HEN2_MOUSE
ID HEN2_MOUSE Reviewed; 135 AA.
AC Q64221; Q8BFV7;
DT 01-NOV-1997, integrated into UniProtKB/Swiss-Prot.
DT 03-OCT-2012, sequence version 2.
DT 03-AUG-2022, entry version 147.
DE RecName: Full=Helix-loop-helix protein 2;
DE Short=HEN-2;
DE AltName: Full=Nescient helix loop helix 2;
DE Short=NSCL-2;
GN Name=Nhlh2; Synonyms=Hen2;
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], TISSUE SPECIFICITY, AND DEVELOPMENTAL STAGE.
RX PubMed=1633105;
RA Gobel V., Lipkowitz S., Kozak C.A., Kirsch I.R.;
RT "NSCL-2: a basic domain helix-loop-helix gene expressed in early
RT neurogenesis.";
RL Cell Growth Differ. 3:143-148(1992).
RN [2]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RC STRAIN=C57BL/6J; TISSUE=Cerebellum, Embryo, Eye, and Head;
RX PubMed=16141072; DOI=10.1126/science.1112014;
RA Carninci P., Kasukawa T., Katayama S., Gough J., Frith M.C., Maeda N.,
RA Oyama R., Ravasi T., Lenhard B., Wells C., Kodzius R., Shimokawa K.,
RA Bajic V.B., Brenner S.E., Batalov S., Forrest A.R., Zavolan M., Davis M.J.,
RA Wilming L.G., Aidinis V., Allen J.E., Ambesi-Impiombato A., Apweiler R.,
RA Aturaliya R.N., Bailey T.L., Bansal M., Baxter L., Beisel K.W., Bersano T.,
RA Bono H., Chalk A.M., Chiu K.P., Choudhary V., Christoffels A.,
RA Clutterbuck D.R., Crowe M.L., Dalla E., Dalrymple B.P., de Bono B.,
RA Della Gatta G., di Bernardo D., Down T., Engstrom P., Fagiolini M.,
RA Faulkner G., Fletcher C.F., Fukushima T., Furuno M., Futaki S.,
RA Gariboldi M., Georgii-Hemming P., Gingeras T.R., Gojobori T., Green R.E.,
RA Gustincich S., Harbers M., Hayashi Y., Hensch T.K., Hirokawa N., Hill D.,
RA Huminiecki L., Iacono M., Ikeo K., Iwama A., Ishikawa T., Jakt M.,
RA Kanapin A., Katoh M., Kawasawa Y., Kelso J., Kitamura H., Kitano H.,
RA Kollias G., Krishnan S.P., Kruger A., Kummerfeld S.K., Kurochkin I.V.,
RA Lareau L.F., Lazarevic D., Lipovich L., Liu J., Liuni S., McWilliam S.,
RA Madan Babu M., Madera M., Marchionni L., Matsuda H., Matsuzawa S., Miki H.,
RA Mignone F., Miyake S., Morris K., Mottagui-Tabar S., Mulder N., Nakano N.,
RA Nakauchi H., Ng P., Nilsson R., Nishiguchi S., Nishikawa S., Nori F.,
RA Ohara O., Okazaki Y., Orlando V., Pang K.C., Pavan W.J., Pavesi G.,
RA Pesole G., Petrovsky N., Piazza S., Reed J., Reid J.F., Ring B.Z.,
RA Ringwald M., Rost B., Ruan Y., Salzberg S.L., Sandelin A., Schneider C.,
RA Schoenbach C., Sekiguchi K., Semple C.A., Seno S., Sessa L., Sheng Y.,
RA Shibata Y., Shimada H., Shimada K., Silva D., Sinclair B., Sperling S.,
RA Stupka E., Sugiura K., Sultana R., Takenaka Y., Taki K., Tammoja K.,
RA Tan S.L., Tang S., Taylor M.S., Tegner J., Teichmann S.A., Ueda H.R.,
RA van Nimwegen E., Verardo R., Wei C.L., Yagi K., Yamanishi H.,
RA Zabarovsky E., Zhu S., Zimmer A., Hide W., Bult C., Grimmond S.M.,
RA Teasdale R.D., Liu E.T., Brusic V., Quackenbush J., Wahlestedt C.,
RA Mattick J.S., Hume D.A., Kai C., Sasaki D., Tomaru Y., Fukuda S.,
RA Kanamori-Katayama M., Suzuki M., Aoki J., Arakawa T., Iida J., Imamura K.,
RA Itoh M., Kato T., Kawaji H., Kawagashira N., Kawashima T., Kojima M.,
RA Kondo S., Konno H., Nakano K., Ninomiya N., Nishio T., Okada M., Plessy C.,
RA Shibata K., Shiraki T., Suzuki S., Tagami M., Waki K., Watahiki A.,
RA Okamura-Oho Y., Suzuki H., Kawai J., Hayashizaki Y.;
RT "The transcriptional landscape of the mammalian genome.";
RL Science 309:1559-1563(2005).
