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Class E basic helix-loop-helix protein 22 (bHLHe22) (Class B basic helix-loop-helix protein 5) (bHLHb5) (Protein BETA3)

 BHE22_MOUSE             Reviewed;         355 AA.
Q8C6A8; Q9JL05;
06-FEB-2007, integrated into UniProtKB/Swiss-Prot.
01-MAR-2003, sequence version 1.
25-OCT-2017, entry version 108.
RecName: Full=Class E basic helix-loop-helix protein 22;
Short=bHLHe22;
AltName: Full=Class B basic helix-loop-helix protein 5;
Short=bHLHb5;
AltName: Full=Protein BETA3;
Name=Bhlhe22; Synonyms=Bhlhb5;
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 TISSUE SPECIFICITY.
PubMed=12213201; DOI=10.1006/geno.2002.6833;
Xu Z.-P., Dutra A., Stellrecht C.M., Wu C., Piatigorsky J.,
Saunders G.F.;
"Functional and structural characterization of the human gene BHLHB5,
encoding a basic helix-loop-helix transcription factor.";
Genomics 80:311-318(2002).
[2]
NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
STRAIN=C57BL/6J; TISSUE=Head;
PubMed=16141072; DOI=10.1126/science.1112014;
Carninci P., Kasukawa T., Katayama S., Gough J., Frith M.C., Maeda N.,
Oyama R., Ravasi T., Lenhard B., Wells C., Kodzius R., Shimokawa K.,
Bajic V.B., Brenner S.E., Batalov S., Forrest A.R., Zavolan M.,
Davis M.J., Wilming L.G., Aidinis V., Allen J.E.,
Ambesi-Impiombato A., Apweiler R., Aturaliya R.N., Bailey T.L.,
Bansal M., Baxter L., Beisel K.W., Bersano T., Bono H., Chalk A.M.,
Chiu K.P., Choudhary V., Christoffels A., Clutterbuck D.R.,
Crowe M.L., Dalla E., Dalrymple B.P., de Bono B., Della Gatta G.,
di Bernardo D., Down T., Engstrom P., Fagiolini M., Faulkner G.,
Fletcher C.F., Fukushima T., Furuno M., Futaki S., Gariboldi M.,
Georgii-Hemming P., Gingeras T.R., Gojobori T., Green R.E.,
Gustincich S., Harbers M., Hayashi Y., Hensch T.K., Hirokawa N.,
Hill D., Huminiecki L., Iacono M., Ikeo K., Iwama A., Ishikawa T.,
Jakt M., Kanapin A., Katoh M., Kawasawa Y., Kelso J., Kitamura H.,
Kitano H., Kollias G., Krishnan S.P., Kruger A., Kummerfeld S.K.,
Kurochkin I.V., Lareau L.F., Lazarevic D., Lipovich L., Liu J.,
Liuni S., McWilliam S., Madan Babu M., Madera M., Marchionni L.,
Matsuda H., Matsuzawa S., Miki H., Mignone F., Miyake S., Morris K.,
Mottagui-Tabar S., Mulder N., Nakano N., Nakauchi H., Ng P.,
Nilsson R., Nishiguchi S., Nishikawa S., Nori F., Ohara O.,
Okazaki Y., Orlando V., Pang K.C., Pavan W.J., Pavesi G., Pesole G.,
Petrovsky N., Piazza S., Reed J., Reid J.F., Ring B.Z., Ringwald M.,
Rost B., Ruan Y., Salzberg S.L., Sandelin A., Schneider C.,
Schoenbach C., Sekiguchi K., Semple C.A., Seno S., Sessa L., Sheng Y.,
Shibata Y., Shimada H., Shimada K., Silva D., Sinclair B.,
Sperling S., Stupka E., Sugiura K., Sultana R., Takenaka Y., Taki K.,
Tammoja K., Tan S.L., Tang S., Taylor M.S., Tegner J., Teichmann S.A.,
Ueda H.R., van Nimwegen E., Verardo R., Wei C.L., Yagi K.,
Yamanishi H., Zabarovsky E., Zhu S., Zimmer A., Hide W., Bult C.,
Grimmond S.M., Teasdale R.D., Liu E.T., Brusic V., Quackenbush J.,
Wahlestedt C., Mattick J.S., Hume D.A., Kai C., Sasaki D., Tomaru Y.,
Fukuda S., Kanamori-Katayama M., Suzuki M., Aoki J., Arakawa T.,
Iida J., Imamura K., Itoh M., Kato T., Kawaji H., Kawagashira N.,
Kawashima T., Kojima M., Kondo S., Konno H., Nakano K., Ninomiya N.,
Nishio T., Okada M., Plessy C., Shibata K., Shiraki T., Suzuki S.,
Tagami M., Waki K., Watahiki A., Okamura-Oho Y., Suzuki H., Kawai J.,
Hayashizaki Y.;
"The transcriptional landscape of the mammalian genome.";
Science 309:1559-1563(2005).
