GLI3_HUMAN - dbPTM
GLI3_HUMAN - PTM Information in dbPTM
Basic Information of Protein
UniProt ID GLI3_HUMAN
UniProt AC P10071
Protein Name Transcriptional activator GLI3
Gene Name GLI3
Organism Homo sapiens (Human).
Sequence Length 1580
Subcellular Localization Nucleus. Cytoplasm. Cell projection, cilium. GLI3FL is localized predominantly in the cytoplasm while GLI3R resides mainly in the nucleus. Ciliary accumulation requires the presence of KIF7 and SMO. Translocation to the nucleus is promoted by interac
Protein Description Has a dual function as a transcriptional activator and a repressor of the sonic hedgehog (Shh) pathway, and plays a role in limb development. The full-length GLI3 form (GLI3FL) after phosphorylation and nuclear translocation, acts as an activator (GLI3A) while GLI3R, its C-terminally truncated form, acts as a repressor. A proper balance between the GLI3 activator and the repressor GLI3R, rather than the repressor gradient itself or the activator/repressor ratio gradient, specifies limb digit number and identity. In concert with TRPS1, plays a role in regulating the size of the zone of distal chondrocytes, in restricting the zone of PTHLH expression in distal cells and in activating chondrocyte proliferation. Binds to the minimal GLI-consensus sequence 5'-GGGTGGTC-3'..
Protein Sequence MEAQSHSSTTTEKKKVENSIVKCSTRTDVSEKAVASSTTSNEDESPGQTYHRERRNAITMQPQNVQGLSKVSEEPSTSSDERASLIKKEIHGSLPHVAEPSVPYRGTVFAMDPRNGYMEPHYHPPHLFPAFHPPVPIDARHHEGRYHYDPSPIPPLHMTSALSSSPTYPDLPFIRISPHRNPTAASESPFSPPHPYINPYMDYIRSLHSSPSLSMISATRGLSPTDAPHAGVSPAEYYHQMALLTGQRSPYADIIPSAATAGTGAIHMEYLHAMDSTRFSSPRLSARPSRKRTLSISPLSDHSFDLQTMIRTSPNSLVTILNNSRSSSSASGSYGHLSASAISPALSFTYSSAPVSLHMHQQILSRQQSLGSAFGHSPPLIHPAPTFPTQRPIPGIPTVLNPVQVSSGPSESSQNKPTSESAVSSTGDPMHNKRSKIKPDEDLPSPGARGQQEQPEGTTLVKEEGDKDESKQEPEVIYETNCHWEGCAREFDTQEQLVHHINNDHIHGEKKEFVCRWLDCSREQKPFKAQYMLVVHMRRHTGEKPHKCTFEGCTKAYSRLENLKTHLRSHTGEKPYVCEHEGCNKAFSNASDRAKHQNRTHSNEKPYVCKIPGCTKRYTDPSSLRKHVKTVHGPEAHVTKKQRGDIHPRPPPPRDSGSHSQSRSPGRPTQGALGEQQDLSNTTSKREECLQVKTVKAEKPMTSQPSPGGQSSCSSQQSPISNYSNSGLELPLTDGGSIGDLSAIDETPIMDSTISTATTALALQARRNPAGTKWMEHVKLERLKQVNGMFPRLNPILPPKAPAVSPLIGNGTQSNNTCSLGGPMTLLPGRSDLSGVDVTMLNMLNRRDSSASTISSAYLSSRRSSGISPCFSSRRSSEASQAEGRPQNVSVADSYDPISTDASRRSSEASQSDGLPSLLSLTPAQQYRLKAKYAAATGGPPPTPLPNMERMSLKTRLALLGDALEPGVALPPVHAPRRCSDGGAHGYGRRHLQPHDAPGHGVRRASDPVRTGSEGLALPRVPRFSSLSSCNPPAMATSAEKRSLVLQNYTRPEGGQSRNFHSSPCPPSITENVTLESLTMDADANLNDEDFLPDDVVQYLNSQNQAGYEQHFPSALPDDSKVPHGPGDFDAPGLPDSHAGQQFHALEQPCPEGSKTDLPIQWNEVSSGSADLSSSKLKCGPRPAVPQTRAFGFCNGMVVHPQNPLRSGPAGGYQTLGENSNPYGGPEHLMLHNSPGSGTSGNAFHEQPCKAPQYGNCLNRQPVAPGALDGACGAGIQASKLKSTPMQGSGGQLNFGLPVAPNESAGSMVNGMQNQDPVGQGYLAHQLLGDSMQHPGAGRPGQQMLGQISATSHINIYQGPESCLPGAHGMGSQPSSLAVVRGYQPCASFGGSRRQAMPRDSLALQSGQLSDTSQTCRVNGIKMEMKGQPHPLCSNLQNYSGQFYDQTVGFSQQDTKAGSFSISDASCLLQGTSAKNSELLSPGANQVTSTVDSLDSHDLEGVQIDFDAIIDDGDHSSLMSGALSPSIIQNLSHSSSRLTTPRASLPFPALSMSTTNMAIGDMSSLLTSLAEESKFLAVMQ
Overview of Protein Modification Sites with Functional and Structural Information
Experimental Post-Translational Modification Sites

