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Homeobox protein MIXL1 (Homeodomain protein MIX) (mMix) (MIX1 homeobox-like protein 1) (Mix.1 homeobox-like protein)

 MIXL1_MOUSE             Reviewed;         231 AA.
Q9WUI0; Q9QZ61;
13-NOV-2007, integrated into UniProtKB/Swiss-Prot.
01-NOV-1999, sequence version 1.
27-SEP-2017, entry version 132.
RecName: Full=Homeobox protein MIXL1;
AltName: Full=Homeodomain protein MIX;
Short=mMix;
AltName: Full=MIX1 homeobox-like protein 1;
AltName: Full=Mix.1 homeobox-like protein;
Name=Mixl1; Synonyms=Mix, Mml;
Mus musculus (Mouse).
Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi;
Mammalia; Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha;
Muroidea; Muridae; Murinae; Mus; Mus.
NCBI_TaxID=10090;
[1]
NUCLEOTIDE SEQUENCE [MRNA], AND DEVELOPMENTAL STAGE.
PubMed=10495285; DOI=10.1016/S0925-4773(99)00135-5;
Pearce J.J.H., Evans M.J.;
"Mml, a mouse Mix-like gene expressed in the primitive streak.";
Mech. Dev. 87:189-192(1999).
[2]
NUCLEOTIDE SEQUENCE [MRNA].
STRAIN=129/Sv; TISSUE=Embryo;
PubMed=11084649;
DOI=10.1002/1097-0177(2000)9999:9999<::AID-DVDY1070>3.0.CO;2-O;
Robb L., Hartley L., Begley C.G., Brodnicki T.C., Copeland N.G.,
Gilbert D.J., Jenkins N.A., Elefanty A.G.;
"Cloning, expression analysis, and chromosomal localization of murine
and human homologues of a Xenopus mix gene.";
Dev. Dyn. 219:497-504(2000).
[3]
NUCLEOTIDE SEQUENCE [GENOMIC DNA], NUCLEOTIDE SEQUENCE [MRNA] OF
13-231, AND DNA-BINDING.
STRAIN=129/SvJ;
PubMed=12095687; DOI=10.1016/S0378-1119(02)00590-5;
Sahr K., Dias D.C., Sanchez R., Chen D., Chen S.W., Gudas L.J.,
Baron M.H.;
"Structure, upstream promoter region, and functional domains of a
mouse and human Mix paired-like homeobox gene.";
Gene 291:135-147(2002).
[4]
NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
PubMed=15489334; DOI=10.1101/gr.2596504;
The MGC Project Team;
"The status, quality, and expansion of the NIH full-length cDNA
project: the Mammalian Gene Collection (MGC).";
Genome Res. 14:2121-2127(2004).
[5]
FUNCTION, AND DISRUPTION PHENOTYPE.
PubMed=12117810;
Hart A.H., Hartley L., Sourris K., Stadler E.S., Li R., Stanley E.G.,
Tam P.P.L., Elefanty A.G., Robb L.;
"Mixl1 is required for axial mesendoderm morphogenesis and patterning
in the murine embryo.";
Development 129:3597-3608(2002).
[6]
SUBCELLULAR LOCATION, TISSUE SPECIFICITY, AND DEVELOPMENTAL STAGE.
PubMed=12619131; DOI=10.1002/dvdy.10263;
Mohn D., Chen S.W., Dias D.C., Weinstein D.C., Dyer M.A., Sahr K.,
Ducker C.E., Zahradka E., Keller G., Zaret K.S., Gudas L.J.,
Baron M.H.;
"Mouse Mix gene is activated early during differentiation of ES and F9
stem cells and induces endoderm in frog embryos.";
Dev. Dyn. 226:446-459(2003).
[7]
INDUCTION.
PubMed=15982639; DOI=10.1016/j.bbrc.2005.06.044;
Hart A.H., Willson T.A., Wong M., Parker K., Robb L.;
"Transcriptional regulation of the homeobox gene Mixl1 by TGF-beta and
FoxH1.";
Biochem. Biophys. Res. Commun. 333:1361-1369(2005).