[3]
NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
STRAIN=C57BL/6J; TISSUE=Brain;
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).
[4]
DEVELOPMENTAL STAGE.
PubMed=14643684; DOI=10.1016/S1567-133X(03)00135-2;
Brunelli S., Innocenzi A., Cossu G.;
"Bhlhb5 is expressed in the CNS and sensory organs during mouse
embryonic development.";
Gene Expr. Patterns 3:755-759(2003).
[5]
FUNCTION, AND DEVELOPMENTAL STAGE.
PubMed=17092954; DOI=10.1242/dev.02664;
Feng L., Xie X., Joshi P.S., Yang Z., Shibasaki K., Chow R.L., Gan L.;
"Requirement for Bhlhb5 in the specification of amacrine and cone
bipolar subtypes in mouse retina.";
Development 133:4815-4825(2006).
[6]
FUNCTION, DEVELOPMENTAL STAGE, AND DISRUPTION PHENOTYPE.
PubMed=18957218; DOI=10.1016/j.neuron.2008.08.006;
Joshi P.S., Molyneaux B.J., Feng L., Xie X., Macklis J.D., Gan L.;
"Bhlhb5 regulates the postmitotic acquisition of area identities in
layers II-V of the developing neocortex.";
Neuron 60:258-272(2008).
[7]
FUNCTION, DEVELOPMENTAL STAGE, AND DISRUPTION PHENOTYPE.
PubMed=20346763; DOI=10.1016/j.neuron.2010.02.025;
Ross S.E., Mardinly A.R., McCord A.E., Zurawski J., Cohen S., Jung C.,
Hu L., Mok S.I., Shah A., Savner E.M., Tolias C., Corfas R., Chen S.,
Inquimbert P., Xu Y., McInnes R.R., Rice F.L., Corfas G., Ma Q.,
Woolf C.J., Greenberg M.E.;
"Loss of inhibitory interneurons in the dorsal spinal cord and
elevated itch in Bhlhb5 mutant mice.";
Neuron 65:886-898(2010).
[8]
FUNCTION, AND INTERACTION WITH PRDM8.
PubMed=22284184; DOI=10.1016/j.neuron.2011.09.035;
Ross S.E., McCord A.E., Jung C., Atan D., Mok S.I., Hemberg M.,
Kim T.K., Salogiannis J., Hu L., Cohen S., Lin Y., Harrar D.,
McInnes R.R., Greenberg M.E.;
"Bhlhb5 and Prdm8 form a repressor complex involved in neuronal
circuit assembly.";
Neuron 73:292-303(2012).
-!- FUNCTION: Inhibits DNA binding of TCF3/E47 homodimers and TCF3
(E47)/NEUROD1 heterodimers and acts as a strong repressor of
Neurod1 and Myod-responsive genes, probably by heterodimerization
with class a basic helix-loop-helix factors. Despite the presence
of an intact basic domain, does not bind to DNA (By similarity).
In the brain, may function as an area-specific transcription
factor that regulates the postmitotic acquisition of area
identities and elucidate the genetic hierarchy between progenitors
and postmitotic neurons driving neocortical arealization. May be
required for the survival of a specific population of inhibitory
neurons in the superficial laminae of the spinal chord dorsal horn
that may regulate pruritis. Seems to play a crucial role in the
retinogenesis, in the specification of amacrine and bipolar
subtypes. Forms with PRDM8 a transcriptional repressor complex
controlling genes involved in neural development and neuronal
differentiation (PubMed:22284184). {ECO:0000250,
ECO:0000269|PubMed:17092954, ECO:0000269|PubMed:18957218,
ECO:0000269|PubMed:20346763, ECO:0000269|PubMed:22284184}.
-!- SUBUNIT: Interacts with PRDM8. {ECO:0000269|PubMed:22284184}.
-!- SUBCELLULAR LOCATION: Nucleus {ECO:0000305}.
-!- TISSUE SPECIFICITY: Brain-specific, with the highest expression in
the cerebellum. {ECO:0000269|PubMed:12213201}.
