MUP20_MOUSE
ID MUP20_MOUSE Reviewed; 181 AA.
AC Q5FW60;
DT 05-OCT-2010, integrated into UniProtKB/Swiss-Prot.
DT 01-MAR-2005, sequence version 1.
DT 03-AUG-2022, entry version 127.
DE RecName: Full=Major urinary protein 20 {ECO:0000312|MGI:MGI:3651981};
DE AltName: Full=Darcin {ECO:0000303|PubMed:20525243};
DE AltName: Full=Major urinary protein 24 {ECO:0000312|EMBL:ACF70718.1};
DE Flags: Precursor;
GN Name=Mup20 {ECO:0000312|MGI:MGI:3651981};
GN Synonyms=Mup24 {ECO:0000312|EMBL:DAA06315.1};
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] {ECO:0000312|EMBL:ACF70718.1}
RP NUCLEOTIDE SEQUENCE [MRNA].
RC STRAIN=C57BL/6J {ECO:0000312|EMBL:ACF70718.1};
RC TISSUE=Liver {ECO:0000312|EMBL:ACF70718.1}, and
RC Submandibular gland {ECO:0000312|EMBL:ACF70718.1};
RX PubMed=18815613; DOI=10.1371/journal.pone.0003280;
RA Logan D.W., Marton T.F., Stowers L.;
RT "Species specificity in major urinary proteins by parallel evolution.";
RL PLoS ONE 3:E3280-E3280(2008).
RN [2] {ECO:0000312|EMBL:CAP58483.1}
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 [3] {ECO:0000312|EMBL:AAH89613.1}
RP NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RC STRAIN=FVB/N {ECO:0000312|EMBL:AAH92096.1};
RC TISSUE=Liver {ECO:0000312|EMBL:AAH89613.1};
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 [4] {ECO:0000305}
RP PROTEIN SEQUENCE OF 59-74 AND 129-141, FUNCTION, SUBCELLULAR LOCATION,
RP TISSUE SPECIFICITY, AND MASS SPECTROMETRY.
RC STRAIN=C57BL/6J {ECO:0000269|PubMed:15934926};
RC TISSUE=Urine {ECO:0000269|PubMed:15934926};
RX PubMed=15934926; DOI=10.1042/bj20050404;
RA Armstrong S.D., Robertson D.H., Cheetham S.A., Hurst J.L., Beynon R.J.;
RT "Structural and functional differences in isoforms of mouse major urinary
RT proteins: a male-specific protein that preferentially binds a male
RT pheromone.";
RL Biochem. J. 391:343-350(2005).
RN [5] {ECO:0000312|EMBL:DAA06315.1}
RP FUNCTION.
RC TISSUE=Urine {ECO:0000269|PubMed:18064011};
RX PubMed=18064011; DOI=10.1038/nature05997;
RA Chamero P., Marton T.F., Logan D.W., Flanagan K., Cruz J.R.,
RA Saghatelian A., Cravatt B.F., Stowers L.;
RT "Identification of protein pheromones that promote aggressive behaviour.";
RL Nature 450:899-902(2007).
RN [6] {ECO:0000305}
RP FUNCTION, SUBCELLULAR LOCATION, TISSUE SPECIFICITY, AND IDENTIFICATION BY
RP MASS SPECTROMETRY.
RC STRAIN=C57BL/6J {ECO:0000269|PubMed:20525243};
RC TISSUE=Urine {ECO:0000269|PubMed:20525243};
RX PubMed=20525243; DOI=10.1186/1741-7007-8-75;
RA Roberts S.A., Simpson D.M., Armstrong S.D., Davidson A.J., Robertson D.H.,
RA McLean L., Beynon R.J., Hurst J.L.;
RT "Darcin: a male pheromone that stimulates female memory and sexual
RT attraction to an individual male's odour.";
RL BMC Biol. 8:75-75(2010).
RN [7]
RP IDENTIFICATION BY MASS SPECTROMETRY [LARGE SCALE ANALYSIS].