[8]
FUNCTION.
PubMed=15673572; DOI=10.1242/dev.01657;
Ng E.S., Azzola L., Sourris K., Robb L., Stanley E.G., Elefanty A.G.;
"The primitive streak gene Mixl1 is required for efficient
haematopoiesis and BMP4-induced ventral mesoderm patterning in
differentiating ES cells.";
Development 132:873-884(2005).
[9]
TISSUE SPECIFICITY.
PubMed=16433620; DOI=10.1089/scd.2005.14.656;
Mossman A.K., Sourris K., Ng E., Stanley E.G., Elefanty A.G.;
"Mixl1 and oct4 proteins are transiently co-expressed in
differentiating mouse and human embryonic stem cells.";
Stem Cells Dev. 14:656-663(2005).
[10]
FUNCTION.
PubMed=16403910; DOI=10.1182/blood-2005-10-4120;
Willey S., Ayuso-Sacido A., Zhang H., Fraser S.T., Sahr K.E.,
Adlam M.J., Kyba M., Daley G.Q., Keller G., Baron M.H.;
"Acceleration of mesoderm development and expansion of hematopoietic
progenitors in differentiating ES cells by the mouse Mix-like
homeodomain transcription factor.";
Blood 107:3122-3130(2006).
[11]
FUNCTION.
PubMed=17060613; DOI=10.1073/pnas.0607776103;
Glaser S., Metcalf D., Wu L., Hart A.H., DiRago L., Mifsud S.,
D'Amico A., Dagger S., Campo C., Chan A.C., Izon D.J., Robb L.;
"Enforced expression of the homeobox gene Mixl1 impairs hematopoietic
differentiation and results in acute myeloid leukemia.";
Proc. Natl. Acad. Sci. U.S.A. 103:16460-16465(2006).
[12]
FUNCTION.
PubMed=17151016; DOI=10.1242/dev.02724;
Tam P.P.L., Khoo P.-L., Lewis S.L., Bildsoe H., Wong N., Tsang T.E.,
Gad J.M., Robb L.;
"Sequential allocation and global pattern of movement of the
definitive endoderm in the mouse embryo during gastrulation.";
Development 134:251-260(2007).
[13]
INDUCTION.
PubMed=17568773; DOI=10.1038/sj.emboj.7601753;
Izzi L., Silvestri C., von Both I., Labbe E., Zakin L., Wrana J.L.,
Attisano L.;
"Foxh1 recruits Gsc to negatively regulate Mixl1 expression during
early mouse development.";
EMBO J. 26:3132-3143(2007).
[14]
FUNCTION.
PubMed=17446562; DOI=10.1634/stemcells.2006-0681;
Izumi N., Era T., Akimaru H., Yasunaga M., Nishikawa S.;
"Dissecting the molecular hierarchy for mesendoderm differentiation
through a combination of embryonic stem cell culture and RNA
interference.";
Stem Cells 25:1664-1674(2007).
-!- FUNCTION: Transcription factor that play a central role in proper
axial mesendoderm morphogenesis and endoderm formation. Required
for efficient differentiation of cells from the primitive streak
stage to blood, by acting early in the recruitment and/or
expansion of mesodermal progenitors to the hemangioblastic and
hematopoietic lineages. Also involved in the morphogenesis of the
heart and the gut during embryogenesis. Acts as a negative
regulator of brachyury expression. {ECO:0000269|PubMed:12117810,
ECO:0000269|PubMed:15673572, ECO:0000269|PubMed:16403910,
ECO:0000269|PubMed:17060613, ECO:0000269|PubMed:17151016,
ECO:0000269|PubMed:17446562}.
-!- INTERACTION:
Self; NbExp=2; IntAct=EBI-13637214, EBI-13637214;
P20293:T; NbExp=7; IntAct=EBI-13637214, EBI-13637160;
-!- SUBCELLULAR LOCATION: Nucleus {ECO:0000255|PROSITE-
ProRule:PRU00108, ECO:0000269|PubMed:12619131}.
