Swiss-Prot entry

ID   GCR_HUMAN      STANDARD;      PRT;   777 AA.
AC   P04150; P04151;
DT   01-NOV-1986 (Rel. 03, Created)
DT   01-NOV-1986 (Rel. 03, Last sequence update)
DT   01-MAY-2005 (Rel. 47, Last annotation update)
DE   Glucocorticoid receptor (GR).
GN   Name=NR3C1; Synonyms=GRL;
OS   Homo sapiens (Human).
OC   Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; 
OC   Mammalia; Eutheria; Euarchontoglires; Primates; Catarrhini; Hominidae; 
OC   Homo. 
OX   NCBI_TaxID=9606;
RN   [1]
RP   NUCLEOTIDE SEQUENCE [MRNA] (ISOFORMS ALPHA AND BETA).
RC   TISSUE=Fibroblast;
RX   MEDLINE=86092206; PubMed=2867473 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Hollenberg S.M., Weinberger C., Ong E.S., Cerelli G., Oro A., Lebo R.,
RA   Thompson E.B., Rosenfeld M.G., Evans R.M.;
RT   "Primary structure and expression of a functional human glucocorticoid
RT   receptor cDNA.";
RL   Nature 318:635-641(1985).
RN   [2]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA] (ISOFORMS ALPHA AND BETA).
RX   MEDLINE=91201378; PubMed=1707881 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Encio I.J., Detera-Wadleigh S.D.;
RT   "The genomic structure of the human glucocorticoid receptor.";
RL   J. Biol. Chem. 266:7182-7188(1991).
RN   [3]
RP   NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM GAMMA).
RA   Munroe D.G., Pang J., Taylor G.R., Lau C., Plante R.K., Zhou L.;
RT   "Alternative splicing within the DNA binding domain creates a novel
RT   isoform of the human glucocorticoid receptor.";
RL   Submitted (SEP-1993) to the EMBL/GenBank/DDBJ databases.
RN   [4]
RP   NUCLEOTIDE SEQUENCE [GENOMIC DNA] (ISOFORM ALPHA), AND VARIANTS LYS-23
RP   AND VAL-65.
RA   Rieder M.J., Livingston R.J., Daniels M.R., Chung M.-W.,
RA   Miyamoto K.E., Nguyen C.P., Nguyen D.A., Poel C.L., Robertson P.D.,
RA   Schackwitz W.S., Sherwood J.K., Witrak L.A., Nickerson D.A.;
RT   "NIEHS-SNPs, environmental genome project, NIEHS ES15478, Department
RT   of Genome Sciences, Seattle, WA (URL: http://egp.gs.washington.edu).";
RL   Submitted (OCT-2003) to the EMBL/GenBank/DDBJ databases.
RN   [5]
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM ALPHA).
RC   TISSUE=Placenta;
RX   MEDLINE=22388257; PubMed=12477932 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1073/pnas.242603899;
RA   Strausberg R.L., Feingold E.A., Grouse L.H., Derge J.G.,
RA   Klausner R.D., Collins F.S., Wagner L., Shenmen C.M., Schuler G.D.,
RA   Altschul S.F., Zeeberg B., Buetow K.H., Schaefer C.F., Bhat N.K.,
RA   Hopkins R.F., Jordan H., Moore T., Max S.I., Wang J., Hsieh F.,
RA   Diatchenko L., Marusina K., Farmer A.A., Rubin G.M., Hong L.,
RA   Stapleton M., Soares M.B., Bonaldo M.F., Casavant T.L., Scheetz T.E.,
RA   Brownstein M.J., Usdin T.B., Toshiyuki S., Carninci P., Prange C.,
RA   Raha S.S., Loquellano N.A., Peters G.J., Abramson R.D., Mullahy S.J.,
RA   Bosak S.A., McEwan P.J., McKernan K.J., Malek J.A., Gunaratne P.H.,
RA   Richards S., Worley K.C., Hale S., Garcia A.M., Gay L.J., Hulyk S.W.,
RA   Villalon D.K., Muzny D.M., Sodergren E.J., Lu X., Gibbs R.A.,
RA   Fahey J., Helton E., Ketteman M., Madan A., Rodrigues S., Sanchez A.,
RA   Whiting M., Madan A., Young A.C., Shevchenko Y., Bouffard G.G.,
RA   Blakesley R.W., Touchman J.W., Green E.D., Dickson M.C.,
RA   Rodriguez A.C., Grimwood J., Schmutz J., Myers R.M.,
RA   Butterfield Y.S.N., Krzywinski M.I., Skalska U., Smailus D.E.,
RA   Schnerch A., Schein J.E., Jones S.J.M., Marra M.A.;
RT   "Generation and initial analysis of more than 15,000 full-length human
RT   and mouse cDNA sequences.";
RL   Proc. Natl. Acad. Sci. U.S.A. 99:16899-16903(2002).
