ISL1_MOUSE
ID ISL1_MOUSE Reviewed; 349 AA.
AC P61372; P20663; P47894; Q812D8;
DT 24-MAY-2004, integrated into UniProtKB/Swiss-Prot.
DT 24-MAY-2004, sequence version 1.
DT 03-AUG-2022, entry version 160.
DE RecName: Full=Insulin gene enhancer protein ISL-1;
DE Short=Islet-1;
GN Name=Isl1;
OS Mus musculus (Mouse).
OC Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia;
OC Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha; Muroidea; Muridae;
OC Murinae; Mus; Mus.
OX NCBI_TaxID=10090;
RN [1]
RP NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1), FUNCTION, ALTERNATIVE SPLICING,
RP SUBCELLULAR LOCATION (ISOFORMS 1 AND 2), AND PHOSPHORYLATION (ISOFORM 1).
RX PubMed=14664703; DOI=10.1677/jme.0.0310419;
RA Ando K., Shioda S., Handa H., Kataoka K.;
RT "Isolation and characterization of an alternatively spliced variant of
RT transcription factor Islet-1.";
RL J. Mol. Endocrinol. 31:419-425(2003).
RN [2]
RP NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 2).
RC STRAIN=C57BL/6J;
RA Koehler K., Dear T.N.;
RT "Mouse Isl1 cDNA.";
RL Submitted (MAR-1999) to the EMBL/GenBank/DDBJ databases.
RN [3]
RP FUNCTION, DISRUPTION PHENOTYPE, AND DEVELOPMENTAL STAGE.
RX PubMed=14667410; DOI=10.1016/s1534-5807(03)00363-0;
RA Cai C.L., Liang X., Shi Y., Chu P.H., Pfaff S.L., Chen J., Evans S.;
RT "Isl1 identifies a cardiac progenitor population that proliferates prior to
RT differentiation and contributes a majority of cells to the heart.";
RL Dev. Cell 5:877-889(2003).
RN [4]
RP FUNCTION, DEVELOPMENTAL STAGE, AND INTERACTION WITH MLIP.
RX PubMed=22343712; DOI=10.1161/circresaha.111.259663;
RA Huang Z.P., Young Seok H., Zhou B., Chen J., Chen J.F., Tao Y., Pu W.T.,
RA Wang D.Z.;
RT "CIP, a cardiac Isl1-interacting protein, represses cardiomyocyte
RT hypertrophy.";
RL Circ. Res. 110:818-830(2012).
RN [5]
RP FUNCTION, DNA-BINDING, INTERACTION WITH POU3F2; POU4F2 AND POU4F3, AND
RP DISRUPTION PHENOTYPE.
RX PubMed=24643061; DOI=10.1371/journal.pone.0092105;
RA Li R., Wu F., Ruonala R., Sapkota D., Hu Z., Mu X.;
RT "Isl1 and Pou4f2 form a complex to regulate target genes in developing
RT retinal ganglion cells.";
RL PLoS ONE 9:E92105-E92105(2014).
RN [6]
RP FUNCTION.
RX PubMed=25775587; DOI=10.1073/pnas.1421535112;
RA Wu F., Kaczynski T.J., Sethuramanujam S., Li R., Jain V., Slaughter M.,
RA Mu X.;
RT "Two transcription factors, Pou4f2 and Isl1, are sufficient to specify the
RT retinal ganglion cell fate.";
RL Proc. Natl. Acad. Sci. U.S.A. 112:E1559-E1568(2015).
RN [7]
RP X-RAY CRYSTALLOGRAPHY (2.05 ANGSTROMS) OF 262-291 IN COMPLEX WITH LHX3, AND
RP FUNCTION.
RX PubMed=18583962; DOI=10.1038/emboj.2008.123;
RA Bhati M., Lee C., Nancarrow A.L., Lee M., Craig V.J., Bach I., Guss J.M.,
RA Mackay J.P., Matthews J.M.;
RT "Implementing the LIM code: the structural basis for cell type-specific
RT assembly of LIM-homeodomain complexes.";
RL EMBO J. 27:2018-2029(2008).