* ASA = Accessible Surface Area

Locations Modification Substrate Peptides
&
Secondary Structure
ASA (%) Reference Orthologous
Protein Cluster
1Acetylation-------MEAQSHSS
-------CCCCCCCC
42.1719413330
19PhosphorylationEKKKVENSIVKCSTR
CHHHHHCCEEEECCC
18.4322817900
24PhosphorylationENSIVKCSTRTDVSE
HCCEEEECCCCCCCH
18.4030177828
25PhosphorylationNSIVKCSTRTDVSEK
CCEEEECCCCCCCHH
47.2430576142
27PhosphorylationIVKCSTRTDVSEKAV
EEEECCCCCCCHHHH
40.6230177828
30PhosphorylationCSTRTDVSEKAVASS
ECCCCCCCHHHHHCC
35.8730177828
32AcetylationTRTDVSEKAVASSTT
CCCCCCHHHHHCCCC
40.4323236377
37PhosphorylationSEKAVASSTTSNEDE
CHHHHHCCCCCCCCC
26.4327251275
38PhosphorylationEKAVASSTTSNEDES
HHHHHCCCCCCCCCC
31.5727251275
39PhosphorylationKAVASSTTSNEDESP
HHHHCCCCCCCCCCC
31.3527251275
40PhosphorylationAVASSTTSNEDESPG
HHHCCCCCCCCCCCC
38.0128555341
45PhosphorylationTTSNEDESPGQTYHR
CCCCCCCCCCCCCCH
46.8025159151
50PhosphorylationDESPGQTYHRERRNA
CCCCCCCCCHHHHHC
7.3230576142
59PhosphorylationRERRNAITMQPQNVQ
HHHHHCCCCCCCCCC
13.8222210691
69PhosphorylationPQNVQGLSKVSEEPS
CCCCCCCHHCCCCCC
37.5222210691
78O-linked_GlycosylationVSEEPSTSSDERASL
CCCCCCCCHHHHHHH
39.8828411811
84PhosphorylationTSSDERASLIKKEIH
CCHHHHHHHHHHHHH
36.5324719451
87SumoylationDERASLIKKEIHGSL
HHHHHHHHHHHHCCC
50.04-
93PhosphorylationIKKEIHGSLPHVAEP
HHHHHHCCCCCCCCC
25.2728348404
101PhosphorylationLPHVAEPSVPYRGTV
CCCCCCCCCCCCCEE
27.51-
104PhosphorylationVAEPSVPYRGTVFAM
CCCCCCCCCCEEEEE
21.37-
163PhosphorylationLHMTSALSSSPTYPD
CCCCCCCCCCCCCCC
28.61-
164PhosphorylationHMTSALSSSPTYPDL
CCCCCCCCCCCCCCC
40.9824719451
165PhosphorylationMTSALSSSPTYPDLP
CCCCCCCCCCCCCCC
20.2028348404
167PhosphorylationSALSSSPTYPDLPFI
CCCCCCCCCCCCCCE
50.0328348404
168PhosphorylationALSSSPTYPDLPFIR
CCCCCCCCCCCCCEE
9.72-
175MethylationYPDLPFIRISPHRNP
CCCCCCEEECCCCCC
24.58-
183PhosphorylationISPHRNPTAASESPF
ECCCCCCCCCCCCCC
38.1029978859
186PhosphorylationHRNPTAASESPFSPP
CCCCCCCCCCCCCCC
35.8229978859
188PhosphorylationNPTAASESPFSPPHP
CCCCCCCCCCCCCCC
28.8429978859
191PhosphorylationAASESPFSPPHPYIN
CCCCCCCCCCCCCCC
40.6929978859
196PhosphorylationPFSPPHPYINPYMDY
CCCCCCCCCCHHHHH
15.4229978859
200PhosphorylationPHPYINPYMDYIRSL
CCCCCCHHHHHHHHH
9.7229978859
203PhosphorylationYINPYMDYIRSLHSS
CCCHHHHHHHHHHCC
5.0229978859
220MethylationLSMISATRGLSPTDA
HHHHEECCCCCCCCC
43.71-
223PhosphorylationISATRGLSPTDAPHA
HEECCCCCCCCCCCC
28.6927251275
257PhosphorylationPYADIIPSAATAGTG
CCHHCCCCCCCCCCC
21.81-
293PhosphorylationARPSRKRTLSISPLS
CCCCCCCEEEECCCC
28.0829523821
295PhosphorylationPSRKRTLSISPLSDH
CCCCCEEEECCCCCC
22.0229523821
297PhosphorylationRKRTLSISPLSDHSF
CCCEEEECCCCCCCC
19.2029523821
312PhosphorylationDLQTMIRTSPNSLVT
CHHHHEECCCCCEEE
37.3024719451
313PhosphorylationLQTMIRTSPNSLVTI
HHHHEECCCCCEEEE
16.5528348404
418PhosphorylationESSQNKPTSESAVSS
CCCCCCCCCHHHCCC