RC TISSUE=Liver;
RX PubMed=21183079; DOI=10.1016/j.cell.2010.12.001;
RA Huttlin E.L., Jedrychowski M.P., Elias J.E., Goswami T., Rad R.,
RA Beausoleil S.A., Villen J., Haas W., Sowa M.E., Gygi S.P.;
RT "A tissue-specific atlas of mouse protein phosphorylation and expression.";
RL Cell 143:1174-1189(2010).
RN [8]
RP FUNCTION.
RX PubMed=23239735; DOI=10.1126/science.1225638;
RA Roberts S.A., Davidson A.J., McLean L., Beynon R.J., Hurst J.L.;
RT "Pheromonal induction of spatial learning in mice.";
RL Science 338:1462-1465(2012).
RN [9]
RP FUNCTION.
RX PubMed=25972792; DOI=10.3389/fnbeh.2015.00106;
RA Hoffman E., Pickavance L., Thippeswamy T., Beynon R.J., Hurst J.L.;
RT "The male sex pheromone darcin stimulates hippocampal neurogenesis and cell
RT proliferation in the subventricular zone in female mice.";
RL Front. Behav. Neurosci. 9:106-106(2015).
RN [10]
RP SUBCELLULAR LOCATION, TISSUE SPECIFICITY, AND POLYMORPHISM.
RX PubMed=28522864; DOI=10.1038/s41598-017-02259-1;
RA Liu Y.J., Guo H.F., Zhang J.X., Zhang Y.H.;
RT "Quantitative inheritance of volatile pheromones and darcin and their
RT interaction in olfactory preferences of female mice.";
RL Sci. Rep. 7:2094-2094(2017).
RN [11]
RP ERRATUM OF PUBMED:28522864.
RX PubMed=29511252; DOI=10.1038/s41598-018-22515-2;
RA Liu Y.J., Guo H.F., Zhang J.X., Zhang Y.H.;
RL Sci. Rep. 8:4300-4300(2018).
RN [12]
RP FUNCTION.
RX PubMed=31996852; DOI=10.1038/s41586-020-1967-8;
RA Demir E., Li K., Bobrowski-Khoury N., Sanders J.I., Beynon R.J.,
RA Hurst J.L., Kepecs A., Axel R.;
RT "The pheromone darcin drives a circuit for innate and reinforced
RT behaviours.";
RL Nature 578:137-141(2020).
RN [13]
RP STRUCTURE BY NMR OF 20-181, FUNCTION, SUBCELLULAR LOCATION, TISSUE
RP SPECIFICITY, POLYMORPHISM, AND DISULFIDE BOND.
RX PubMed=25279835; DOI=10.1371/journal.pone.0108415;
RA Phelan M.M., McLean L., Armstrong S.D., Hurst J.L., Beynon R.J., Lian L.Y.;
RT "The structure, stability and pheromone binding of the male mouse protein
RT sex pheromone darcin.";
RL PLoS ONE 9:E108415-E108415(2014).
CC -!- FUNCTION: Male pheromone which stimulates female sexual attraction to
CC male urinary scent and promotes a strong learned attraction to the
CC airborne urinary odor of an individual male (PubMed:20525243,
CC PubMed:31996852). Promotes spatial learning by rapidly conditioning
CC preference for its remembered location among females and competitor
CC males so that animals prefer to spend time in the site even when scent
CC is absent (PubMed:23239735). In addition to promoting a rapid
CC attraction response, also elicits ultrasonic vocalizations and urinary
CC scent marking in females which do not occur immediately after exposure
CC (PubMed:31996852). Stimulates hippocampal neurogenesis and cell
CC proliferation in the subventricular zone in females (PubMed:25972792).
CC Promotes male aggressive behavior (PubMed:18064011). Response to Mup20
CC is mediated by a neural circuit extending from the accessory olfactory
CC bulb to a subset of nitric oxidase synthase-expressing neurons in the
CC medial amygdala (PubMed:31996852). As well as acting as a pheromone
CC itself, binds most of the male pheromone, 2-sec-butyl-4,5-
CC dihydrothiazole, in urine and is responsible for its slow release from
CC scent marks (PubMed:15934926, PubMed:25279835).