CC -!- FUNCTION: DNA-binding transcriptional activator (PubMed:14664703,
CC PubMed:24643061, PubMed:25775587, PubMed:22343712). Recognizes and
CC binds to the consensus octamer binding site 5'-ATAATTAA-3' in promoter
CC of target genes (PubMed:24643061, PubMed:25775587). Plays a fundamental
CC role in the gene regulatory network essential for retinal ganglion cell
CC (RGC) differentiation (PubMed:25775587). Cooperates with the
CC transcription factor POU4F2 to achieve maximal levels of expression of
CC RGC target genes and RGC fate specification in the developing retina
CC (PubMed:24643061, PubMed:25775587). Involved in the specification of
CC motor neurons in cooperation with LHX3 and LDB1 (PubMed:18583962).
CC Binds to insulin gene enhancer sequences (By similarity). Essential for
CC heart development. Marker of one progenitor cell population that give
CC rise to the outflow tract, right ventricle, a subset of left
CC ventricular cells, and a large number of atrial cells as well, its
CC function is required for these progenitors to contribute to the heart.
CC Controls the expression of FGF and BMP growth factors in this cell
CC population and is required for proliferation and survival of cells
CC within pharyngeal foregut endoderm and adjacent splanchnic mesoderm as
CC well as for migration of cardiac progenitors into the heart
CC (PubMed:14667410). {ECO:0000250|UniProtKB:P61374,
CC ECO:0000269|PubMed:14664703, ECO:0000269|PubMed:14667410,
CC ECO:0000269|PubMed:18583962, ECO:0000269|PubMed:22343712,
CC ECO:0000269|PubMed:24643061, ECO:0000269|PubMed:25775587}.
CC -!- SUBUNIT: At neuronal promoters, displaces LDB1 from LHX3 LIM domain to
CC form a ternary complex in which ISL1 contacts both LHX3 and LDB1
CC (PubMed:18583962). Interacts (via C-terminus) with POU4F2 (via C-
CC terminus) isoform 1 (PubMed:24643061). Interacts with POU3F2
CC (PubMed:24643061). Interacts with POU4F3 (PubMed:24643061). Interacts
CC (via N-terminal domain) with MLIP; the interaction represses ISL1
CC transactivator activity (PubMed:22343712).
CC {ECO:0000269|PubMed:18583962, ECO:0000269|PubMed:22343712,
CC ECO:0000269|PubMed:24643061}.
CC -!- INTERACTION:
CC P61372; P70662: Ldb1; NbExp=4; IntAct=EBI-7988215, EBI-6272082;
CC P61372; P50481: Lhx3; NbExp=7; IntAct=EBI-7988215, EBI-7988290;
CC -!- SUBCELLULAR LOCATION: [Isoform 1]: Nucleus
CC {ECO:0000269|PubMed:14664703}.
CC -!- SUBCELLULAR LOCATION: [Isoform 2]: Nucleus
CC {ECO:0000269|PubMed:14664703}.
CC -!- ALTERNATIVE PRODUCTS:
CC Event=Alternative splicing; Named isoforms=2;
CC Name=1; Synonyms=Isl1-alpha {ECO:0000303|PubMed:14664703};
CC IsoId=P61372-1; Sequence=Displayed;
CC Name=2; Synonyms=Isl1-beta {ECO:0000303|PubMed:14664703};
CC IsoId=P61372-2; Sequence=VSP_010338;
CC -!- DEVELOPMENTAL STAGE: Between 7.5 dpc and 8.5 dpc, as the heart tube
CC forms, expressed in splanchnic mesenchyme comprising the mesocardium
CC and adjacent to foregut endoderm as well as in both splanchnic mesoderm
CC and in ventral foregut endoderm. At 10 dpc, continues to be expressed
CC in ventral endoderm and splanchnic mesoderm but is not expressed in the
CC myocardium of the heart (PubMed:14667410). At 10.5 dpc, expressed in
CC cardiomyocytes located in the outflow tract (PubMed:22343712).
CC {ECO:0000269|PubMed:14667410, ECO:0000269|PubMed:22343712}.