46.8930206219
419PhosphorylationSSQNKPTSESAVSST
CCCCCCCCHHHCCCC
37.7730206219
421PhosphorylationQNKPTSESAVSSTGD
CCCCCCHHHCCCCCC
33.4630206219
424PhosphorylationPTSESAVSSTGDPMH
CCCHHHCCCCCCCCC
23.1530206219
425PhosphorylationTSESAVSSTGDPMHN
CCHHHCCCCCCCCCC
29.7930206219
426PhosphorylationSESAVSSTGDPMHNK
CHHHCCCCCCCCCCC
37.7930206219
435PhosphorylationDPMHNKRSKIKPDED
CCCCCCCCCCCCCCC
40.1030206219
436MethylationPMHNKRSKIKPDEDL
CCCCCCCCCCCCCCC
59.80-
438SumoylationHNKRSKIKPDEDLPS
CCCCCCCCCCCCCCC
50.5928112733
445PhosphorylationKPDEDLPSPGARGQQ
CCCCCCCCCCCCCCC
43.1729255136
462SumoylationPEGTTLVKEEGDKDE
CCCCEEEECCCCCCC
53.8428112733
541PhosphorylationVVHMRRHTGEKPHKC
EEEEHHCCCCCCCCC
45.2723532336
564UbiquitinationYSRLENLKTHLRSHT
HHHHHHHHHHHHHHC
45.54-
571PhosphorylationKTHLRSHTGEKPYVC
HHHHHHHCCCCCEEE
48.5629214152
595MethylationSNASDRAKHQNRTHS
CCHHHHHHHCCCCCC
46.76-
618PhosphorylationIPGCTKRYTDPSSLR
CCCCCCCCCCHHHHH
19.6329496907
619PhosphorylationPGCTKRYTDPSSLRK
CCCCCCCCCHHHHHH
45.53-
622PhosphorylationTKRYTDPSSLRKHVK
CCCCCCHHHHHHHHE
44.8321082442
623PhosphorylationKRYTDPSSLRKHVKT
CCCCCHHHHHHHHEE
37.2724719451
639PhosphorylationHGPEAHVTKKQRGDI
CCCCCCCCCCCCCCC
23.7421082442
640AcetylationGPEAHVTKKQRGDIH
CCCCCCCCCCCCCCC
45.917681179
640UbiquitinationGPEAHVTKKQRGDIH
CCCCCCCCCCCCCCC
45.91-
641AcetylationPEAHVTKKQRGDIHP
CCCCCCCCCCCCCCC
35.547681189
656PhosphorylationRPPPPRDSGSHSQSR
CCCCCCCCCCCCCCC
43.4023403867
658PhosphorylationPPPRDSGSHSQSRSP
CCCCCCCCCCCCCCC
24.7423403867
660PhosphorylationPRDSGSHSQSRSPGR
CCCCCCCCCCCCCCC
31.3923403867
662PhosphorylationDSGSHSQSRSPGRPT
CCCCCCCCCCCCCCC
37.4323403867
664PhosphorylationGSHSQSRSPGRPTQG
CCCCCCCCCCCCCCC
36.6330266825
669PhosphorylationSRSPGRPTQGALGEQ
CCCCCCCCCCCCCCC
37.9729255136
680PhosphorylationLGEQQDLSNTTSKRE
CCCCCCCCCCCCCHH
40.1823403867
682PhosphorylationEQQDLSNTTSKREEC
CCCCCCCCCCCHHHH
29.1323403867
683PhosphorylationQQDLSNTTSKREECL
CCCCCCCCCCHHHHE
36.1823927012
684PhosphorylationQDLSNTTSKREECLQ
CCCCCCCCCHHHHEE
27.9023927012
696SumoylationCLQVKTVKAEKPMTS
HEECEEEECCCCCCC
56.94-
773UbiquitinationRRNPAGTKWMEHVKL
HHCCCCCCHHHHHHH
43.5616705181
779SumoylationTKWMEHVKLERLKQV
CCHHHHHHHHHHHHH
46.6828112733
779UbiquitinationTKWMEHVKLERLKQV
CCHHHHHHHHHHHHH
46.6816705181
784UbiquitinationHVKLERLKQVNGMFP
HHHHHHHHHHCCCCC
59.4216705181
800UbiquitinationLNPILPPKAPAVSPL
CCCCCCCCCCCCCCC
66.1316705181
849PhosphorylationNMLNRRDSSASTISS
HCCCCCCCCCHHHHH
26.4530576142
850PhosphorylationMLNRRDSSASTISSA
CCCCCCCCCHHHHHH
30.4430576142
852PhosphorylationNRRDSSASTISSAYL
CCCCCCCHHHHHHHH
28.3730576142
853PhosphorylationRRDSSASTISSAYLS
CCCCCCHHHHHHHHH
25.2923312004
855PhosphorylationDSSASTISSAYLSSR
CCCCHHHHHHHHHHC
14.7030576142
856PhosphorylationSSASTISSAYLSSRR
CCCHHHHHHHHHHCC
19.6530576142
858PhosphorylationASTISSAYLSSRRSS