RN   [6]
RP   NUCLEOTIDE SEQUENCE OF 1-394.
RX   MEDLINE=91224961; PubMed=2026589 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Leclerc S., Xie B.X., Roy R., Govindan M.V.;
RT   "Purification of a human glucocorticoid receptor gene promoter-binding
RT   protein. Production of polyclonal antibodies against the purified
RT   factor.";
RL   J. Biol. Chem. 266:8711-8719(1991).
RN   [7]
RP   NUCLEOTIDE SEQUENCE OF 1-394.
RX   PubMed=1958537 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Govindan M.V., Pothier F., Leclerc S., Palaniswami R., Xie B.;
RT   "Human glucocorticoid receptor gene promotor-homologous down
RT   regulation.";
RL   J. Steroid Biochem. Mol. Biol. 40:317-323(1991).
RN   [8]
RP   NUCLEOTIDE SEQUENCE OF 396-630.
RA   Kimmerly W., Bondoc M., Cheng J., Connolly K.S., Gunning K.M.,
RA   Kadner K., Miguel T., Miller C., Pitluck S., Pollard M., Rojeski H.,
RA   Subramanian S., Martin C.H.;
RL   Submitted (SEP-1998) to the EMBL/GenBank/DDBJ databases.
RN   [9]
RP   ALTERNATIVE INITIATION, AND MUTAGENESIS OF MET-1 AND MET-27.
RX   MEDLINE=21329577; PubMed=11435610 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1210/me.15.7.1093;
RA   Yudt M.R., Cidlowski J.A.;
RT   "Molecular identification and characterization of A and B forms of the
RT   glucocorticoid receptor.";
RL   Mol. Endocrinol. 15:1093-1103(2001).
RN   [10]
RP   DOMAINS.
RX   MEDLINE=86092211; PubMed=3841189 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Weinberger C., Hollenberg S.M., Rosenfeld M.G., Evans R.M.;
RT   "Domain structure of human glucocorticoid receptor and its
RT   relationship to the v-erb-A oncogene product.";
RL   Nature 318:670-672(1985).
RN   [11]
RP   INTERACTIONS WITH TADA2L AND THE ADA COMPLEX, AND MUTAGENESIS OF
RP   PHE-191; ILE-193; LEU-194; LEU-197; TRP-213; LEU-224; LEU-225; PHE-235
RP   AND LEU-236.
RX   MEDLINE=97299656; PubMed=9154805 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Henriksson A., Almloef T., Ford J., McEwan I.J., Gustafsson J.-A.,
RA   Wright A.P.H.;
RT   "Role of the Ada adaptor complex in gene activation by the
RT   glucocorticoid receptor.";
RL   Mol. Cell. Biol. 17:3065-3073(1997).
RN   [12]
RP   INTERACTIONS WITH THE SMARCA4 COMPLEX; NCOA1; NCOA2 AND THE
RP   CREBBP/EP300 COMPLEX.
RX   MEDLINE=98250578; PubMed=9590696 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1038/30032;
RA   Fryer C.J., Archer T.K.;
RT   "Chromatin remodelling by the glucocorticoid receptor requires the
RT   BRG1 complex.";
RL   Nature 393:88-91(1998).
RN   [13]
RP   INTERACTION WITH BAG1.
RX   MEDLINE=99408777; PubMed=10477749 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1083/jcb.146.5.929;
RA   Schneikert J., Huebner S., Martin E., Cato A.B.C.;
RT   "A nuclear action of the eukaryotic cochaperone RAP46 in
RT   downregulation of glucocorticoid receptor activity.";
RL   J. Cell Biol. 146:929-940(1999).
RN   [14]
RP   ALTERNATIVE SPLICING (ISOFORM GAMMA).
RX   MEDLINE=20030962; PubMed=10566686 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1210/jc.84.11.4283;
RA   Rivers C., Levy A., Hancock J., Lightman S., Norman M.;
RT   "Insertion of an amino acid in the DNA-binding domain of the
RT   glucocorticoid receptor as a result of alternative splicing.";
RL   J. Clin. Endocrinol. Metab. 84:4283-4286(1999).
RN   [15]
RP   ALTERNATIVE SPLICING (ISOFORMS GP-P AND GP-A).
RX   PubMed=8358712 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Moalli P.A., Pillay S., Krett N.L., Rosen S.T.;
RT   "Alternatively spliced glucocorticoid receptor messenger RNAs in
RT   glucocorticoid-resistant human multiple myeloma cells.";
RL   Cancer Res. 53:3877-3879(1993).
RN   [16]
RP   REVIEW ON ALTERNATIVE SPLICING, ALTERNATIVE INITIATION, AND
RP   POSTTRANSLATION MODIFICATIONS.