CC -!- PTM: Isoform 1 is phosphorylated. {ECO:0000269|PubMed:14664703}.
CC -!- DISRUPTION PHENOTYPE: Embryonic mutants exhibit growth retardation at
CC approximately 9.5 dpc and die at approximately 10.5 dpc. Between 9.0
CC dpc and 9.5 dpc hearts are severely abnormal, appear misshapen and
CC unlooped. Hearts are completely missing the outflow tract, right
CC ventricle, and much of the atria (PubMed:14667410). Conditional mutants
CC for retina expression show a decrease in several gene expression levels
CC involved in the differentiation of retinal ganglion cells (RGC)
CC (PubMed:24643061). {ECO:0000269|PubMed:14667410,
CC ECO:0000269|PubMed:24643061}.
CC -!- MISCELLANEOUS: [Isoform 2]: Preferentially expressed in insulinoma cell
CC lines. Expression is much lower than that of isoform 1. Shows
CC relatively higher transcriptional activity than isoform 1.
CC {ECO:0000269|PubMed:14664703}.
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DR EMBL; AB104633; BAC57891.1; -; mRNA.
DR EMBL; AJ132765; CAB38446.1; -; mRNA.
DR CCDS; CCDS26790.1; -. [P61372-1]
DR RefSeq; NP_067434.3; NM_021459.4. [P61372-1]
DR RefSeq; XP_006517596.1; XM_006517533.2.
DR PDB; 2RGT; X-ray; 2.05 A; A/B=262-291.
DR PDB; 4JCJ; X-ray; 3.00 A; A/B/C=123-138.
DR PDBsum; 2RGT; -.
DR PDBsum; 4JCJ; -.
DR AlphaFoldDB; P61372; -.
DR SASBDB; P61372; -.
DR SMR; P61372; -.
DR BioGRID; 200812; 8.
DR CORUM; P61372; -.
DR IntAct; P61372; 4.
DR MINT; P61372; -.
DR STRING; 10090.ENSMUSP00000044879; -.
DR iPTMnet; P61372; -.
DR PhosphoSitePlus; P61372; -.
DR MaxQB; P61372; -.
DR PaxDb; P61372; -.
DR PRIDE; P61372; -.
DR ProteomicsDB; 269000; -. [P61372-1]
DR ProteomicsDB; 269001; -. [P61372-2]
DR Antibodypedia; 23291; 596 antibodies from 37 providers.
DR DNASU; 16392; -.
DR Ensembl; ENSMUST00000036060; ENSMUSP00000044879; ENSMUSG00000042258. [P61372-1]
DR Ensembl; ENSMUST00000176044; ENSMUSP00000135567; ENSMUSG00000042258. [P61372-2]
DR GeneID; 16392; -.
DR KEGG; mmu:16392; -.
DR UCSC; uc007rye.2; mouse. [P61372-1]
DR UCSC; uc007ryf.2; mouse. [P61372-2]
DR CTD; 3670; -.
DR MGI; MGI:101791; Isl1.
DR VEuPathDB; HostDB:ENSMUSG00000042258; -.
DR eggNOG; KOG0490; Eukaryota.
DR GeneTree; ENSGT00940000153731; -.
DR HOGENOM; CLU_027802_2_0_1; -.
DR InParanoid; P61372; -.
DR OMA; VSQMHGY; -.
DR PhylomeDB; P61372; -.
DR TreeFam; TF315442; -.
DR BioGRID-ORCS; 16392; 0 hits in 74 CRISPR screens.
DR ChiTaRS; Isl1; mouse.
DR EvolutionaryTrace; P61372; -.
DR PRO; PR:P61372; -.
DR Proteomes; UP000000589; Chromosome 13.
DR RNAct; P61372; protein.
DR Bgee; ENSMUSG00000042258; Expressed in superior cervical ganglion and 224 other tissues.
DR ExpressionAtlas; P61372; baseline and differential.
DR Genevisible; P61372; MM.
DR GO; GO:0005654; C:nucleoplasm; TAS:Reactome.
DR GO; GO:0005634; C:nucleus; IDA:UniProtKB.
DR GO; GO:0043425; F:bHLH transcription factor binding; ISO:MGI.