CHHHHHHHHHHCCCC
14.7423312004
860PhosphorylationTISSAYLSSRRSSGI
HHHHHHHHHCCCCCC
14.7730576142
861PhosphorylationISSAYLSSRRSSGIS
HHHHHHHHCCCCCCC
29.1723312004
864PhosphorylationAYLSSRRSSGISPCF
HHHHHCCCCCCCCCC
31.6323403867
865PhosphorylationYLSSRRSSGISPCFS
HHHHCCCCCCCCCCC
38.0323403867
868PhosphorylationSRRSSGISPCFSSRR
HCCCCCCCCCCCCCC
21.0923090842
872PhosphorylationSGISPCFSSRRSSEA
CCCCCCCCCCCCHHH
29.65-
873PhosphorylationGISPCFSSRRSSEAS
CCCCCCCCCCCHHHH
16.0923403867
876PhosphorylationPCFSSRRSSEASQAE
CCCCCCCCHHHHHCC
31.2727251275
877PhosphorylationCFSSRRSSEASQAEG
CCCCCCCHHHHHCCC
35.0610693759
880PhosphorylationSRRSSEASQAEGRPQ
CCCCHHHHHCCCCCC
26.3528348404
895PhosphorylationNVSVADSYDPISTDA
CCCCCCCCCCCCCCH
26.7027642862
899PhosphorylationADSYDPISTDASRRS
CCCCCCCCCCHHHHC
26.14-
900PhosphorylationDSYDPISTDASRRSS
CCCCCCCCCHHHHCC
35.98-
903PhosphorylationDPISTDASRRSSEAS
CCCCCCHHHHCCHHH
30.91-
906PhosphorylationSTDASRRSSEASQSD
CCCHHHHCCHHHHCC
31.2725850435
907PhosphorylationTDASRRSSEASQSDG
CCHHHHCCHHHHCCC
35.0625850435
910PhosphorylationSRRSSEASQSDGLPS
HHHCCHHHHCCCCCH
26.6025850435
912PhosphorylationRSSEASQSDGLPSLL
HCCHHHHCCCCCHHH
31.0223403867
917PhosphorylationSQSDGLPSLLSLTPA
HHCCCCCHHHHCCHH
47.1623090842
920PhosphorylationDGLPSLLSLTPAQQY
CCCCHHHHCCHHHHH
35.1723090842
933PhosphorylationQYRLKAKYAAATGGP
HHHHHHHHHHHHCCC
13.3128634298
937PhosphorylationKAKYAAATGGPPPTP
HHHHHHHHCCCCCCC
37.6328634298
943PhosphorylationATGGPPPTPLPNMER
HHCCCCCCCCCCHHH
43.1428634298
980PhosphorylationVHAPRRCSDGGAHGY
CCCCCCCCCCCCCCC
37.1626699800
987PhosphorylationSDGGAHGYGRRHLQP
CCCCCCCCCCCCCCC
9.5226699800
1006PhosphorylationGHGVRRASDPVRTGS
CCCCCCCCCCCCCCC
40.419302279
1013PhosphorylationSDPVRTGSEGLALPR
CCCCCCCCCCCCCCC
28.0722210691
1025PhosphorylationLPRVPRFSSLSSCNP
CCCCCCCCCCCCCCC
31.4822210691
1026PhosphorylationPRVPRFSSLSSCNPP
CCCCCCCCCCCCCCC
29.5628348404
1028PhosphorylationVPRFSSLSSCNPPAM
CCCCCCCCCCCCCHH
34.4928348404
1029PhosphorylationPRFSSLSSCNPPAMA
CCCCCCCCCCCCHHC
24.1728348404
1254PhosphorylationQPCKAPQYGNCLNRQ
CCCCCCCCCCCCCCC
14.83-
1401PhosphorylationRQAMPRDSLALQSGQ
CCCCCCCCEEHHCCC
19.47-
1412PhosphorylationQSGQLSDTSQTCRVN
HCCCCCCCCCEEEEC
21.25-
1413PhosphorylationSGQLSDTSQTCRVNG
CCCCCCCCCEEEECC
28.60-
1415PhosphorylationQLSDTSQTCRVNGIK
CCCCCCCEEEECCEE
11.31-
1422SumoylationTCRVNGIKMEMKGQP
EEEECCEEEEECCCC
30.15-
1444PhosphorylationQNYSGQFYDQTVGFS
HHCCCCCCCCCCCCC
10.1527642862
1459PhosphorylationQQDTKAGSFSISDAS
CCCCCCCEEECHHHH
22.2327732954
1461PhosphorylationDTKAGSFSISDASCL
CCCCCEEECHHHHHH
23.9227732954
1463PhosphorylationKAGSFSISDASCLLQ
CCCEEECHHHHHHHC
26.4027732954
1466PhosphorylationSFSISDASCLLQGTS
EEECHHHHHHHCCCC
15.4127732954
1539PhosphorylationSHSSSRLTTPRASLP
CCCCCCCCCCCCCCC
33.3724719451
1563PhosphorylationNMAIGDMSSLLTSLA
CCHHHCHHHHHHHHH
23.6722210691