RX   PubMed=15265776 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Lu N.Z., Cidlowski J.A.;
RT   "The origin and functions of multiple human glucocorticoid receptor
RT   isoforms.";
RL   Ann. N. Y. Acad. Sci. 1024:102-123(2004).
RN   [17]
RP   INTERACTION WITH NCOA6.
RX   MEDLINE=20325329; PubMed=10866662 [NCBI, ExPASy, EBI, Israel, Japan];
RX   DOI=10.1128/MCB.20.14.5048-5063.2000;
RA   Mahajan M.A., Samuels H.H.;
RT   "A new family of nuclear receptor coregulators that integrates nuclear
RT   receptor signaling through CBP.";
RL   Mol. Cell. Biol. 20:5048-5063(2000).
RN   [18]
RP   GLUCOCORTICOID-MEDIATED DOWN-REGULATION.
RX   MEDLINE=21560972; PubMed=11555652 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1074/jbc.M106033200;
RA   Wallace A.D., Cidlowski J.A.;
RT   "Proteasome-mediated glucocorticoid receptor degradation restricts
RT   transcriptional signaling by glucocorticoids.";
RL   J. Biol. Chem. 276:42714-42721(2001).
RN   [19]
RP   SUMOYLATION, AND MUTAGENESIS OF LYS-277; LYS-293 AND LYS-703.
RX   MEDLINE=22280317; PubMed=12144530 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1042/BJ20021085;
RA   Tian S., Poukka H., Palvimo J.J., Jaenne O.A.;
RT   "Small ubiquitin-related modifier-1 (SUMO-1) modification of the
RT   glucocorticoid receptor.";
RL   Biochem. J. 367:907-911(2002).
RN   [20]
RP   PHOSPHORYLATION SITES SER-203 AND SER-211.
RX   MEDLINE=22113030; PubMed=12000743 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1074/jbc.M110530200;
RA   Wang Z., Frederick J., Garabedian M.J.;
RT   "Deciphering the phosphorylation 'code' of the glucocorticoid receptor
RT   in vivo.";
RL   J. Biol. Chem. 277:26573-26580(2002).
RN   [21]
RP   X-RAY CRYSTALLOGRAPHY (2.5 ANGSTROMS) OF 521-777 OF MUTANT SER-602 IN
RP   COMPLEX WITH NCOA2; DEXAMETHASONE AND RU-486, AND MUTAGENESIS OF
RP   ARG-585; ASP-590; PHE-602; PRO-625 AND ILE-628.
RX   PubMed=12151000 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Bledsoe R.K., Montana V.G., Stanley T.B., Delves C.J., Apolito C.J.,
RA   McKee D.D., Consler T.G., Parks D.J., Stewart E.L., Willson T.M.,
RA   Lambert M.H., Moore J.T., Pearce K.H., Xu H.E.;
RT   "Crystal structure of the glucocorticoid receptor ligand binding
RT   domain reveals a novel mode of receptor dimerization and coactivator
RT   recognition.";
RL   Cell 110:93-105(2002).
RN   [22]
RP   X-RAY CRYSTALLOGRAPHY (2.3 ANGSTROMS) OF 500-777 OF MUTANT SER-602 IN
RP   COMPLEX WITH COACTIVATOR PEPTIDE; DEXAMETHASONE AND WITH RU-486.
RX   PubMed=12686538 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1074/jbc.M212711200;
RA   Kauppi B., Jakob C., Faernegaardh M., Yang J., Ahola H., Alarcon M.,
RA   Calles K., Engstrom O., Harlan J., Muchmore S., Ramqvist A.-K.,
RA   Thorell S., Oehman L., Greer J., Gustafsson J.-A., Carlstedt-Duke J.,
RA   Carlquist M.;
RT   "The three-dimensional structures of antagonistic and agonistic forms
RT   of the glucocorticoid receptor ligand-binding domain: RU-486 induces a
RT   transconformation that leads to active antagonism.";
RL   J. Biol. Chem. 278:22748-22754(2003).
RN   [23]
RP   CHARACTERIZATION OF VARIANT GLUCOCORTICOID RESISTANCE VAL-641.
RX   MEDLINE=91123468; PubMed=1704018 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Hurley D.M., Accili D., Stratakis C.A., Karl M., Vamvakopoulos N.,
RA   Rorer E., Constantine K., Taylor S.I., Chrousos G.P.;
RT   "Point mutation causing a single amino acid substitution in the
RT   hormone binding domain of the glucocorticoid receptor in familial
RT   glucocorticoid resistance.";
RL   J. Clin. Invest. 87:680-686(1991).
RN   [24]
RP   VARIANTS TYR-421 AND PHE-753.