-!- DEVELOPMENTAL STAGE: Expressed at 11.5 dpc within the neuroblast
layer (NBL) of the central retina. As retinogenesis progressed
from central to peripheral retina from 12 dpc to 15.5 dpc,
expression expanded to the entire retina with the majority of
Bhlhb5+ cells being detected in the proliferating NBL. From 17.5
dpc to birth (P0), expression became restricted to the ganglion
cell layer (GCL) and to the inner boundary of the NBL (INL; inner
nuclear layer), presumably the newly formed ACL (amacrine cells).
Predominantly expressed in post-mitotic cells in the developing
retina. Expressed from 9.5 dpc in the neural tube, restricted to
longitudinal ventral columns of neurons, extending from the
hindbrain to the caudal spinal cord. In the developing cortex, at
12.5 dpc, expressed in the nascent cortical plate of the dorsal
telencephalon (at protein level). During peak production of deep
layer neurons between 12.5 and 13.5 dpc, restricted to
postmigrational neurons, with no expression in the proliferative
ventricular zone (VZ) or in migrating neurons. Between 15.5 and
17.5 dpc, when superficial layer neurons are generated, strongly
expressed in the cortical plate and weakly in presumptive
migrating neurons and in the subventricular zone (SVZ). Does not
colocalize with proliferating progenitors in S or M phase in
either the VZ or the SVZ; restricted to postmitotic glutamatergic
projection neurons during neurogenesis. Not detected in cortical
GABAergic interneurons or their extracortical sites of genesis,
the medial and caudal ganglionic eminences (at protein level).
Postnatally, down-regulation begins at the junction of the
cingulate cortex and neocortex; by postnatal day 4 (P4), down-
regulation well into the neocortex is observed anteriorly, with
the exception of the most superficial layer. At P4, expressed in
neocortical layers II, III, IV, and V. Within layer VI, expressed
in only very rare scattered TBR1 negative neurons. Down-regulation
continues from medial to lateral, exhibiting a markedly reduced
expression by P14. Expression varies strikingly along the A-P axis
with a precipitous decrease in the rostral cortical plate. At 12.5
dpc, highly expressed medially in the cingulate cortex with weak
expression in the rest of the cortex. At 15.5 dpc, expressed in a
high caudomedial to a low rostrolateral gradient. Between 15.5dpc
and P0, expression increases laterally and the gradient gradually
transforms into a sharp border between the presumptive rostral
motor and sensory domains. During the first postnatal week, there
is a further transformation from homogeneous expression across
sensory cortex into discrete areas of high expression coincident
with the primary sensory areas. At P4 and P7, expressed in primary
visual cortex, primary auditory cortex and distinct primary
somatosensory representations, including the vibrissal barrel
field. In the spinal chord, transiently expressed in V1, V2, and
dI6 interneurons at 10.5 dpc. Also observed in a subpopulation of
late-born neurons that migrate to the superficial layers of the
dorsal horn. Expression starts shortly after the neurons exit
mitosis at 13.5 dpc and persisting for up to 2 weeks postnatally.
At birth, about one-third of dorsal horn neurons expressing the
protein are excitatory and two-thirds inhibitory. Also expressed
in the developing eye and hair follicles, in the epithelial layer
of the cochlea in the developing inner, and in the nasal
epithelium. At 16.5 dpc, expressed outside the CNS, in particular
in all sensory organs; in the nasal pits, transcripts can be
detected in the olfactory epithelium. In the developing eye it can
be found in the inner and outer retinal layer, and it is also
detectable in the sensory layer of the cochlea in the developing
inner ear. In addition, expression is found in the developing hair
follicles, both in the epithelial component and in the dermal
papilla, and in the skin. {ECO:0000269|PubMed:14643684,
ECO:0000269|PubMed:17092954, ECO:0000269|PubMed:18957218,
ECO:0000269|PubMed:20346763}.
-!- DISRUPTION PHENOTYPE: Null mice exhibits aberrant expression of
brain area-specific genes and structural organization in the
somatosensory and caudal motor cortices. In somatosensory cortex,
vibrissal barrels display postsynaptic disorganization. In caudal
motor cortex, anomalous differentiation of corticospinal motor
neurons is observed, accompanied by failure of corticospinal tract
formation. Mice also develop self-inflicted skin lesions and show
significantly enhanced scratching responses to pruritic agents,
due to the selective loss of a subset of spinal chord inhibitory
interneurons. {ECO:0000269|PubMed:18957218,
ECO:0000269|PubMed:20346763}.
-----------------------------------------------------------------------
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EMBL; AF218865; AAF32324.1; -; mRNA.
EMBL; AK076228; BAC36262.1; -; mRNA.
EMBL; BC053007; AAH53007.1; -; mRNA.
CCDS; CCDS17253.1; -.