DR GO; GO:0003682; F:chromatin binding; IDA:MGI.
DR GO; GO:0000987; F:cis-regulatory region sequence-specific DNA binding; ISO:MGI.
DR GO; GO:0003677; F:DNA binding; IDA:MGI.
DR GO; GO:0001228; F:DNA-binding transcription activator activity, RNA polymerase II-specific; IDA:MGI.
DR GO; GO:0000981; F:DNA-binding transcription factor activity, RNA polymerase II-specific; ISO:MGI.
DR GO; GO:0046872; F:metal ion binding; IEA:UniProtKB-KW.
DR GO; GO:0030331; F:nuclear estrogen receptor binding; ISO:MGI.
DR GO; GO:0016922; F:nuclear receptor binding; ISO:MGI.
DR GO; GO:1990841; F:promoter-specific chromatin binding; IDA:UniProtKB.
DR GO; GO:0000978; F:RNA polymerase II cis-regulatory region sequence-specific DNA binding; ISO:MGI.
DR GO; GO:0061629; F:RNA polymerase II-specific DNA-binding transcription factor binding; ISO:MGI.
DR GO; GO:0043565; F:sequence-specific DNA binding; IDA:UniProtKB.
DR GO; GO:1990837; F:sequence-specific double-stranded DNA binding; ISO:MGI.
DR GO; GO:0060413; P:atrial septum morphogenesis; IGI:BHF-UCL.
DR GO; GO:0031103; P:axon regeneration; IEA:Ensembl.
DR GO; GO:0007409; P:axonogenesis; IBA:GO_Central.
DR GO; GO:0060070; P:canonical Wnt signaling pathway; IGI:MGI.
DR GO; GO:0060913; P:cardiac cell fate determination; IMP:MGI.
DR GO; GO:0060379; P:cardiac muscle cell myoblast differentiation; IMP:MGI.
DR GO; GO:0003215; P:cardiac right ventricle morphogenesis; IGI:BHF-UCL.
DR GO; GO:0008283; P:cell population proliferation; IDA:MGI.
DR GO; GO:0071385; P:cellular response to glucocorticoid stimulus; IEA:Ensembl.
DR GO; GO:0003203; P:endocardial cushion morphogenesis; IGI:BHF-UCL.
DR GO; GO:0007507; P:heart development; IMP:MGI.
DR GO; GO:0003007; P:heart morphogenesis; IGI:MGI.
DR GO; GO:0060384; P:innervation; IMP:BHF-UCL.
DR GO; GO:0048762; P:mesenchymal cell differentiation; IMP:MGI.
DR GO; GO:0090090; P:negative regulation of canonical Wnt signaling pathway; IGI:MGI.
DR GO; GO:0050680; P:negative regulation of epithelial cell proliferation; ISO:MGI.
DR GO; GO:0050728; P:negative regulation of inflammatory response; IMP:BHF-UCL.
DR GO; GO:0033147; P:negative regulation of intracellular estrogen receptor signaling pathway; ISO:MGI.
DR GO; GO:0043524; P:negative regulation of neuron apoptotic process; IMP:BHF-UCL.
DR GO; GO:0045665; P:negative regulation of neuron differentiation; IGI:MGI.
DR GO; GO:0031333; P:negative regulation of protein-containing complex assembly; ISO:MGI.
DR GO; GO:0000122; P:negative regulation of transcription by RNA polymerase II; IMP:BHF-UCL.
DR GO; GO:0045892; P:negative regulation of transcription, DNA-templated; ISS:UniProtKB.
DR GO; GO:0001755; P:neural crest cell migration; IGI:MGI.
DR GO; GO:0030182; P:neuron differentiation; IMP:MGI.
DR GO; GO:0048663; P:neuron fate commitment; IGI:MGI.
DR GO; GO:0048665; P:neuron fate specification; IMP:BHF-UCL.
DR GO; GO:0003151; P:outflow tract morphogenesis; IGI:BHF-UCL.
DR GO; GO:0003148; P:outflow tract septum morphogenesis; IGI:BHF-UCL.
DR GO; GO:0031016; P:pancreas development; IMP:MGI.