RN [3]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC STRAIN=C57BL/6J;
RX PubMed=19468303; DOI=10.1371/journal.pbio.1000112;
RA Church D.M., Goodstadt L., Hillier L.W., Zody M.C., Goldstein S., She X.,
RA Bult C.J., Agarwala R., Cherry J.L., DiCuccio M., Hlavina W., Kapustin Y.,
RA Meric P., Maglott D., Birtle Z., Marques A.C., Graves T., Zhou S.,
RA Teague B., Potamousis K., Churas C., Place M., Herschleb J., Runnheim R.,
RA Forrest D., Amos-Landgraf J., Schwartz D.C., Cheng Z., Lindblad-Toh K.,
RA Eichler E.E., Ponting C.P.;
RT "Lineage-specific biology revealed by a finished genome assembly of the
RT mouse.";
RL PLoS Biol. 7:E1000112-E1000112(2009).
RN [4]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RA Mural R.J., Adams M.D., Myers E.W., Smith H.O., Venter J.C.;
RL Submitted (JUL-2005) to the EMBL/GenBank/DDBJ databases.
RN [5]
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RC STRAIN=C57BL/6J; TISSUE=Brain;
RX PubMed=15489334; DOI=10.1101/gr.2596504;
RG The MGC Project Team;
RT "The status, quality, and expansion of the NIH full-length cDNA project:
RT the Mammalian Gene Collection (MGC).";
RL Genome Res. 14:2121-2127(2004).
RN [6]
RP TISSUE SPECIFICITY.
RX PubMed=9011755; DOI=10.1016/0169-328x(95)00282-w;
RA Haire M.F., Chiaramello A.;
RT "Transient expression of the basic helix-loop-helix protein NSCL-2 in the
RT mouse cerebellum during postnatal development.";
RL Brain Res. Mol. Brain Res. 36:174-178(1996).
RN [7]
RP FUNCTION, TISSUE SPECIFICITY, DEVELOPMENTAL STAGE, AND DISRUPTION
RP PHENOTYPE.
RX PubMed=9090387; DOI=10.1038/ng0497-397;
RA Good D.J., Porter F.D., Mahon K.A., Parlow A.F., Westphal H., Kirsch I.R.;
RT "Hypogonadism and obesity in mice with a targeted deletion of the Nhlh2
RT gene.";
RL Nat. Genet. 15:397-401(1997).
RN [8]
RP FUNCTION, AND DISRUPTION PHENOTYPE.
RX PubMed=12419415; DOI=10.1016/s0031-9384(02)00885-5;
RA Coyle C.A., Jing E., Hosmer T., Powers J.B., Wade G., Good D.J.;
RT "Reduced voluntary activity precedes adult-onset obesity in Nhlh2 knockout
RT mice.";
RL Physiol. Behav. 77:387-402(2002).
RN [9]
RP FUNCTION, DEVELOPMENTAL STAGE, AND DISRUPTION PHENOTYPE.
RX PubMed=15470499; DOI=10.1038/sj.emboj.7600431;
RA Krueger M., Ruschke K., Braun T.;
RT "NSCL-1 and NSCL-2 synergistically determine the fate of GnRH-1 neurons and
RT control necdin gene expression.";
RL EMBO J. 23:4353-4364(2004).