RefSeq; NP_067535.3; NM_021560.4.
UniGene; Mm.149938; -.
ProteinModelPortal; Q8C6A8; -.
SMR; Q8C6A8; -.
STRING; 10090.ENSMUSP00000026120; -.
BindingDB; Q8C6A8; -.
PhosphoSitePlus; Q8C6A8; -.
PaxDb; Q8C6A8; -.
PRIDE; Q8C6A8; -.
GeneID; 59058; -.
KEGG; mmu:59058; -.
UCSC; uc008orl.2; mouse.
CTD; 27319; -.
MGI; MGI:1930001; Bhlhe22.
eggNOG; KOG3898; Eukaryota.
eggNOG; ENOG410ZDH5; LUCA.
HOGENOM; HOG000060094; -.
InParanoid; Q8C6A8; -.
KO; K09086; -.
PhylomeDB; Q8C6A8; -.
TreeFam; TF322733; -.
PRO; PR:Q8C6A8; -.
Proteomes; UP000000589; Unplaced.
GO; GO:0005634; C:nucleus; IDA:MGI.
GO; GO:0003682; F:chromatin binding; IDA:MGI.
GO; GO:0042803; F:protein homodimerization activity; IPI:MGI.
GO; GO:0000981; F:RNA polymerase II transcription factor activity, sequence-specific DNA binding; IMP:MGI.
GO; GO:0021960; P:anterior commissure morphogenesis; IMP:MGI.
GO; GO:0021952; P:central nervous system projection neuron axonogenesis; IMP:MGI.
GO; GO:0021796; P:cerebral cortex regionalization; IMP:MGI.
GO; GO:0021540; P:corpus callosum morphogenesis; IMP:MGI.
GO; GO:0021957; P:corticospinal tract morphogenesis; IMP:MGI.
GO; GO:0045892; P:negative regulation of transcription, DNA-templated; IDA:MGI.
GO; GO:0030182; P:neuron differentiation; IMP:MGI.
GO; GO:0060042; P:retina morphogenesis in camera-type eye; IMP:MGI.
GO; GO:0060040; P:retinal bipolar neuron differentiation; IMP:MGI.
GO; GO:0006351; P:transcription, DNA-templated; IEA:UniProtKB-KW.
CDD; cd00083; HLH; 1.
Gene3D; 4.10.280.10; -; 1.
InterPro; IPR011598; bHLH_dom.
InterPro; IPR032654; Bhlhe22.
InterPro; IPR036638; HLH_DNA-bd_sf.
PANTHER; PTHR19290:SF52; PTHR19290:SF52; 1.
Pfam; PF00010; HLH; 1.
SMART; SM00353; HLH; 1.
SUPFAM; SSF47459; SSF47459; 1.
PROSITE; PS50888; BHLH; 1.
1: Evidence at protein level;
Complete proteome; Neurogenesis; Nucleus; Reference proteome;
Repressor; Transcription; Transcription regulation.
CHAIN 1 355 Class E basic helix-loop-helix protein
22.
/FTId=PRO_0000274286.
DOMAIN 216 270 bHLH. {ECO:0000255|PROSITE-
ProRule:PRU00981}.
COMPBIAS 56 59 Poly-Ser.
COMPBIAS 71 206 Gly-rich.
COMPBIAS 289 331 Ala-rich.
CONFLICT 75 75 R -> L (in Ref. 1; AAF32324).
{ECO:0000305}.
CONFLICT 149 149 L -> R (in Ref. 1; AAF32324).
{ECO:0000305}.
CONFLICT 166 166 R -> G (in Ref. 1; AAF32324).
{ECO:0000305}.
SEQUENCE 355 AA; 35217 MW; CA221A9169FCB897 CRC64;
MERGLHLGAA AASEDDLFLH KSLGTSAAKR LEAAFRSTPP GMDLSLAPPT RERPASSSSP
LGCFEPADPE GAGLRLPPPG GGGGASGGGG GVSVPGLLVG SAGVGGEPSL SSLPAGAALC
LKYGESAGRG SVAESSGGEQ SPDDDSDGLC ELVLRAGGPD PRASPRAGGG SAKVAEGCSN
AHLHGGSGLP PGGPTSGGGS GGGGGGSSKK SKEQKALRLN INARERRRMH DLNDALDELR
AVIPYAHSPS VRKLSKIATL LLAKNYILMQ AQALEEMRRL VAYLNQGQAI SAASLPSSAA
AAAAAAALHP ALGAYEQAAG YPFSAGLPPA ASCPEKCALF NSVSSSLCKQ CTEKP


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