DR GO; GO:0048936; P:peripheral nervous system neuron axonogenesis; IMP:BHF-UCL.
DR GO; GO:0048935; P:peripheral nervous system neuron development; IGI:MGI.
DR GO; GO:0060037; P:pharyngeal system development; IGI:BHF-UCL.
DR GO; GO:0021983; P:pituitary gland development; IMP:MGI.
DR GO; GO:0045766; P:positive regulation of angiogenesis; IMP:BHF-UCL.
DR GO; GO:0045597; P:positive regulation of cell differentiation; IMP:UniProtKB.
DR GO; GO:0008284; P:positive regulation of cell population proliferation; IDA:MGI.
DR GO; GO:0043388; P:positive regulation of DNA binding; IDA:MGI.
DR GO; GO:0010718; P:positive regulation of epithelial to mesenchymal transition; ISO:MGI.
DR GO; GO:0071657; P:positive regulation of granulocyte colony-stimulating factor production; IDA:BHF-UCL.
DR GO; GO:0032725; P:positive regulation of granulocyte macrophage colony-stimulating factor production; IDA:BHF-UCL.
DR GO; GO:0035066; P:positive regulation of histone acetylation; IMP:CACAO.
DR GO; GO:0032024; P:positive regulation of insulin secretion; ISO:MGI.
DR GO; GO:0032729; P:positive regulation of interferon-gamma production; IDA:BHF-UCL.
DR GO; GO:0032730; P:positive regulation of interleukin-1 alpha production; IDA:BHF-UCL.
DR GO; GO:0032731; P:positive regulation of interleukin-1 beta production; IDA:BHF-UCL.
DR GO; GO:0032735; P:positive regulation of interleukin-12 production; IDA:BHF-UCL.
DR GO; GO:0032755; P:positive regulation of interleukin-6 production; IDA:BHF-UCL.
DR GO; GO:1901258; P:positive regulation of macrophage colony-stimulating factor production; IDA:BHF-UCL.
DR GO; GO:0045944; P:positive regulation of transcription by RNA polymerase II; IDA:MGI.
DR GO; GO:0032760; P:positive regulation of tumor necrosis factor production; IDA:BHF-UCL.
DR GO; GO:0042531; P:positive regulation of tyrosine phosphorylation of STAT protein; IDA:MGI.
DR GO; GO:0010575; P:positive regulation of vascular endothelial growth factor production; IDA:BHF-UCL.
DR GO; GO:0010468; P:regulation of gene expression; IMP:MGI.
DR GO; GO:0086091; P:regulation of heart rate by cardiac conduction; IMP:BHF-UCL.
DR GO; GO:0003266; P:regulation of secondary heart field cardioblast proliferation; IMP:MGI.
DR GO; GO:0006357; P:regulation of transcription by RNA polymerase II; IBA:GO_Central.
DR GO; GO:0006355; P:regulation of transcription, DNA-templated; ISO:MGI.
DR GO; GO:0031290; P:retinal ganglion cell axon guidance; IMP:MGI.
DR GO; GO:0003139; P:secondary heart field specification; ISO:MGI.
DR GO; GO:0048880; P:sensory system development; IMP:BHF-UCL.
DR GO; GO:0060931; P:sinoatrial node cell development; IMP:BHF-UCL.
DR GO; GO:0021520; P:spinal cord motor neuron cell fate specification; IGI:MGI.
DR GO; GO:0021522; P:spinal cord motor neuron differentiation; IMP:MGI.
DR GO; GO:0048863; P:stem cell differentiation; IMP:MGI.
DR GO; GO:0006366; P:transcription by RNA polymerase II; IDA:MGI.
DR GO; GO:0021559; P:trigeminal nerve development; IMP:BHF-UCL.
DR GO; GO:0055010; P:ventricular cardiac muscle tissue morphogenesis; IGI:BHF-UCL.
DR GO; GO:0021524; P:visceral motor neuron differentiation; IGI:MGI.
DR CDD; cd00086; homeodomain; 1.
DR IDEAL; IID50047; -.
DR InterPro; IPR009057; Homeobox-like_sf.