RN [10]
RP FUNCTION, AND DISRUPTION PHENOTYPE.
RX PubMed=15465527; DOI=10.1016/j.yhbeh.2004.03.006;
RA Johnson S.A., Marin-Bivens C.L., Miele M., Coyle C.A., Fissore R.,
RA Good D.J.;
RT "The Nhlh2 transcription factor is required for female sexual behavior and
RT reproductive longevity.";
RL Horm. Behav. 46:420-427(2004).
RN [11]
RP FUNCTION, DEVELOPMENTAL STAGE, AND DISRUPTION PHENOTYPE.
RX PubMed=17573818; DOI=10.1111/j.1471-4159.2007.04694.x;
RA Schmid T., Krueger M., Braun T.;
RT "NSCL-1 and -2 control the formation of precerebellar nuclei by
RT orchestrating the migration of neuronal precursor cells.";
RL J. Neurochem. 102:2061-2072(2007).
RN [12]
RP FUNCTION, AND DISRUPTION PHENOTYPE.
RX PubMed=17717072; DOI=10.1210/me.2005-0337;
RA Cogliati T., Delgado-Romero P., Norwitz E.R., Guduric-Fuchs J.,
RA Kaiser U.B., Wray S., Kirsch I.R.;
RT "Pubertal impairment in Nhlh2 null mice is associated with hypothalamic and
RT pituitary deficiencies.";
RL Mol. Endocrinol. 21:3013-3027(2007).
RN [13]
RP FUNCTION, INTERACTION WITH STAT3, AND DISRUPTION PHENOTYPE.
RX PubMed=18356286; DOI=10.1210/me.2008-0010;
RA Fox D.L., Good D.J.;
RT "Nescient helix-loop-helix 2 interacts with signal transducer and activator
RT of transcription 3 to regulate transcription of prohormone convertase
RT 1/3.";
RL Mol. Endocrinol. 22:1438-1448(2008).
RN [14]
RP FUNCTION, AND DISRUPTION PHENOTYPE.
RX PubMed=19436734; DOI=10.1371/journal.pone.0005516;
RA Ruschke K., Ebelt H., Kloeting N., Boettger T., Raum K., Blueher M.,
RA Braun T.;
RT "Defective peripheral nerve development is linked to abnormal architecture
RT and metabolic activity of adipose tissue in Nscl-2 mutant mice.";
RL PLoS ONE 4:e5516-e5516(2009).
RN [15]
RP FUNCTION, AND DISRUPTION PHENOTYPE.
RX PubMed=20808804; DOI=10.1371/journal.pone.0012324;
RA Wankhade U.D., Vella K.R., Fox D.L., Good D.J.;
RT "Deletion of Nhlh2 results in a defective torpor response and reduced Beta
RT adrenergic receptor expression in adipose tissue.";
RL PLoS ONE 5:e12324-e12324(2010).
RN [16]
RP FUNCTION, AND DISRUPTION PHENOTYPE.
RX PubMed=23785158; DOI=10.1523/jneurosci.5287-12.2013;
RA Schmid T., Guenther S., Mendler L., Braun T.;
RT "Loss of NSCL-2 in gonadotropin releasing hormone neurons leads to
RT reduction of pro-opiomelanocortin neurons in specific hypothalamic nuclei
RT and causes visceral obesity.";
RL J. Neurosci. 33:10459-10470(2013).
CC -!- FUNCTION: Transcription factor which binds the E box motif 5'-
CC CA[TC][AG]TG-3' (PubMed:15470499, PubMed:18356286). Involved in
CC regulating energy expenditure, body mass, voluntary physical activity,
CC mating behavior and reproductive longevity, acting through the
CC hypothalamic-pituitary-gonadal axis (PubMed:15470499, PubMed:12419415,
CC PubMed:9090387, PubMed:17717072, PubMed:23785158, PubMed:15465527).
CC Acts as a transcriptional activator of target genes, including Ndn and
CC Pcsk1 (PubMed:15470499, PubMed:18356286). May act centrally to regulate
CC function of both white and brown adipose tissue (PubMed:20808804,
CC PubMed:19436734). Together with NHLH1, required to maintain migration
CC and survival of cells in the anterior extramural migration stream
CC (aes), which forms the precerebellar nuclei (PubMed:17573818). Also, in
CC concert with Nhlh1, may determine fate of gonadotropin releasing
CC hormone-1 (GnRH-1) neurons (PubMed:15470499).