Upstream regulatory proteins (kinases for phosphorylation sites, E3 ubiquitin ligases of ubiquitination sites, ...)
Modified Location Modified Residue Modification Type of Upstream Proteins Gene Name of Upstream Proteins UniProt AC of Upstream Proteins Sources
849SPhosphorylationKinasePRKACAP17612
GPS
849SPhosphorylationKinasePKA-Uniprot
849SPhosphorylationKinasePKA-FAMILY-GPS
865SPhosphorylationKinasePKA-FAMILY-GPS
865SPhosphorylationKinasePKA-Uniprot
865SPhosphorylationKinasePRKACAP17612
GPS
877SPhosphorylationKinasePKA-FAMILY-GPS
877SPhosphorylationKinasePRKACAP17612
GPS
877SPhosphorylationKinasePKA-Uniprot
907SPhosphorylationKinasePKA-FAMILY-GPS
907SPhosphorylationKinasePRKACAP17612
GPS
907SPhosphorylationKinasePKA-Uniprot
980SPhosphorylationKinasePRKACAP17612
GPS
980SPhosphorylationKinasePKA-FAMILY-GPS
980SPhosphorylationKinasePKA-Uniprot
1006SPhosphorylationKinasePRKACAP17612
GPS
1006SPhosphorylationKinasePKA-FAMILY-GPS
1006SPhosphorylationKinasePKA-Uniprot
-KUbiquitinationE3 ubiquitin ligaseBTRCQ9Y297
PMID:24658274
-KUbiquitinationE3 ubiquitin ligaseSPOPO43791
PMID:22199232