DR InterPro; IPR017970; Homeobox_CS.
DR InterPro; IPR001356; Homeobox_dom.
DR InterPro; IPR001781; Znf_LIM.
DR Pfam; PF00046; Homeodomain; 1.
DR Pfam; PF00412; LIM; 2.
DR SMART; SM00389; HOX; 1.
DR SMART; SM00132; LIM; 2.
DR SUPFAM; SSF46689; SSF46689; 1.
DR PROSITE; PS00027; HOMEOBOX_1; 1.
DR PROSITE; PS50071; HOMEOBOX_2; 1.
DR PROSITE; PS00478; LIM_DOMAIN_1; 2.
DR PROSITE; PS50023; LIM_DOMAIN_2; 2.
PE 1: Evidence at protein level;
KW 3D-structure; Activator; Alternative splicing; Developmental protein;
KW Differentiation; DNA-binding; Homeobox; LIM domain; Metal-binding; Nucleus;
KW Phosphoprotein; Reference proteome; Repeat; Transcription;
KW Transcription regulation; Zinc.
FT CHAIN 1..349
FT /note="Insulin gene enhancer protein ISL-1"
FT /id="PRO_0000075748"
FT DOMAIN 17..70
FT /note="LIM zinc-binding 1"
FT /evidence="ECO:0000255|PROSITE-ProRule:PRU00125"
FT DOMAIN 79..133
FT /note="LIM zinc-binding 2"
FT /evidence="ECO:0000255|PROSITE-ProRule:PRU00125"
FT DNA_BIND 181..240
FT /note="Homeobox"
FT /evidence="ECO:0000255|PROSITE-ProRule:PRU00108"
FT REGION 262..291
FT /note="LIM-binding domain (LID)"
FT REGION 312..349
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT COMPBIAS 312..341
FT /note="Polar residues"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT VAR_SEQ 256..278
FT /note="Missing (in isoform 2)"
FT /evidence="ECO:0000303|PubMed:14664703"
FT /id="VSP_010338"
FT TURN 18..20
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 26..32
FT /evidence="ECO:0007829|PDB:4JCJ"
FT TURN 33..35
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 36..38
FT /evidence="ECO:0007829|PDB:4JCJ"
FT HELIX 40..42
FT /evidence="ECO:0007829|PDB:4JCJ"
FT TURN 46..48
FT /evidence="ECO:0007829|PDB:4JCJ"
FT HELIX 52..54
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 55..61
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 64..66
FT /evidence="ECO:0007829|PDB:4JCJ"
FT HELIX 68..74
FT /evidence="ECO:0007829|PDB:4JCJ"
FT TURN 80..82
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 91..95
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 98..101
FT /evidence="ECO:0007829|PDB:4JCJ"
FT HELIX 102..104
FT /evidence="ECO:0007829|PDB:4JCJ"
FT TURN 108..110
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 119..122
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 127..129
FT /evidence="ECO:0007829|PDB:4JCJ"
FT TURN 130..132
FT /evidence="ECO:0007829|PDB:4JCJ"
FT STRAND 145..148
FT /evidence="ECO:0007829|PDB:2RGT"
SQ SEQUENCE 349 AA; 39036 MW; 4DB82FD09154F404 CRC64;
MGDMGDPPKK KRLISLCVGC GNQIHDQYIL RVSPDLEWHA ACLKCAECNQ YLDESCTCFV
RDGKTYCKRD YIRLYGIKCA KCSIGFSKND FVMRARSKVY HIECFRCVAC SRQLIPGDEF
ALREDGLFCR ADHDVVERAS LGAGDPLSPL HPARPLQMAA EPISARQPAL RPHVHKQPEK
TTRVRTVLNE KQLHTLRTCY AANPRPDALM KEQLVEMTGL SPRVIRVWFQ NKRCKDKKRS
IMMKQLQQQQ PNDKTNIQGM TGTPMVAASP ERHDGGLQAN PVEVQSYQPP WKVLSDFALQ
SDIDQPAFQQ LVNFSEGGPG SNSTGSEVAS MSSQLPDTPN SMVASPIEA