CC {ECO:0000269|PubMed:12419415, ECO:0000269|PubMed:15465527,
CC ECO:0000269|PubMed:15470499, ECO:0000269|PubMed:17573818,
CC ECO:0000269|PubMed:17717072, ECO:0000269|PubMed:18356286,
CC ECO:0000269|PubMed:19436734, ECO:0000269|PubMed:20808804,
CC ECO:0000269|PubMed:23785158, ECO:0000269|PubMed:9090387}.
CC -!- SUBUNIT: Homodimer. Interacts and may form heterodimers with STAT3.
CC {ECO:0000269|PubMed:18356286}.
CC -!- INTERACTION:
CC Q64221; Q923E4: Sirt1; NbExp=2; IntAct=EBI-5378529, EBI-1802585;
CC -!- SUBCELLULAR LOCATION: Nucleus {ECO:0000255|PROSITE-ProRule:PRU00981}.
CC -!- TISSUE SPECIFICITY: Expressed in developing neurons (PubMed:1633105).
CC Transiently expressed in the cerebellum during postnatal development,
CC exclusively in the premigratory zone of the external granule layer
CC where postmitotic neurons undergo initial stages of neuronal
CC differentiation (PubMed:9011755). Expression is not detected in mature
CC neurons (PubMed:9011755). Expressed in the anterior lobe of the adult
CC pituitary (PubMed:9090387). {ECO:0000269|PubMed:1633105,
CC ECO:0000269|PubMed:9011755, ECO:0000269|PubMed:9090387}.
CC -!- DEVELOPMENTAL STAGE: Expressed in embryos at 10.5-13 days post coitus
CC (dpc), including developing ventral hypothalamus, Rathke's pouch and
CC subventricular layers of the developing diencephalon (PubMed:1633105,
CC PubMed:9090387). Expressed in the vomeronasal organ, olfactory
CC epithelium, and vomeronasal nerve fibers at 14.5 to 16.5 dpc
CC (PubMed:15470499). Expressed in various areas of the developing
CC hypothalamus at 18.5 dpc, including the septal areas, the diagonal band
CC of Broca (DBB), within the organum vasculosum lateralis terminalis
CC (OVLT) region, in the paraventricular nucleus (PVN), in lateral areas
CC of the hypothalamus (LH), in the arcuate nucleus (ARC), and in the
CC dorsomedial hypothalamic nucleus (DMH) (PubMed:15470499). Also
CC expressed in the developing pons from 14.5 dpc onwards, including the
CC pontine and reticulotegmental nuclei (PubMed:17573818).
CC {ECO:0000269|PubMed:15470499, ECO:0000269|PubMed:1633105,
CC ECO:0000269|PubMed:17573818, ECO:0000269|PubMed:9090387}.
CC -!- DISRUPTION PHENOTYPE: Disruption of the hypothalamic-pituitary axis in
CC both genders, but female pubertal development is influenced by the
CC presence or absence of male mice (PubMed:9090387, PubMed:17717072).
CC Altered female sexual behavior and reproductive longevity, with fewer
CC pregnancies over a shorter period, abnormal estrous cycles and reduced
CC ovulation with aging (PubMed:15465527). Reduction in neuroendocrine
CC gonadotropin releasing hormone-1 (GnRH-1) cell number and altered
CC location of these cells, and morphological and functional abnormalities
CC of the adenohypophysis (PubMed:17717072, PubMed:15470499). Ndn
CC expression almost abolished at the level of the optic chiasma in the
CC hypothalamus (PubMed:15470499). Hypothalamic Pcsk1 expression does not
CC respond to signals of energy availability (PubMed:18356286).