CC {ECO:0000269|PubMed:15934926, ECO:0000269|PubMed:18064011,
CC ECO:0000269|PubMed:20525243, ECO:0000269|PubMed:23239735,
CC ECO:0000269|PubMed:25279835, ECO:0000269|PubMed:25972792,
CC ECO:0000269|PubMed:31996852}.
CC -!- SUBCELLULAR LOCATION: Secreted {ECO:0000269|PubMed:15934926,
CC ECO:0000269|PubMed:20525243, ECO:0000269|PubMed:25279835,
CC ECO:0000269|PubMed:28522864}.
CC -!- TISSUE SPECIFICITY: Detected in urine of males but absent from female
CC urine (at protein level). {ECO:0000269|PubMed:15934926,
CC ECO:0000269|PubMed:20525243, ECO:0000269|PubMed:25279835,
CC ECO:0000269|PubMed:28522864}.
CC -!- MASS SPECTROMETRY: Mass=18894; Mass_error=2; Method=Electrospray;
CC Evidence={ECO:0000269|PubMed:15934926};
CC -!- POLYMORPHISM: Constitutes approximately 10% of the total major urinary
CC protein composition in the urine of C57BL/6 males but is barely
CC detectable in the urine of BALB/c males. {ECO:0000269|PubMed:25279835,
CC ECO:0000269|PubMed:28522864}.
CC -!- SIMILARITY: Belongs to the calycin superfamily. Lipocalin family.
CC {ECO:0000255}.
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DR EMBL; EU882234; ACF70718.1; -; mRNA.
DR EMBL; BX088584; -; NOT_ANNOTATED_CDS; Genomic_DNA.
DR EMBL; CT990635; CAP58483.1; -; Genomic_DNA.
DR EMBL; CT990636; CAQ11567.1; -; Genomic_DNA.
DR EMBL; BC089613; AAH89613.1; -; mRNA.
DR EMBL; BC092096; AAH92096.1; -; mRNA.
DR EMBL; BK006677; DAA06315.1; -; Genomic_DNA.
DR CCDS; CCDS18233.1; -.
DR RefSeq; NP_001012323.1; NM_001012323.1.
DR RefSeq; XP_006538108.1; XM_006538045.2.
DR PDB; 2L9C; NMR; -; A=20-181.
DR PDBsum; 2L9C; -.
DR AlphaFoldDB; Q5FW60; -.
DR BMRB; Q5FW60; -.
DR SMR; Q5FW60; -.
DR STRING; 10090.ENSMUSP00000073667; -.
DR Allergome; 478; Mus m 1.
DR iPTMnet; Q5FW60; -.
DR PhosphoSitePlus; Q5FW60; -.
DR CPTAC; non-CPTAC-3659; -.
DR jPOST; Q5FW60; -.
DR MaxQB; Q5FW60; -.
DR PaxDb; Q5FW60; -.
DR PeptideAtlas; Q5FW60; -.
DR PRIDE; Q5FW60; -.
DR ProteomicsDB; 290225; -.
DR DNASU; 381530; -.
DR Ensembl; ENSMUST00000074018; ENSMUSP00000073667; ENSMUSG00000078672.
DR GeneID; 381530; -.
DR KEGG; mmu:381530; -.
DR UCSC; uc008tbr.1; mouse.
DR CTD; 381530; -.
DR MGI; MGI:3651981; Mup20.
DR VEuPathDB; HostDB:ENSMUSG00000078672; -.
DR eggNOG; ENOG502S6GK; Eukaryota.
DR GeneTree; ENSGT01050000244868; -.
DR HOGENOM; CLU_094061_4_0_1; -.
DR InParanoid; Q5FW60; -.
DR OMA; GVYADEY; -.
DR OrthoDB; 1475169at2759; -.