RX   MEDLINE=93364907; PubMed=8358735 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Powers J.H., Hillmann A.G., Tang D.C., Harmon J.M.;
RT   "Cloning and expression of mutant glucocorticoid receptors from
RT   glucocorticoid-sensitive and -resistant human leukemic cells.";
RL   Cancer Res. 53:4059-4065(1993).
RN   [25]
RP   VARIANT SER-363.
RX   MEDLINE=93187003; PubMed=8445027 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1210/jc.76.3.683;
RA   Karl M., Lamberts S.W.J., Detera-Wadleigh S.D., Encio I.J.,
RA   Stratakis C.A., Hurley D.M., Accili D., Chrousos G.P.;
RT   "Familial glucocorticoid resistance caused by a splice site deletion
RT   in the human glucocorticoid receptor gene.";
RL   J. Clin. Endocrinol. Metab. 76:683-689(1993).
RN   [26]
RP   VARIANT GLUCOCORTICOID RESISTANCE ILE-729.
RX   MEDLINE=93253031; PubMed=7683692 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Malchoff D.M., Brufsky A., Reardon G., McDermott P., Javier E.C.,
RA   Bergh C.H., Rowe D., Malchoff C.D.;
RT   "A mutation of the glucocorticoid receptor in primary cortisol
RT   resistance.";
RL   J. Clin. Invest. 91:1918-1925(1993).
RN   [27]
RP   VARIANT PHE-753.
RX   MEDLINE=93302771; PubMed=8316249 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1210/me.7.5.631;
RA   Ashraf J., Thompson E.B.;
RT   "Identification of the activation-labile gene: a single point mutation
RT   in the human glucocorticoid receptor presents as two distinct receptor
RT   phenotypes.";
RL   Mol. Endocrinol. 7:631-642(1993).
RN   [28]
RP   VARIANTS LYS-23 AND SER-363.
RX   MEDLINE=97295085; PubMed=9150737 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Koper J.W., Stolk R.P., de Lange P., Huizenga N.A.T.M., Molijn G.-J.,
RA   Pols H.A.P., Grobbee D.E., Karl M., de Jong F.H., Brinkmann A.O.,
RA   Lamberts S.W.J.;
RT   "Lack of association between five polymorphisms in the human
RT   glucocorticoid receptor gene and glucocorticoid resistance.";
RL   Hum. Genet. 99:663-668(1997).
RN   [29]
RP   VARIANTS LYS-23; VAL-65 AND SER-363.
RX   MEDLINE=99318093; PubMed=10391209 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1038/10290;
RA   Cargill M., Altshuler D., Ireland J., Sklar P., Ardlie K., Patil N.,
RA   Shaw N., Lane C.R., Lim E.P., Kalyanaraman N., Nemesh J., Ziaugra L.,
RA   Friedland L., Rolfe A., Warrington J., Lipshutz R., Daley G.Q.,
RA   Lander E.S.;
RT   "Characterization of single-nucleotide polymorphisms in coding regions
RT   of human genes.";
RL   Nat. Genet. 22:231-238(1999).
RN   [30]
RP   ERRATUM.
RA   Cargill M., Altshuler D., Ireland J., Sklar P., Ardlie K., Patil N.,
RA   Shaw N., Lane C.R., Lim E.P., Kalyanaraman N., Nemesh J., Ziaugra L.,
RA   Friedland L., Rolfe A., Warrington J., Lipshutz R., Daley G.Q.,
RA   Lander E.S.;
RL   Nat. Genet. 23:373-373(1999).
RN   [31]
RP   VARIANTS LYS-23; LEU-29; PHE-112; ASN-233 AND SER-363.
RX   MEDLINE=20357652; PubMed=10898924 [NCBI, ExPASy, EBI, Israel, Japan];
RX   DOI=10.1002/1096-8628(20000612)96:3<412::AID-AJMG33>3.0.CO;2-C;
RA   Feng J., Zheng J., Bennett W.P., Heston L.L., Jones I.R., Craddock N.,
RA   Sommer S.S.;
RT   "Five missense variants in the amino-terminal domain of the
RT   glucocorticoid receptor: no association with puerperal psychosis or
RT   schizophrenia.";
RL   Am. J. Med. Genet. 96:412-417(2000).
RN   [32]
RP   VARIANTS GLUCOCORTICOID RESISTANCE HIS-477 AND SER-679.
RX   MEDLINE=21473978; PubMed=11589680 [NCBI, ExPASy, EBI, Israel, Japan];
RA   Ruiz M., Lind U., Gaafvels M., Eggertsen G., Carlstedt-Duke J.,
RA   Nilsson L., Holtmann M., Stierna P., Wikstroem A.-C., Werner S.;
RT   "Characterization of two novel mutations in the glucocorticoid
RT   receptor gene in patients with primary cortisol resistance.";
RL   Clin. Endocrinol. (Oxf.) 55:363-371(2001).