CC Progressive adult-onset obesity, preceded by reduced physical activity
CC (PubMed:9090387, PubMed:12419415). Significant reduction in the
CC innervation and vascularization of white adipose tissue and
CC accumulation of preadipocyte/macrophage-like cells, prior to onset of
CC obesity (PubMed:19436734). Defective torpor response and altered serum
CC leptin levels, body temperature and adipose inflammation
CC (PubMed:20808804). Conditional knockout in GnRH neurons but not in pro-
CC opiomelanocortin (POMC) neurons reduces POMC neuron number and
CC increases visceral fat mass (PubMed:23785158). Double knockout of Nhlh1
CC and Nhlh2 genes causes neonatal lethality, complete absence of GnRH-1
CC neurons in the posterior parts of the brain at 18.5 days post coitus
CC (dpc) and aberrant morphology of the remaining GnRH-1 neurons in the
CC anterior parts of the brain (PubMed:15470499). Double knockout of Nhlh1
CC and Nhlh2 genes causes absence of pontine nuclei, which belong to the
CC precerebellar nuclei and are located either side of the midline of the
CC ventral rhombencephalon (PubMed:17573818).
CC {ECO:0000269|PubMed:12419415, ECO:0000269|PubMed:15465527,
CC ECO:0000269|PubMed:15470499, ECO:0000269|PubMed:17573818,
CC ECO:0000269|PubMed:17717072, ECO:0000269|PubMed:18356286,
CC ECO:0000269|PubMed:19436734, ECO:0000269|PubMed:20808804,
CC ECO:0000269|PubMed:23785158, ECO:0000269|PubMed:9090387}.
CC -!- CAUTION: Has been reported to be expressed in Rathke's pouch and the
CC developing pituitary gland (PubMed:9090387). However, a later report
CC found no expression in these tissues and speculated that earlier
CC results may represent cross-hybridization with the very similar Nhlh1
CC gene (PubMed:15470499). {ECO:0000269|PubMed:15470499,
CC ECO:0000269|PubMed:9090387}.
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DR EMBL; S40532; AAB22580.1; -; mRNA.
DR EMBL; AK031747; BAC27535.1; -; mRNA.
DR EMBL; AK053837; BAC35549.1; -; mRNA.
DR EMBL; AK079121; BAC37549.1; -; mRNA.
DR EMBL; AK082728; BAC38590.1; -; mRNA.
DR EMBL; AK084160; BAC39129.1; -; mRNA.
DR EMBL; AK140559; BAE24423.1; -; mRNA.
DR EMBL; AC166072; -; NOT_ANNOTATED_CDS; Genomic_DNA.
DR EMBL; CH466608; EDL07620.1; -; Genomic_DNA.
DR EMBL; CH466608; EDL07621.1; -; Genomic_DNA.
DR EMBL; CH466608; EDL07622.1; -; Genomic_DNA.
DR EMBL; BC058413; AAH58413.1; -; mRNA.
DR CCDS; CCDS17685.1; -.
DR PIR; A49005; A49005.
DR RefSeq; NP_848892.1; NM_178777.3.
DR RefSeq; XP_006501175.1; XM_006501112.2.
DR AlphaFoldDB; Q64221; -.
DR SMR; Q64221; -.
DR IntAct; Q64221; 1.
DR STRING; 10090.ENSMUSP00000064355; -.
DR PhosphoSitePlus; Q64221; -.
DR PaxDb; Q64221; -.
DR PRIDE; Q64221; -.
DR ProteomicsDB; 269590; -.
DR Antibodypedia; 33870; 195 antibodies from 24 providers.
DR DNASU; 18072; -.
DR Ensembl; ENSMUST00000066187; ENSMUSP00000064355; ENSMUSG00000048540.
DR Ensembl; ENSMUST00000196324; ENSMUSP00000142746; ENSMUSG00000048540.
DR Ensembl; ENSMUST00000198675; ENSMUSP00000143362; ENSMUSG00000048540.
DR GeneID; 18072; -.
DR KEGG; mmu:18072; -.
DR UCSC; uc008qro.2; mouse.
DR CTD; 4808; -.
DR MGI; MGI:97324; Nhlh2.
DR VEuPathDB; HostDB:ENSMUSG00000048540; -.
DR eggNOG; KOG4029; Eukaryota.
DR GeneTree; ENSGT00940000162602; -.
DR InParanoid; Q64221; -.