-!- TISSUE SPECIFICITY: Expressed in the primitive streak of the
gastrulating embryo, and marks cells destined to form mesoderm and
endoderm. Present in differentiating embryonic stem cells (at
protein level). {ECO:0000269|PubMed:12619131,
ECO:0000269|PubMed:16433620}.
-!- DEVELOPMENTAL STAGE: Initially expressed in visceral endoderm and
becomes restricted to primitive streak and nascent mesoderm at
gastrulation. This includes the hemangioblast, a precursor of
hematopoietic and vascular stem cells. At E5.5, it is expressed
symmetrically in the visceral endoderm but by E6.0 this expression
is noticeably asymmetric. At E6.5, expression is restricted to the
nascent primitive streak and persists in the primitive streak
through E7.5-E9.5, marking those cells fated to form extra-
embryonic and lateral mesoderm. {ECO:0000269|PubMed:10495285,
ECO:0000269|PubMed:12619131}.
-!- INDUCTION: By TGF-beta. Regulated by Nodal in mesendoderm
morphogenesis. Regulated by FOXH1, which acts as a negative
regulator by recruiting GSC to promoter during early development.
{ECO:0000269|PubMed:15982639, ECO:0000269|PubMed:17568773}.
-!- DISRUPTION PHENOTYPE: Mice display defects in mesendoderm
morphogenesis and patterning during development. Embryos show a
marked thickening of the primitive streak. By the early somite
stage, embryonic development is arrested, with the formation of
abnormal head folds, foreshortened body axis, absence of heart
tube and gut, deficient paraxial mesoderm, and an enlarged midline
tissue mass that replaces the notochord. Development of extra-
embryonic structures is generally normal except that the allantois
is often disproportionately large for the size of the mutant
embryo. {ECO:0000269|PubMed:12117810}.
-!- SIMILARITY: Belongs to the paired homeobox family. {ECO:0000305}.
-----------------------------------------------------------------------
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EMBL; AF135063; AAD25543.1; -; mRNA.
EMBL; AF154573; AAK59867.1; -; mRNA.
EMBL; AF201959; AAF08314.1; -; mRNA.
EMBL; AF447591; AAM77218.1; -; Genomic_DNA.
EMBL; BC099876; AAH99876.1; -; mRNA.
EMBL; BC099877; AAH99877.1; -; mRNA.
CCDS; CCDS15571.1; -.
RefSeq; NP_038757.1; NM_013729.3.
RefSeq; XP_006496934.1; XM_006496871.1.
UniGene; Mm.103647; -.
ProteinModelPortal; Q9WUI0; -.
SMR; Q9WUI0; -.
BioGrid; 205138; 10.
IntAct; Q9WUI0; 6.
STRING; 10090.ENSMUSP00000027778; -.
PhosphoSitePlus; Q9WUI0; -.
PaxDb; Q9WUI0; -.
PeptideAtlas; Q9WUI0; -.
PRIDE; Q9WUI0; -.
Ensembl; ENSMUST00000027778; ENSMUSP00000027778; ENSMUSG00000026497.
GeneID; 27217; -.
KEGG; mmu:27217; -.
UCSC; uc007dwp.1; mouse.
CTD; 83881; -.
MGI; MGI:1351322; Mixl1.
eggNOG; KOG0849; Eukaryota.
eggNOG; ENOG410XS01; LUCA.
GeneTree; ENSGT00860000133740; -.
HOGENOM; HOG000113594; -.
HOVERGEN; HBG106595; -.
InParanoid; Q9WUI0; -.
OMA; TMYPDIH; -.
OrthoDB; EOG091G15I5; -.
PhylomeDB; Q9WUI0; -.
TreeFam; TF334098; -.
PRO; PR:Q9WUI0; -.
Proteomes; UP000000589; Chromosome 1.
Bgee; ENSMUSG00000026497; -.
CleanEx; MM_MIXL1; -.