RN   [33]
RP   CHARACTERIZATION OF VARIANT GLUCOCORTICOID RESISTANCE ASN-559.
RX   MEDLINE=21558592; PubMed=11701741 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1210/jc.86.11.5600;
RA   Kino T., Stauber R.H., Resau J.H., Pavlakis G.N., Chrousos G.P.;
RT   "Pathologic human GR mutant has a transdominant negative effect on the
RT   wild-type GR by inhibiting its translocation into the nucleus:
RT   importance of the ligand-binding domain for intracellular GR
RT   trafficking.";
RL   J. Clin. Endocrinol. Metab. 86:5600-5608(2001).
RN   [34]
RP   VARIANT GLUCOCORTICOID RESISTANCE MET-747, AND ALTERED INTERACTION
RP   WITH THE COACTIVATOR NCOA2.
RX   MEDLINE=22045363; PubMed=12050230 [NCBI, ExPASy, EBI, Israel, Japan]; DOI=10.1210/jc.87.6.2658;
RA   Vottero A., Kino T., Combe H., Lecomte P., Chrousos G.P.;
RT   "A novel, C-terminal dominant negative mutation of the GR causes
RT   familial glucocorticoid resistance through abnormal interactions with
RT   p160 steroid receptor coactivators.";
RL   J. Clin. Endocrinol. Metab. 87:2658-2667(2002).
CC   -!- FUNCTION: Receptor for glucocorticoids (GC). Has a dual mode of
CC       action: as a transcription factor that binds to glucocorticoid
CC       response elements (GRE) and as a modulator of other transcription
CC       factors. Affects inflammatory responses, cellular proliferation
CC       and differentiation in target tissues.
CC   -!- SUBUNIT: Heteromultimeric cytoplasmic complex with HSP90, HSP70,
CC       and FKBP5 or another immunophilin, or the immunophilin homolog
CC       PPP5C. Upon ligand binding FKBP5 dissociates from the complex and
CC       FKBP4 takes its place, thereby linking the complex to dynein and
CC       mediating transport to the nucleus, where the complex dissociates
CC       (By similarity). Binds to DNA as a homodimer, and as a heterodimer
CC       with NR3C2 or the retinoid X receptor. Binds STAT5A and STAT5B
CC       homodimers and heterodimers. Interacts with NRIP1, POU2F1, POU2F2
CC       and TRIM28 (By similarity). Interacts with several coactivator
CC       complexes, including the SMARCA4 complex, CREBBP/EP300, TADA2L and
CC       p160 coactivators such as NCOA2 and NCOA6. Interaction with BAG1
CC       inhibits transactivation.
CC   -!- SUBCELLULAR LOCATION: Cytoplasmic in the absence of ligand;
CC       nuclear after ligand-binding.
CC   -!- ALTERNATIVE PRODUCTS:
CC       Event=Alternative splicing; Named isoforms=5;
CC         Comment=Additional isoforms seem to exist;
CC       Name=Alpha; Synonyms=Alpha-A;
CC         IsoId=P04150-1; Sequence=Displayed;
CC         Note=Predominant physiological form. Isoform Alpha-B is produced
CC         by alternative initiation at Met-27 of isoform Alpha. Both
CC         isoforms exhibit similar subcellular location and nuclear
CC         translocation after ligand activation. Isoform Alpha-B appears
CC         to be more susceptible to degradation, at least when expressed
CC         in mammalian cells, but more effective in transcriptional
CC         activation and not in transrepression;
CC       Name=Beta; Synonyms=Beta-A;
CC         IsoId=P04150-2; Sequence=VSP_003703;
CC         Note=No hormone-binding activity. Widely expressed at low level.
CC         Localized largely in the nucleus. Isoform Beta-B is produced by
CC         alternative initiation at Met-27 of isoform Beta;
CC       Name=Gamma; Synonyms=Alpha-2, Gamma-A, Alpha-2-A;
CC         IsoId=P04150-3; Sequence=VSP_007363;
CC         Note=Lower transcriptional activity. Expressed at low level;
CC       Name=GR-P;
CC         IsoId=P04150-4; Sequence=Not described;
CC         Note=Encoded by exons 2-7 plus several basepairs from the
CC         subsequent intron region. Lacks the ligand binding domain.
CC         Accounts for up to 10-20% of mRNAs;
CC       Name=GR-A;
CC         IsoId=P04150-5; Sequence=VSP_013340;
CC         Note=Lacks exons 5, 6 and 7. Found in glucocorticoid-resistant
CC         myeloma patients;
CC       Event=Alternative initiation;
CC         Comment=At least 4 isoforms, Alpha (shown here), Alpha-B, Beta
CC         and Beta-B, are produced by alternative initiation at Met-1 and
CC         Met-27. The existence of isoform Alpha and isoform Alpha-B has
CC         been proved by mutagenesis. As the sequence environment of the 2
CC         potential ATG initiator codons is the same for the other
CC         isoforms, alternative initiation of translation could also occur
CC         on these transcripts;
CC   -!- TISSUE SPECIFICITY: Widely expressed.