DR PhylomeDB; Q5FW60; -.
DR TreeFam; TF338197; -.
DR BioGRID-ORCS; 381530; 3 hits in 39 CRISPR screens.
DR ChiTaRS; Mup20; mouse.
DR PRO; PR:Q5FW60; -.
DR Proteomes; UP000000589; Chromosome 4.
DR RNAct; Q5FW60; protein.
DR Bgee; ENSMUSG00000078672; Expressed in liver and 21 other tissues.
DR GO; GO:0005576; C:extracellular region; IDA:UniProtKB.
DR GO; GO:0005615; C:extracellular space; IBA:GO_Central.
DR GO; GO:0000772; F:mating pheromone activity; IDA:UniProtKB.
DR GO; GO:0005549; F:odorant binding; IBA:GO_Central.
DR GO; GO:0005550; F:pheromone binding; IDA:UniProtKB.
DR GO; GO:0036094; F:small molecule binding; IEA:InterPro.
DR GO; GO:0008355; P:olfactory learning; IDA:UniProtKB.
DR GO; GO:2000179; P:positive regulation of neural precursor cell proliferation; IDA:UniProtKB.
DR GO; GO:0050769; P:positive regulation of neurogenesis; IDA:UniProtKB.
DR GO; GO:0071625; P:vocalization behavior; IDA:UniProtKB.
DR Gene3D; 2.40.128.20; -; 1.
DR InterPro; IPR012674; Calycin.
DR InterPro; IPR002345; Lipocalin.
DR InterPro; IPR022272; Lipocalin_CS.
DR InterPro; IPR000566; Lipocln_cytosolic_FA-bd_dom.
DR InterPro; IPR002971; Maj_urinary.
DR PANTHER; PTHR11430; PTHR11430; 1.
DR Pfam; PF00061; Lipocalin; 1.
DR PRINTS; PR01221; MAJORURINARY.
DR SUPFAM; SSF50814; SSF50814; 1.
DR PROSITE; PS00213; LIPOCALIN; 1.
PE 1: Evidence at protein level;
KW 3D-structure; Direct protein sequencing; Disulfide bond; Pheromone;
KW Pheromone-binding; Reference proteome; Secreted; Signal.
FT SIGNAL 1..19
FT /evidence="ECO:0000250|UniProtKB:P11590"
FT CHAIN 20..181
FT /note="Major urinary protein 20"
FT /evidence="ECO:0000250|UniProtKB:P11590"
FT /id="PRO_0000398791"
FT DISULFID 83..176
FT /evidence="ECO:0000269|PubMed:25279835,
FT ECO:0007744|PDB:2L9C"
FT HELIX 31..34
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 35..37
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 40..46
FT /evidence="ECO:0007829|PDB:2L9C"
FT TURN 48..51
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 60..66
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 68..76
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 80..82
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 88..92
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 98..110
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 116..128
FT /evidence="ECO:0007829|PDB:2L9C"
FT STRAND 131..144
FT /evidence="ECO:0007829|PDB:2L9C"
FT HELIX 147..157
FT /evidence="ECO:0007829|PDB:2L9C"
FT HELIX 158..160
FT /evidence="ECO:0007829|PDB:2L9C"
FT TURN 164..166
FT /evidence="ECO:0007829|PDB:2L9C"
FT HELIX 170..172
FT /evidence="ECO:0007829|PDB:2L9C"
FT TURN 177..179
FT /evidence="ECO:0007829|PDB:2L9C"
SQ SEQUENCE 181 AA; 20930 MW; CBAF1D33E1B03074 CRC64;
MKLLVLLLCL GLTLVCVHAE EASSMERNFN VEKINGEWYT IMLATDKREK IEEHGSMRVF
VEYIHVLENS LALKFHIIIN EECSEIFLVA DKTEKAGEYS VTYDGSNTFT ILKTDYDNYI
MIHLINKKDG ETFQLMELYG REPDLSSDIK EKFAQLSEEH GIVRENIIDL TNANRCLEAR
E