Genevisible; Q9WUI0; MM.
GO; GO:0000790; C:nuclear chromatin; IDA:BHF-UCL.
GO; GO:0005634; C:nucleus; IDA:BHF-UCL.
GO; GO:0000987; F:core promoter proximal region sequence-specific DNA binding; IDA:BHF-UCL.
GO; GO:0042802; F:identical protein binding; IPI:IntAct.
GO; GO:0042803; F:protein homodimerization activity; IPI:BHF-UCL.
GO; GO:0000978; F:RNA polymerase II core promoter proximal region sequence-specific DNA binding; IDA:BHF-UCL.
GO; GO:0000980; F:RNA polymerase II distal enhancer sequence-specific DNA binding; IDA:BHF-UCL.
GO; GO:0001103; F:RNA polymerase II repressing transcription factor binding; IPI:BHF-UCL.
GO; GO:0001085; F:RNA polymerase II transcription factor binding; IPI:BHF-UCL.
GO; GO:0001077; F:transcriptional activator activity, RNA polymerase II core promoter proximal region sequence-specific binding; IDA:BHF-UCL.
GO; GO:0001205; F:transcriptional activator activity, RNA polymerase II distal enhancer sequence-specific binding; IDA:BHF-UCL.
GO; GO:0001190; F:transcriptional activator activity, RNA polymerase II transcription factor binding; IDA:BHF-UCL.
GO; GO:0042074; P:cell migration involved in gastrulation; IMP:MGI.
GO; GO:0048565; P:digestive tract development; IMP:MGI.
GO; GO:0007492; P:endoderm development; IMP:MGI.
GO; GO:0001706; P:endoderm formation; IMP:MGI.
GO; GO:0035987; P:endodermal cell differentiation; IDA:BHF-UCL.
GO; GO:0007369; P:gastrulation; IMP:MGI.
GO; GO:0007507; P:heart development; IMP:MGI.
GO; GO:0002244; P:hematopoietic progenitor cell differentiation; IDA:BHF-UCL.
GO; GO:0030097; P:hemopoiesis; IDA:MGI.
GO; GO:1901533; P:negative regulation of hematopoietic progenitor cell differentiation; IDA:BHF-UCL.
GO; GO:2000382; P:positive regulation of mesoderm development; IDA:BHF-UCL.
GO; GO:0045944; P:positive regulation of transcription from RNA polymerase II promoter; IDA:BHF-UCL.
CDD; cd00086; homeodomain; 1.
InterPro; IPR009057; Homeobox-like.
InterPro; IPR017970; Homeobox_CS.
InterPro; IPR001356; Homeobox_dom.
Pfam; PF00046; Homeobox; 1.
SMART; SM00389; HOX; 1.
SUPFAM; SSF46689; SSF46689; 1.
PROSITE; PS00027; HOMEOBOX_1; 1.
PROSITE; PS50071; HOMEOBOX_2; 1.
1: Evidence at protein level;
Complete proteome; Developmental protein; Differentiation;
DNA-binding; Homeobox; Nucleus; Reference proteome; Transcription;
Transcription regulation.
CHAIN 1 231 Homeobox protein MIXL1.
/FTId=PRO_0000311334.
DNA_BIND 86 145 Homeobox. {ECO:0000255|PROSITE-
ProRule:PRU00108}.
SEQUENCE 231 AA; 24844 MW; 59DA80A5E7E6EE33 CRC64;
MAAAGSQQLQ FAEGAAFPIF PAAHPGGQLL PAMRPASGLP AAPHDSRAPA ATQCFPNRDS
SPTAQTPAGL DPPGPSKGSA APSAPQRRKR TSFSSEQLQL LELVFRQTMY PDIHLRERLA
ALTLLPESRI QVWFQNRRAK SRRQSGKSFQ PLSSRRGVFL HCPAPGTEAR CLKPQLPLEA
DVNHVPDPSM TGGGVCTSGS QSFETYSSLS EDIGSKLDSW EEHIFSALGN F


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