CC   -!- DOMAIN: Composed of three domains: a modulating N-terminal domain,
CC       a DNA-binding domain and a C-terminal steroid-binding domain.
CC   -!- PTM: Increased proteasome-mediated degradation in response to
CC       glucocorticoids.
CC   -!- PTM: Phosphorylated in the absence of hormone; becomes
CC       hyperphosphorylated in the presence of glucocorticoid. The Ser-
CC       203-phosphorylated form is mainly cytoplasmic, and the Ser-211-
CC       phosphorylated form is nuclear. Transcriptional activity
CC       correlates with the amount of phosphorylation at Ser-211.
CC   -!- PTM: Sumoylated; this reduces transcription transactivation.
CC   -!- DISEASE: Defects in NR3C1 are a cause of glucocorticoid resistance
CC       [MIM:138040]; also known as cortisol resistance. It is a
CC       hypertensive, hyperandrogenic disorder characterized by increased
CC       serum cortisol concentrations. Inheritance is autosomal dominant.
CC   -!- SIMILARITY: Belongs to the nuclear hormone receptor family. NR3
CC       subfamily.
CC   -!- SIMILARITY: Contains 1 nuclear receptor DNA-binding domain.
CC   --------------------------------------------------------------------------
CC   This Swiss-Prot entry is copyright. It is produced through a collaboration
CC   between  the Swiss Institute of Bioinformatics  and the  EMBL outstation -
CC   the European Bioinformatics Institute.  There are no  restrictions on  its
CC   use as long as its content is in no way modified and this statement is not
CC   removed.
CC   --------------------------------------------------------------------------
DR   EMBL; X03225; CAA26976.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; X03348; CAA27054.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U80946; AAB64353.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78506; AAB64353.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78507; AAB64353.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78508; AAB64353.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78509; AAB64353.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78510; AAB64353.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78511; AAB64353.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78512; AAB64353.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U80947; AAB64354.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78506; AAB64354.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78507; AAB64354.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78508; AAB64354.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78509; AAB64354.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78510; AAB64354.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78511; AAB64354.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U78512; AAB64354.1; JOINED. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; U01351; AAA16603.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; AY436590; AAQ97180.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; BC015610; AAH15610.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; M69104; AAA88049.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; M73816; AAA53151.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; S68378; AAB20466.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   EMBL; AC005601; AAC34207.1; -. [EMBL / GenBank / DDBJ] [CoDingSequence]
DR   PIR; A93370; QRHUGA.
DR   PIR; B93370; QRHUGB.
DR   PDB; 1M2Z; X-ray; A/D=521-777. [ExPASy / RCSB]
DR   PDB; 1NHZ; X-ray; A=500-777. [ExPASy / RCSB]
DR   PDB; 1P93; X-ray; A/B/C/D=500-777. [ExPASy / RCSB]
DR   SMR; P04150; 417-491.
DR   IntAct; P04150; -.
DR   TRANSFAC; T00337; -.
DR   TRANSFAC; T01920; -.
DR   Ensembl; ENSG00000113580; Homo_sapiens
DR   Genew; HGNC:7978; NR3C1.
DR   CleanEx; HGNC:7978; NR3C1.
DR   H-InvDB; HIX0005273; -.
DR   MIM; 138040; -. [NCBI / EBI]
DR   GeneCards; NR3C1.
DR   GeneLynx; NR3C1.
DR   GenAtlas; NR3C1.
DR   SOURCE; NR3C1.
DR   GO; GO:0005737; C:cytoplasm; TAS.
DR   GO; GO:0005759; C:mitochondrial matrix; TAS.
DR   GO; GO:0005634; C:nucleus; TAS.
DR   GO; GO:0004883; F:glucocorticoid receptor activity; TAS.
DR   GO; GO:0003700; F:transcription factor activity; TAS.
DR   GO; GO:0007165; P:signal transduction; TAS.
DR   GO; GO:0006366; P:transcription from Pol II promoter; TAS.
DR   InterPro; IPR001409; Glcrtcd_receptor.
DR   InterPro; IPR000536; Hrmon_recept_lig.
DR   InterPro; IPR001723; Stdhrmn_receptor.
DR   InterPro; IPR008946; Str_ncl_receptor.
DR   InterPro; IPR001628; Znf_C4steroid.
DR   InterPro; Graphical view of domain structure.
DR   Pfam; PF02155; GCR; 1.
DR   Pfam; PF00104; Hormone_recep; 1.
DR   Pfam; PF00105; zf-C4; 1.
