) 131544352(glceb)
PPRM120475357(glcea) 120491029(glceb)
RKG130073707(glcea) 130082981(glceb)
MAMB125246813(glcea) 125264777(glceb)
CIDE127507476 127515223
MASI127416399 127436102 127437561 127443782
TROS130551922(glcea) 130557793(glceb)
TDW130414305(glcea) 130427134(glceb)
MANU129433850(glcea) 129443782(glceb)
IPU108259276(glceb) 108274589(glcea)
IFU128602706(glceb) 128617895(glcea)
PHYP113529971 113540832
SMEO124379867(glceb) 124391984(glcea)
TFD113636763(glceb) 113653913(glcea)
TVC132838729(glceb) 132856931(glcea)
AMEX103040160 103040217
CMAO118811890(glcea) 118817488(glceb)
EEE113572322 113584499
CHAR105892465(glcea) 105906560(glceb)
TRU101074194(glce)
TFS130517173(glceb)
TNGGSTEN00018120G001
LCO104920418(glce)
NCC104944436(glce)
TBEN117471173(glceb)
CGOB115005637(glce)
PGEO117464774(glceb)
GACU117549342(glceb)
EMAC134863718(glceb)
ELY117253633(glceb)
EFO125899220(glceb)
PLEP121946907(glceb)
SLUC116043452(glceb)
ECRA117950179(glceb)
ESP116689425(glce)
PFLV114556531(glce)
GAT120812554(glceb)
PPUG119198574(glceb)
AFB129108435(glceb)
CLUM117728632(glceb)
PSWI130193270(glceb)
MSAM119890080(glceb)
SCHU122861321(glceb)
CUD121507882(glceb)
ALAT119018323(glceb)
MZE101468415(glce)
ONL100689967(glce)
OAU116323950(glceb)
OLA101168104(glce)
OML112160751(glceb)
CSAI133459566(glceb)
XMA102220367(glce)
XCO114142896(glce)
XHE116718446(glce)
PRET103462118(glce)
PFOR103146186(glce)
PLAI106964982(glce)
PMEI106930363(glce)
GAF122841808(glceb)
PPRL129369432(glceb)
CVG107104897(glce)
CTUL119776853(glceb)
GMU124867026(glceb)
NFU107381748(glce)
KMR108234798(glceb)
ALIM106512969(glce)
NWH119424012(glceb)
AOCE111564420(glce)
MCEP125005684(glceb)
CSEM103398539(glce)
POV109637279(glce)
SSEN122772339(glceb)
HHIP117756084(glceb)
HSP118109205(glceb)
SMAU118302159(glceb)
LCF108887864(glceb)
SDU111223714(glce)
SLAL111670127(glce)
XGL120797172(glceb)
HCQ109514358(glce)
SSCV125967138
SBIA133498478
PEE133399155(glceb)
PTAO133471157(glceb)
BPEC110174517(glce)
MALB109963362(glce)
BSPL114852278(glceb)
SJO128362782(glceb)
SASA106562601 106562708 106593990(glce)
STRU115166801 115203527
OTW112264250 121845510
OMY110506198 110526997
OGO124035485(glceb)
OKI109879929(glce)
OKE118372181
SALP111980622(glce)
SNH120049768(glceb)
CCLU121548171(glceb)
ELS105013975(glce)
SFM108934585(glce)
PKI111845625(glce)
AANG118215235
LOC102694794(glce)
PSPA121294489 121299204
ARUT117427879 117429202
PSEX120541503
LCM102357523(GLCE)
CMK103190379
CPLA122540910
HOC132834294
LERI129712750(glceb)
PMRN116941988(GLCE)
LRJ133343759(GLCE)
BFO118429968
BBEL109471711
CIN100181100 113473995
SCLV120335536
SPU115918110
LPIC129275791
APLC110979098
AJC117109373
SKO100370244
DMEDmel_CG3194(Hsepi)
DER6541647
DSE6607870
DSIDsimw501_GD10315(Dsim_GD10315)
DYADyak_GE24632
DAN6496639
DSR110189521
DPO4803422
DPE6591115
DMN108159497 117190075
DWI6652210
DGR6559652
DMODmoj_GI20241
DAZ108616232
DNV108654278
DHE111597636
DVI6626477
CCAT101459521
BOD106626692
BDR105230624
RZE108353930
AOQ129244787
TDA119680919
MDE101895364
SCAC106091743
LCQ111679193
LSQ119606790 119614316
GFS119642228
ECOE129950263
CLON129607435
HIS119649614
AGA1276368
ACOZ120949916
AARA120894259
AMER121592948
ASTE118510552
AFUN125772131
AMOU128303949
AALI118462703
AAG5565650
AALB109397352 109420263
CQUCpipJ_CPIJ014213
CPII120415752
CNS116352573
BCOO119081361
AME410118
ACER107994528
ALAB122719649
ADR102671555
AFLR100867340
BIM100746913
BBIF117211829
BVK117231548
BVAN117166639
BTER100648289
BAFF126917459
BPYO122572307
BPAS132906620
FVI122537017
CCAL108625516
OBB114873294
OLG117601985
MGEN117225220
NMEA116425382
CGIG122397182
SOC105205016
MPHA105839326
AEC105152283
ACEP105621248
PBAR105428344
VEM105569445
HST105180547
DQU106743638
CFO105252887
FEX115245702
LHU105667459
PGC109856517
OBO105275234
PCF106785179
PFUC122525806
VPS122633444
VCRB124429623
VVE124953901
NVI100123867
CSOL105359880
TPRE106654805
LHT122511160
LBD127287345
MDL103572179
CGLO123265683