DR OMA; RSCCASD; -.
DR OrthoDB; 1491224at2759; -.
DR PhylomeDB; Q64221; -.
DR BioGRID-ORCS; 18072; 2 hits in 72 CRISPR screens.
DR ChiTaRS; Nhlh2; mouse.
DR PRO; PR:Q64221; -.
DR Proteomes; UP000000589; Chromosome 3.
DR RNAct; Q64221; protein.
DR Bgee; ENSMUSG00000048540; Expressed in habenula and 91 other tissues.
DR Genevisible; Q64221; MM.
DR GO; GO:0000785; C:chromatin; IDA:UniProtKB.
DR GO; GO:0005634; C:nucleus; IEA:UniProtKB-SubCell.
DR GO; GO:0001228; F:DNA-binding transcription activator activity, RNA polymerase II-specific; IDA:NTNU_SB.
DR GO; GO:0000981; F:DNA-binding transcription factor activity, RNA polymerase II-specific; IBA:GO_Central.
DR GO; GO:0046983; F:protein dimerization activity; IEA:InterPro.
DR GO; GO:0000978; F:RNA polymerase II cis-regulatory region sequence-specific DNA binding; IDA:NTNU_SB.
DR GO; GO:0000977; F:RNA polymerase II transcription regulatory region sequence-specific DNA binding; IDA:UniProtKB.
DR GO; GO:0061629; F:RNA polymerase II-specific DNA-binding transcription factor binding; IPI:UniProtKB.
DR GO; GO:1990837; F:sequence-specific double-stranded DNA binding; ISO:MGI.
DR GO; GO:0006915; P:apoptotic process; IGI:UniProtKB.
DR GO; GO:0021535; P:cell migration in hindbrain; IGI:UniProtKB.
DR GO; GO:0021888; P:hypothalamus gonadotrophin-releasing hormone neuron development; IMP:UniProtKB.
DR GO; GO:0008584; P:male gonad development; IMP:UniProtKB.
DR GO; GO:0060179; P:male mating behavior; IMP:UniProtKB.
DR GO; GO:0042698; P:ovulation cycle; IMP:MGI.
DR GO; GO:0007422; P:peripheral nervous system development; IMP:UniProtKB.
DR GO; GO:0045944; P:positive regulation of transcription by RNA polymerase II; IDA:UniProtKB.
DR GO; GO:0006357; P:regulation of transcription by RNA polymerase II; IBA:GO_Central.
DR Gene3D; 4.10.280.10; -; 1.
DR InterPro; IPR011598; bHLH_dom.
DR InterPro; IPR036638; HLH_DNA-bd_sf.
DR InterPro; IPR040238; TAL-like.
DR PANTHER; PTHR13864; PTHR13864; 1.
DR Pfam; PF00010; HLH; 1.
DR SMART; SM00353; HLH; 1.
DR SUPFAM; SSF47459; SSF47459; 1.
DR PROSITE; PS50888; BHLH; 1.
PE 1: Evidence at protein level;
KW Activator; Developmental protein; Differentiation; DNA-binding; Nucleus;
KW Reference proteome; Transcription; Transcription regulation.
FT CHAIN 1..135
FT /note="Helix-loop-helix protein 2"
FT /id="PRO_0000127200"
FT DOMAIN 77..129
FT /note="bHLH"
FT /evidence="ECO:0000255|PROSITE-ProRule:PRU00981"
FT REGION 1..81
FT /note="Disordered"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT COMPBIAS 9..23
FT /note="Polar residues"
FT /evidence="ECO:0000256|SAM:MobiDB-lite"
FT CONFLICT 55
FT /note="A -> V (in Ref. 1; AAB22580)"
FT /evidence="ECO:0000305"
SQ SEQUENCE 135 AA; 15004 MW; E7872BF22A601405 CRC64;
MMLSPDQAAD SDHPSSTHSD PESLGGADTK VLGSVSDLEP VEEADGDGKG GSRAALYPHP
QQLSREEKRR RRRATAKYRS AHATRERIRV EAFNLAFAEL RKLLPTLPPD KKLSKIEILR
LAICYISYLN HVLDV