DR   Pfam; Graphical view of domain structure.
DR   PRINTS; PR00528; GLCORTICOIDR.
DR   PRINTS; PR00398; STRDHORMONER.
DR   PRINTS; PR00047; STROIDFINGER.
DR   ProDom; PD000035; Znf_C4steroid; 1.
DR   ProDom [Domain structure / List of seq. sharing at least 1 domain ]
DR   SMART; SM00430; HOLI; 1.
DR   SMART; SM00399; ZnF_C4; 1.
DR   PROSITE; PS00031; NUCLEAR_REC_DBD_1; 1.
DR   PROSITE; PS51030; NUCLEAR_REC_DBD_2; 1.
DR   CMR; P04150.
DR   HOVERGEN [Family / Alignment / Tree]
DR   BLOCKS; P04150.
DR   ProtoNet; P04150.
DR   ProtoMap; P04150.
DR   PRESAGE; P04150.
DR   DIP; P04150.
DR   ModBase; P04150.
DR   SWISS-2DPAGE; GET REGION ON 2D PAGE.
KW   3D-structure; Alternative initiation; Alternative splicing;
KW   Disease mutation; DNA-binding; Nuclear protein; Phosphorylation;
KW   Polymorphism; Receptor; Steroid-binding; Trans-acting factor;
KW   Transcription; Transcription regulation; Ubl conjugation; Zinc-finger.
FT   CHAIN         1    777       Glucocorticoid receptor, A-type isoforms.
FT   CHAIN        27    777       Glucocorticoid receptor, B-type isoforms.
FT   INIT_MET     27     27       For B-type isoforms.
FT   DOMAIN        1    420       Modulating.
FT   DOMAIN      399    418       Glu/Ser/Pro/Thr-rich (PEST region)
FT                                (Potential).
FT   DNA_BIND    421    486       Nuclear receptor-type.
FT   ZN_FING     421    441       C4-type.
FT   ZN_FING     457    481       C4-type.
FT   DOMAIN      487    527       Hinge.
FT   DOMAIN      528    777       Steroid-binding.
FT   MOD_RES     113    113       Phosphoserine (By similarity).
FT   MOD_RES     141    141       Phosphoserine (By similarity).
FT   MOD_RES     203    203       Phosphoserine.
FT   MOD_RES     211    211       Phosphoserine.
FT   MOD_RES     226    226       Phosphoserine (By similarity).
FT   VARSPLIC    728    777       VVENLLNYCFQTFLDKTMSIEFPEMLAEIITNQIPKYSNGN
FT                                IKKLLFHQK -> NVMWLKPESTSHTLI (in isoform
FT                                Beta).
FT                                /FTId=VSP_003703.
FT   VARSPLIC    451    451       G -> GR (in isoform Gamma).
FT                                /FTId=VSP_007363.
FT   VARSPLIC    491    674       Missing (in isoform GR-A).
FT                                /FTId=VSP_013340.
FT   VARIANT      23     23       R -> K (in dbSNP:6190).
FT                                /FTId=VAR_014140.
FT   VARIANT      29     29       F -> L.
FT                                /FTId=VAR_015628.
FT   VARIANT      65     65       F -> V (in dbSNP:6192).
FT                                /FTId=VAR_014622.
FT   VARIANT     112    112       L -> F.
FT                                /FTId=VAR_015629.
FT   VARIANT     233    233       D -> N.
FT                                /FTId=VAR_015630.
FT   VARIANT     363    363       N -> S (may increase sensitivity to
FT                                exogenously administered glucocorticoids;
FT                                dbSNP:6195).
FT                                /FTId=VAR_004675.
FT   VARIANT     421    421       C -> Y (in a glucocorticoid resistant
FT                                leukemia cell line).
FT                                /FTId=VAR_015631.
FT   VARIANT     477    477       R -> H (in glucocorticoid resistance).
FT                                /FTId=VAR_013472.
FT   VARIANT     559    559       I -> N (in glucocorticoid resistance;
FT                                interferes with translocation to the
FT                                nucleus and thereby strongly reduces
FT                                transcription activation. Is equally
FT                                impaired in nuclear export. Acts as
FT                                dominant negative mutant).
FT                                /FTId=VAR_015632.
FT   VARIANT     641    641       D -> V (in glucocorticoid resistance).
FT                                /FTId=VAR_004676.
FT   VARIANT     679    679       G -> S (in glucocorticoid resistance; has
FT                                50% binding affinity).
FT                                /FTId=VAR_013473.
FT   VARIANT     729    729       V -> I (in glucocorticoid resistance).
FT                                /FTId=VAR_004677.
FT   VARIANT     747    747       I -> M (in glucocorticoid resistance;
FT                                alters interaction with NCOA2 and
FT                                strongly reduces transcription
FT                                activation. Acts as dominant negative
FT                                mutant).