FAS105271878
DAM107037394
AGIF122848386
CINS118068612
VCAN122409058
CCIN107269276
DSM124404319
NPT124213295
NFB124176075
NLO107221644
NVG124298264
AROA105690027
TCA663342
DPA109539260
SOY115890532
AGRG126746177
ATD109604348
CSET123314239
AGB108914173
LDC111501961
NVL108565207
APLN108733612
PPYR116175551
OTU111415151
BMOR101736603
BMAN114251952
MSEX115451466
DPLKGM_205946
BANY112052728
MJU123881054
NIQ126770001
VCD124533535
MCIX123654105
PMAC106718979
PPOT106105194
PXU106128471
PRAP110999574
PBX123708233
PNAP125049785
ZCE119835154
CCRC123690903
LSIN126965641
HAW110374867
HZE124631208
TNL113504444
SLIU111351881
OFU114351332
PXY105382951
PGW126367326
CFEL113363902 113377896
CCRN123295251
API100164972
DNX107166129
AGS114124434
RMD113555611 113556575
ACOO126844264
DVT126897373
BTAB109035995
DCI103509657
CLEC106670169
HHAL106680682
NLU111045855
HVI124366335
MQU129002806
PHUPhum_PHUM045460
FOC113205627
TPAL117649391
ZNE110840150
CSEC111862967
SGRE126356007
BROR134536896
IEL124158479
FCD110862144
DPXDAPPUDRAFT_325820
DMK116919414
DPZ124341667
PVM113806085
PJA122266525
PCHN125032807
PMOO119595439
HAME121861192
PCLA123762054
CQD128695462
PTRU123512008
ESN126997011
HAZT108678715
EAF111700314
LSM121117465
TCF131882839
PPOI119109679
DSV119446560
RSAN119389727
RMP119177059
VDE111253483
VJA111264172
TUT107372174
DPTE113793709
DFR124494060
CSCU111642384
PTEP107446424
ABRU129960790
UDV129217698
SDM118202796
LPOL106459105 106467792 106477692
PMEO129583932
CELCELE_B0285.5(hse-5)
CBRCBG_18086(Cbr-hse-5)
CRQGCK72_010119
NAINECAME_11020
BMYBM_BM1737(Bma-hse-5)
LOALOAG_05488
TSPTsp_05727
HROHELRODRAFT_142574
LGILOTGIDRAFT_123168
PVUL126826994
PCAN112556648
BGT106071274 106078711
GAE121390640
HRF124149284
HRJ124280958
CRG105324867 105329756
CVN111135541 111135825
CANU128188555 128188865
OED125680051 125680093
MYI110455274
PMAX117344629
MCAF127720411
MMER123526242
RPHI132748127
DPOL127867599 127867600
OBI106876518
OSN115227227
LAK106161783 106167628 106169258 106173383
SMMSmp_126320
SHXMS3_00002945(HSE5)
OVIT265_06313
EGLEGR_02731
NVE5520075
EPA110241410
ATEN116299102
ADF107346476
AMIL114975962
PDAM113672224
SPIS111329368
DGT114518014
XEN124444471
HMG100207938
HSY130645104
TADTRIADDRAFT_27335
PTTVY86_07540
BMARHF888_03430
KSDKS2013_1451
CHACHAB381_1476
AANAAANAER_2044
NAMNAMH_1657
NAPC3L23_08545
GSBGSUB_15310
VNECFK40_05110
LAOAOX59_11990
AQTFN924_03240
CKLCKL_1010 CKL_1095
CKRCKR_0914 CKR_0997
CLJCLJU_c25590 CLJU_c26810
CAHCAETHG_0628 CAETHG_0762
CSQCSCA_4800
CDRKB9W14_20875
RCHRUM_09550
DMTDESME_14470
BPROPMF13cell1_00168
TTETTE0659
NOWGBF35_13825
HHGXM38_017590
RULUC8_39190
ABAAcid345_3516
AMOAnamo_1192
BUYD8S85_14480
BVIRI6J59_01675
FUVJR347_05040
MYRMYRA21_1118
MPWMPR_3643
MODAS202_05235
FLUCHH17_10220
IALIALB_2082
MSJMSSAC_1842
MSWMSSIT_2435
 » show all
Reference
1  [PMID:107165]
  Authors
Backstrom G, Hook M, Lindahl U, Feingold DS, Malmstrom A, Roden L, Jacobsson I.
  Title
Biosynthesis of heparin. Assay and properties of the microsomal uronosyl C-5 epimerase.
  Journal
J Biol Chem 254:2975-82 (1979)
Reference
2  [PMID:6420398]
  Authors
Jacobsson I, Lindahl U, Jensen JW, Roden L, Prihar H, Feingold DS
  Title
Biosynthesis of heparin. Substrate specificity of heparosan N-sulfate D-glucuronosyl 5-epimerase.
  Journal
J Biol Chem 259:1056-63 (1984)
Reference
3  [PMID:10642607]
  Authors
Hagner-McWhirter A, Hannesson HH, Campbell P, Westley J, Roden L, Lindahl U, Li JP
  Title
Biosynthesis of heparin/heparan sulfate: kinetic studies of the glucuronyl C5-epimerase with N-sulfated derivatives of the Escherichia coli K5 capsular polysaccharide as substrates.
  Journal
Glycobiology 10:159-71 (2000)
DOI:10.1093/glycob/10.2.159
Other DBs
ExplorEnz - The Enzyme Database: 5.1.3.17
IUBMB Enzyme Nomenclature: 5.1.3.17
ExPASy - ENZYME nomenclature database: 5.1.3.17
BRENDA, the Enzyme Database: 5.1.3.17
CAS: 112567-86-9

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