Functions of PTM Sites
Modified Location Modified Residue Modification Function Reference

Oops, there are no descriptions of PTM sites of GLI3_HUMAN !!

Disease-associated PTM Sites based on SAP

* Distance = the distance between SAP position and PTM sites.

Modified Location Modification Variant Position
(Distance <= 10)
Residue Change SAP Related Disease Reference

Oops, there are no SNP-PTM records of GLI3_HUMAN !!

Protein-Protein Interaction
Interacting Protein Gene Name Interaction Type PPI Reference Domain-Domain Interactions
ZIC1_HUMANZIC1physical
11238441
ZIC2_HUMANZIC2physical
11238441
ZIC3_HUMANZIC3physical
11238441
STK36_HUMANSTK36physical
10806483
SAP18_HUMANSAP18genetic
11960000
SUFU_MOUSESufugenetic
11960000
SIN3A_MOUSESin3agenetic
11960000
MED12_HUMANMED12physical
17000779
CBP_HUMANCREBBPphysical
17000779
FBW1A_HUMANBTRCphysical
16705181
SKI_HUMANSKIphysical
12435627
FBW1A_HUMANBTRCphysical
16371461
UBP7_HUMANUSP7physical
26120032

Drug and Disease Associations
Kegg Disease
There are no disease associations of PTM sites.
OMIM Disease
175700Greig cephalo-poly-syndactyly syndrome (GCPS)
146510Pallister-Hall syndrome (PHS)
174200Polydactyly, postaxial A1 (PAPA1)
174200Polydactyly, postaxial B (PAPB)
174700Polydactyly preaxial 4 (POP4)
Kegg Drug
There are no disease associations of PTM sites.
DrugBank
There are no disease associations of PTM sites.
Regulatory Network of GLI3_HUMAN

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Related Literatures of Post-Translational Modification
Acetylation
ReferencePubMed
"Lys-N and trypsin cover complementary parts of the phosphoproteome ina refined SCX-based approach.";
Gauci S., Helbig A.O., Slijper M., Krijgsveld J., Heck A.J.,Mohammed S.;
Anal. Chem. 81:4493-4501(2009).
Cited for: ACETYLATION [LARGE SCALE ANALYSIS] AT MET-1, AND MASS SPECTROMETRY.
Phosphorylation
ReferencePubMed
"Global, in vivo, and site-specific phosphorylation dynamics insignaling networks.";
Olsen J.V., Blagoev B., Gnad F., Macek B., Kumar C., Mortensen P.,Mann M.;
Cell 127:635-648(2006).
Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-19, AND MASSSPECTROMETRY.
"Hedgehog-regulated processing of Gli3 produces an anterior/posteriorrepressor gradient in the developing vertebrate limb.";
Wang B., Fallon J.F., Beachy P.A.;
Cell 100:423-434(2000).
Cited for: FUNCTION, PROTEOLYTIC PROCESSING, PHOSPHORYLATION AT SER-849; SER-865;SER-877; SER-907; SER-980 AND SER-1006, AND MUTAGENESIS OF SER-849;SER-865; SER-877; SER-907; SER-980 AND SER-1006.
Ubiquitylation
ReferencePubMed
"Multisite protein kinase A and glycogen synthase kinase 3betaphosphorylation leads to Gli3 ubiquitination by SCFbetaTrCP.";
Tempe D., Casas M., Karaz S., Blanchet-Tournier M.F., Concordet J.P.;
Mol. Cell. Biol. 26:4316-4326(2006).
Cited for: PROTEOLYTIC PROCESSING, PHOSPHORYLATION, INTERACTION WITH BTRC,UBIQUITINATION AT LYS-773; LYS-779; LYS-784 AND LYS-800, ANDMUTAGENESIS OF LYS-773; LYS-779; LYS-784; LYS-800; SER-849; SER-855;SER-856; SER-861; SER-864; SER-873; SER-877; SER-903 AND SER-907.

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