FT                                /FTId=VAR_015633.
FT   VARIANT     753    753       L -> F (in two glucocorticoid resistant
FT                                leukemia cell lines lacking the normal
FT                                allele).
FT                                /FTId=VAR_004678.
FT   MUTAGEN       1      1       M->T: Abolishes expression of A-type
FT                                isoforms.
FT   MUTAGEN      27     27       M->T: Abolishes expression of B-type
FT                                isoforms.
FT   MUTAGEN     191    191       F->D: Reduces transactivation by the ADA
FT                                complex.
FT   MUTAGEN     193    193       I->D: Reduces transactivation by the ADA
FT                                complex.
FT   MUTAGEN     194    194       L->A: Strongly reduces transactivation by
FT                                the ADA complex; when associated with V-
FT                                224 and F-225.
FT   MUTAGEN     197    197       L->E: Reduces transactivation by the ADA
FT                                complex.
FT   MUTAGEN     213    213       W->A: Strongly reduces transactivation by
FT                                the ADA complex.
FT   MUTAGEN     224    224       L->V: Strongly reduces transactivation by
FT                                the ADA complex; when associated with A-
FT                                194 and F-225.
FT   MUTAGEN     225    225       L->F: Strongly reduces transactivation by
FT                                the ADA complex; when associated with A-
FT                                194 and V-224.
FT   MUTAGEN     235    235       F->L: Strongly reduces transactivation by
FT                                the ADA complex; when associated with V-
FT                                236.
FT   MUTAGEN     236    236       L->V: Strongly reduces transactivation by
FT                                the ADA complex; when associated with L-
FT                                235.
FT   MUTAGEN     277    277       K->R: Strongly reduces sumoylation.
FT                                Almost complete loss of sumoylation; when
FT                                associated with R-293.
FT   MUTAGEN     293    293       K->R: Strongly reduces sumoylation.
FT                                Almost complete loss of sumoylation; when
FT                                associated with R-277.
FT   MUTAGEN     585    585       R->A: Reduces activation mediated by
FT                                ligand binding domain; when associated
FT                                with A-590.
FT   MUTAGEN     590    590       D->A: Reduces activation mediated by
FT                                ligand binding domain; when associated
FT                                with A-585.
FT   MUTAGEN     602    602       F->S: Increases solubility. No effect on
FT                                transactivation by dexamethasone.
FT   MUTAGEN     625    625       P->A: Decreases transactivation by
FT                                dexamethasone by 95%.
FT   MUTAGEN     628    628       I->A: Decreases dimerization and
FT                                transactivation by dexamethasone; when
FT                                associated with S-602.
FT   MUTAGEN     703    703       K->R: Slightly reduces sumoylation.
SQ   SEQUENCE   777 AA;  85659 MW;  C5C90C9A5DD16AAB CRC64;
     MDSKESLTPG REENPSSVLA QERGDVMDFY KTLRGGATVK VSASSPSLAV ASQSDSKQRR
     LLVDFPKGSV SNAQQPDLSK AVSLSMGLYM GETETKVMGN DLGFPQQGQI SLSSGETDLK
     LLEESIANLN RSTSVPENPK SSASTAVSAA PTEKEFPKTH SDVSSEQQHL KGQTGTNGGN
     VKLYTTDQST FDILQDLEFS SGSPGKETNE SPWRSDLLID ENCLLSPLAG EDDSFLLEGN
     SNEDCKPLIL PDTKPKIKDN GDLVLSSPSN VTLPQVKTEK EDFIELCTPG VIKQEKLGTV
     YCQASFPGAN IIGNKMSAIS VHGVSTSGGQ MYHYDMNTAS LSQQQDQKPI FNVIPPIPVG
     SENWNRCQGS GDDNLTSLGT LNFPGRTVFS NGYSSPSMRP DVSSPPSSSS TATTGPPPKL
     CLVCSDEASG CHYGVLTCGS CKVFFKRAVE GQHNYLCAGR NDCIIDKIRR KNCPACRYRK
     CLQAGMNLEA RKTKKKIKGI QQATTGVSQE TSENPGNKTI VPATLPQLTP TLVSLLEVIE
     PEVLYAGYDS SVPDSTWRIM TTLNMLGGRQ VIAAVKWAKA IPGFRNLHLD DQMTLLQYSW
     MFLMAFALGW RSYRQSSANL LCFAPDLIIN EQRMTLPCMY DQCKHMLYVS SELHRLQVSY
     EEYLCMKTLL LLSSVPKDGL KSQELFDEIR MTYIKELGKA IVKREGNSSQ NWQRFYQLTK
     LLDSMHEVVE NLLNYCFQTF LDKTMSIEFP EMLAEIITNQ IPKYSNGNIK KLLFHQK
//